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PROCEEDINGS
OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS
ZOOLOGICAL SOOLETY
OF LONDON
1923, pp. 483-1097,
with 3 Prarns, 191 Texr-rigurns, AnD 1 Map.
rare TN oe
ee
goal yn Ins HG
tha ‘ns iti %
SU In nD 7 th hus ae Dy
PRINTED FOR THE SOCIETY,
SOLD AT ITS HOUSE IN REGENTS PARK,
LONDON:
MESSRS. LONGMANS, GREEN, AN) CO,
PATRERNOSTER ROW,
4
Pst
OF THE
COUNCIL.
AND. OR LOERS
- OF THE
ZOOLOGICAL
SOCIETY OF LONDON.
1923,
| Patron.
His Masrsry Tur Kine.
COUNCIL.
His Grace Tar Duke or Beprorp, K.G.,
Cou. 8S. Moncx'ron
M.D., F.R.S.
Cuarites Drummonn, Esq,
Treasurer.
AuFreD HKizra, Ksq., O.B.K.
Huen S. Guapstonr, Esq.,
M.A., F.R.S.E.
Tne Rr. Hon. Tur Viscount
Grey or FaLnopon, K.G.,
P.C., Vice-President.
Sir Sipney F. Harmer, K.B.E.,
M.A., Se.D., F.R.S., Vice-
President.
Pror. James P. Ilin1n, D.Sce.,
F.R.S., Vice-President.
WILLIAM ‘ Howesman, sq.
Lr.-Con. Tins Lorp ALASTAtr
Roser Innus-Kur, D.S.0.
Str Warren Roper LAWRENCE,
Br.,G.C.1.E., G.C.V.O., C.B.
CoPrEMAN,
FE.R.S., President.
Cot. Sm Henry McManon,
G.C.M.G., K.C.L.1.
KE. G. B. Mrape- WALDbo, sq.,
Vice-President.
P, CHALMERS Mivoueen Esq.,
C.B.M., M.A., D.Se., LL.D.,
E.R.S., Secretary.
Tar Hart or Onstow, O.B.K.,
Vice-President.
Magor Aturrnr Pam, O.B.1K.
Liz.-Con. Sim Davin Pratn,
C.M.G., C.LE., F.R.S.
Tue Lord QuEENBOROUGH.
Warotp Jonn ITlastinas
RussEnn, Esq., IK.C.
Ricnard Streruens ‘Taynor,
Lisa.
A. Surri Woopwarp, Ksq.,
LLD., Y.R.S., Vice-Presi-
dent.
PRINCIPAL OFFICERS.
P, Cuatmers Mircnmny, C.B.M., M.A., D.Se.
Secretary.
, LL.D. ERS.
G. M. Verviers, M.R.C.S., L.R.C.P., Superintendent of the
Gardens.
D, Sern-Surrn, Resident Curator of Mammals and Birds.
Epwanp G. Bourenar n, Director of the Aquarium.
Miss Joan B. Procrrn, F.L
S., Curator of Reptiles.
Miss L. Ii. Currsman, E.LS., Curator of Insects.
C. F. Sonnac, M.D.,
Viens
N.S. Lucas, M.B., Ch.B., Pathologist.
Prof. G. H. Woorpripar, F. R.C.V.S., Hon. Consulting Veterinary
Be geon.
I Wi AL.
oh advologist.
SatMANnD, O.B.E., M.D.,
Ton, Consulting
If, Martin Duncan, F.R.M.S., Librarian.
I, W. Bonn, Accountant.
W.Y, Wannis, Chief Clerk,
LIST OF CONTENTS.
1923, pp. 483-1097.
EXHIBITIONS AND NOTICKS.
Tue Secretary. Exhibition of Photographs of Big Game
eon Clioyrrna, ING IRINOCIT RY — Goonasecuocnonsononoooousor
Miss L. Mi. Crmesman, F.1S., F.Z.S. Exhibition of
(1) living specimens of Peripatus; and (2) of section
of nest of Stingless Bee from Australia..................
Mr. F. Martin Duncan, F.R.M.S., F.Z.S. Account of his
recent work on microscopic structure of Mammalian
IDENT pee Sho meen oTe eee USM le Am icc at eRe
The Sscrerary. Report on Additions to the Society’s
Menagerie during the month of April, 1923............
Dr. G. M. Vevens, F.Z.S. Exhibition of the parasitic Pro-
tozoon Bealantidiamn colt ....c. ccc ccc cee ese ccc e cence en sent ces
Prof, J. P. in, F.R.S., Vice-President. Exhibition of
Photographs of the Australian . Lung-Fish (Cera-
UTILS) ec cas Sondese bapa oso denon se des sede nocdcbdc baa scuNsacbonesat
Dr. N. 8. Lucas, F.Z.S. Exhibition of a mucous cyst......
Mr, C. Tare Reean, M.A.;,-F.R.S. Exhibition of lantern-
slides from Photographs of the Spotted Basking
Shark (himodon typicws) .......csscesecsssasernecnecenoes
he Secrerary. Report on Additions to the Society’s
Menagerie during the month of May, 1923 ............
Mr. D. Seru-Smuirn, F.Z.8., and Dr. P. HW. Manson Bann, |
DS OF A: 8. Exhibition of photographs and
sketches showing the display of Hunstein’s Magnifi-
cent Bird-of- Paradise (Dipha ee magnifica hun
GUIAUOO))) | Ben ogdeed sense qoecco nes ucAacde var onqdanendspaonaacos00 cn
Vago
lv
Miss Atysr Cunninenam. Exhibition of cinematograph
records of the Gorilla “ John” ........cc cece cee e cence ees
Mr. BE. G. Bounuyarr, F.Z.S. Exhibition of models for
the Rockwork of the Society’s new Aquarium .........
Mrs. Neate. Exhibition of a living Tree-Hyrax (Dendro-
(EGIRTAD GHORSANIS)) conncoshsoossueocounssades8He ceo canobedueccocon
Dr. H. A. Bayuis, F.Z.S. Exhibition of a Nematode
On, MMEASAUPOS UGOICOD sonsoscossscocnc00s0aponae002000
My. BE, A. Spavt, F.Z.S. Exhibition of specimens showing
metamorphosis of Axolotls induced by injection of
pituitary gland anterior lobe extract.................66:
The Sxcretary. Report on Additions to the Society’s
Menagerie during the months of June, July, and
ANUS, US) | cnosdosonsosbosubsd25 .obdedcedaosyoocneoncopone
Dr. C. W. Anprews, F.R.S., F.Z.8. Exhibition of phalan-
geal bone of a clawed Perissodactyl........................
Mr. D. Seru-Suivu, 7.2.8. (1) Exhibition of photographs
of Lion-cubs ; and (2) of rave varieties of Black
IN larayaloeyy WMLOIMENY covcsoonos soso ccssccacdnoosoanerpoaDa00004
Dr. Cart Ansonon. Exhibition of drawings and photo-
graphs of Cave Animals from the Balkans ......... ..
Mr. E. A. Sraun, B.Se., 1.2.8. Remarks on acceleration
of metamorphoses of Wrog-tadpoles .....................
eeeeee
The Secrerary. Uxhibition of photograph of a Ratel
Mr. D. Seru-Smurru, F.Z.8. Kxhibition of Young Pigmy
Hippopotamus lexb-momrel |) ererer eee eee eceer
Mr. F. A. Mrrcneni-Herpexs, F.L.8., "2.5, Exhibition
of photographs on his recent expedition to the jungle-
recionol Vanamac.aeneey shel. atte avacnsebutiaci. 5 fae eerase ee
The Srcrrrary. Report on Additions to the Society’s
Menagerie during the month of October, 1923.........
The Secretary. Exhibition of an autograph letter of Sir
Stamford Raflles, and of a collection of autographs
recently presented to the Society .........,....:00000 ane
Page
657
1093
1095
1095
1095
1095
1096
1096
1096
1097
1097
Vv
Page
Mr. R. T. Gunter, M.A., 12.8. Exhibition of (1) Ver-
tebre of Mesozoic Crocodiles ; and (2) of a jaw-bone
(OPC GSTS CDG TINGED Waniek aap bee abada: Sd bok ose abnnae neue ssalct: 1097
MiphabeticalMinistobs Comb wtorsrace sre er-ccsmeersics: steerer: Vill
Thvdlexe ote illtsticbronse meaner eci meme cacee yas slices bites XV
UDC AN ee eee NepOUeL al Date PRIA M RS ac, OUNRL Lac Ista cpanel tater cB x1x
PAPERS.
26. On Mammals collected by Capt. Shortridge during the
Percy Sladen and Kaffrarian Expedition to the Orange
River. By Oxnprity Tuomas, F.R.S., F.Z.8., and
Wbaramy IN (GS IRI INMOST, INZAIS), \ soosasnondonscoboreonoede co 483
27. The Chondrocranium of the 'leleostean Fish Sebastes
marinus. By N. A. Macxinrosu, B.Sc. (Text-
figures 1-9.) 2... cccececeeneeee tent et ec tee ces et en eneeeceees d01
98. The Comparative Anatomy of the Tongues of the
Mammialia.—LX. Kdentata, Dermoptera, and Insecti-
vora. By Cuas. F. Sonwrac, M.D., F.Z.5. (Text-
fieures S0=O7.)) 20.00. cccseee ees ver cress css seerensnscessessn: 515
29. The External Characters of the Pigmy Hippopotamus
(Cheeropsis liberiensis) and of the Suid and Camelide.
By R. I. Pocock, F.R.S., F.2.8. (‘Text-figures 30-46.) 531
30. A Revision of the Family Panthophthalmide [Diptera],
with Descriptions of New Species and a New Genus.
By Major E. KE. Ausrey, D.S.0., F.Z.5. (Text-
(Huse TEIN) godgendeqorancandoacspasnccanasneasosasocndc Vuaek DOL
31. New Cryptosome Beetles. By 8. Mavuuix, F.Z.8.
(Text-figures 1-8.) .........ccccereneee eee ees ee ee ee eeeaveees 599
32. On the Display of the Magnificent Bird-of-Paradise
Diphyllodes magnifica hunsteint. By D. Suru-Smrru,
TAs, (CU vextratileriers dhs) Gaps poosbosseoentsadapcagaeodosoe 609
33. The Brain of the Zeuglodontide (Cetacea), By
Raymonp A. Darr, M.Sc., M.B., Ch. M.; with a Note
on the Skulls from which the Endocranial Casts were
taken. By C. W. Anprews, D.Sc., F.R.S., F.Z.S.
(Lext-figuves 1-24.)..........cceeecees tenes eens a eeneenenenes 615
34.
36.
37,
38.
39.
AQ.
4).
42.
43.
44,
45.
v1
On the Structure of the Skull in the Carnivorous Dino-
cephalian Reptiles. By R. Broom, M.D., .R.S.,
C.M.Z.8, (Wext-figures 1-17.) ................5. PANNE Nes
. Notes on Mast African Mamainals collected 1920-1923.
By Arritur Lovurtpasr, W.K.8., C.M.Z.8. (Map.) .
Observations on the Development of the Sympathetic
Nervous System and Suprarenal Bodies in the
Sparrow. By A. Supa Rav, B.A., M.Sce., F.RMLS.,
and P. H. Jomnson, B.A., B.Sc., F.L.8. (Text-
figures 1=20.) ... ccc eccnc eee eee cee te eter retorts t eee ee ees
A New Spider of the Genus Ziphistius from the Malay
Peninsula, and some Observations on its Habits. By
ll. C. Aprauam, F.Z.8., V.L.S. (Platel.; Text-
sPsAUNHs) Uh) pedcoossodosododbane soboacodes, andobnessoepeogeaden
A Review of the Lizards of the Genus 7opidophorus
on the Asiatic Mainland. By Matcoim A. Surru,
WEIR OLS 4 Ip OLE, I Adskonsasodosqadeosach congo oboceuess
On the Larval Anatomy of the Gout-Ily of Barley
(Chlorops teniopus Meig.) and two Related Acalyp-.
trate Muscids, with Notes on their Winter Host-
Plants. By J. H. Frew, M.Sce., F.Z.8. (Text-figures
A Revison of the Isopod Genus Ligidiwm Brandt—
Crustacea. By HaronpGorpon Jackson, F.Z.5.
(Rext=tigures TNO) iT icaceeceees arse aeede cerns sora sir
A List of the Lizards of British Territories in Kast
Africa (Uganda, Kenya Colony, Tanganyika Territory,
and Zanibar), with Keys for the Diagnosis of the
Species. By Arruur Loveriver, F.V.S.,C.M.Z.S. .
On some Mammals from Jugoslavia. By Ivor G. 5.
IOISHONCLO INGA baosonoasoougancocget camo sbeeos opoubne ound
Notes on East African Snakes collected 1918-1923.
By Arruur Loveripesr, F.E.S., C.M.Z.5.......0- 1
Notes on East African Birds (chiefly nesting habits and
endo-parasites) collected 1920-1923. By Arvitun
Theojiiorebrenoy INI DiSh, OLWIVAISs cadaancacebatenodo ooondooerds
Notes on East African Tortoises collected 1921-1923,
with the description of a new species of Soft Land
Tortoise. By Arruurn Loverspan, I1.8., C.M.Z.8.
(Blenirag 16 03 TE)) conoptonossdoossasptoonse noon BG Arta Aan
Page
661
685
741
769
783
823
590)
46.
47.
48.
vil
Notes on Kast Afvican Lizards collected 1920-1923,
with the Description of two new Races of Agama
lionatus Blgr. By <Anrtnur Loveriver, F.E.S.,
CAME ZS ee entree tenet easter care mer ME EN eacuNin gh
On some new or little-known Species of Acari. B
STANLEY Hirsr, W.L.S. (‘Text-figures 1-24.).........
On the Pelvic Muscles and Generative Organs in the
Male Chimpanzee. By Cuaruus F. Sonnac, M.D.,
¥.L.8. (Anatomist to the Society). (Text-figures
DO OA a) 2 awe. SRD See eect ARM Ca ae ewe tre
. Notes on Mast African Insects collected 1915-1922. By
ANAM IDONnoRUT EO, WIGS, (CW AISIy Goosneeassousnencs
. On the Guernsey Crocidura. By IvornG. 8. Monvracu,
I.Z.8., and Grace Pickronn...... 5 een epee WRI.
. Reversible and Irreversible Hvolution.; a Study based
on Reptiles. By Dr. Francis, Baron Norcsa. (‘Text-
AOUTES Bice Ye) iy hase SMe a cen Ryn an all on ONE
. On New and Rare Reptiles from South America. By
Joan B. Procren, F.Z.5., F.L.5. (Curator of Reptiles).
(llexcG-fionesielNc, 2/5) crn mane entree teriac: cus s2a099osweane
. On New and Rare Reptiles and Batrachians from
the Australian Region. By Joan B. Procrer, F.Z.5.,
F.L.8. (Curator of Reptiles). (Text-figures 3 & 4.)
. Ov Additions to the Snake Fauna of Egypt. By Major
oso Noonan KOLB ID. T/Aass dgsuéiodoasabnboenobubonsoobe
. Cervical Vertebrie of a Gigantic Blue Whale from
Panama. By Sir Sipney FE. Warmer, K.B. K., Se.D.,
nel eecbe lile/Aas\s (Ass. qian) Ea hcebavecacnesns oboe galt
. Notes on the African Crested Rat (Lophiomys unhaustt).
IB yiG ld’ CLOUD FUN GUG baer cae as as we miyle ears eee cer weeeteaee
Page
935
oa
1001
1015
1043
1045
1061
1069
1079
1085
AGM WwiE Tat NIB DANG AV tn Se
Or THK
CONTRIBUTORS,
With References to the several Articles contributed by each.
(1923, pp. 483-1097.)
AsrauaM, H. C., F.Z.8., F.L.8.
A New Spider of the Genus Liphistiws from the
Malay Peninsula, and some Observations on its Iabits.
(atric) Tee A Nesqnas is (bU Op Ihe)! codnntadecodcasand@hasdonoaneddstnednan
Axnsoon, Dr, Carn.
Kxhibition of drawings and photographs of Cave
Animals from the Balkans
CO CCC Oe Cn CC ey
ANDREWS, C. W., D.Sc., F.R.S.
Exhibition of phalangeal bone of a clawed Perisso-
dactyl
Austen, Major KE. K., D.S.O., F.Z.S.
A Revision of the Family Panthopthalnide [ Diptera],
with Descriptions of New Species and a New Genus.
(‘Text-figures 1-11.)
Bann, P. 1. Manson. See Suivi, D. Servite,
Bayuis, H. A., M.A., D.Sc.
Mxhibition of a Nematode worm, Cowascaris lconina
Page
769
1095
1095
ix
Boutrenerr, K. G., F.Z.S.
Exhibition of models for the Rockwork of the Society's
TES EAC (ETT MUNIN etn Ree ene Rien ee Na AE er asc) oR aa
Broom, R., M.D., ¥.1.8., C.M.Z.S.
On the Structure of the Skull in the Carnivorous
Dinocephalian Reptiles. (Text-figures 1-17.) ............
Curresman, L. H., F.1AS., 1.7.5.
Exhibition of (1) living specimens of Peripatus; and
(2) of section of nest of Stingless Bee from Australia
CuNNINGHAM, ALYSE.
Exhibition of cinematograph records of the Gorilla
Ce OUI tartrate Wea ata Sevenee Ce enc ee Sea yatta Me ey ae oP
Darr, Rayvmonp A., M.Se., M.B., Ch.M.
The Brain of the Zeuglodontide (Cetacea). With a
Note on the Skulls from which the Endocranial Casts
were taken. By 0. W. Anprews, D.Sc., F.R.S. (Text-
floures sLSZA.\i usa sAiee. anvhtioemaaten n ER acter oot Neer ay ;
Duncan, F. Marvin, F.R.M.S., I.Z.8.
Account of his recent work on microscopic structure
ObMiaimanaeuli minllnauswer ne atece son seine ra amienene soe mmeceaeOne
Friower, Major 8. 8., O.B.E., F.Z.8S.
On Additions to the Snake Fauna of Weypt ............
Frew, J. H., M.Sc., F.Z.8.
On the Larval Anatomy of the Gout-Fly of Barley
(Chlorops tuniopus Meig.) and two Nelated Acalyp-
trate Muscids, with Notes on their Winter Host-Plants.
(Gestofioumes 23. iwc nensteas sn. ners sence ean) eee neeer cet
GovprincH, G. IL.
Notes on the African Crested Rat (Lophiomys
inahuanesti.) TGS SOC AER HES AB CHICO RAT RSE eee ISTE gn RPO Ue
GunrueEr, R. T,, M.A,, F.Z.S.
Exhibition of (1) Vertebree of Mesozoic Crocodile ; and
(2) of a jaw-bone of Ursus anglicus ........0.0. ces e cess aye
Page
657
661
655
655
1079
783
i091
x
Hanmer, Sir Srpney F., K.B.E., 8c.D., F.R.S.
Cervical Vertebrie of a Gigantic Blue Whale from
‘Panamaten @lext-hietmenll)) sna sce ee sep er ee er prereer sha ee cae
Hupans, Fr. A. Mivcnenn-, F.L.8., F.Z.8.
Exhibition of photographs and remarks on his recent
expedition to the jungle-region of Panama..................
Hit, J.P., D.S., F.R.S.
Exhibition of Photographs of the Australian Lung-
Mish (Cenatod us) 0 ce croc tevets cn eee saiogs ois ects stasis atone
Hiyton, Marvin A.C. See ‘tHomMas, OLDFINLD.
Hirst, Sranuey, |".Z.S.
On some new or little-known Species of Acari. (‘Text-
fires) l= 24) eee tes ie Ha NUR Rea tal ars eitivo ne Out
Jackson, Haronip Gorpon, F.Z.S.
A. Revision of the Isopod Genus Ligidiwm Brandt—
Crustaceal y(Lext-fouress Oey nhac. ero lcn sa iesece
Jounsron, P. H. See Rau, A. Susp.
Loventpcr, Anruur, F.E.S., 0.M.Z.8.
Notes on Kast African Mammals collected 1920-
ESP Hiner 1) oe) eet ren SA MnP nen icee ae en U nec h
A List of the Lizards of British ‘erritories in Hast
Africa (Uganda, Kenya Colony, Tanganyika ‘Territory,
and Zanzibar), with Keys for the Diagnosis of the
SPEClESHiee ts is Delhi eeMeUL eh ea PE ety ae es et
Notes on Hast African Snakes collected 1918-1923 .
Notes on Hast African Birds (chiefly nesting habits
and endo-parasites) collected 1920-1923 ..................405
Notes on Kast African Tortoises collected 1921-1923,
with the description of a new species of Soft Land
AMorenorseh (Cites My 6S JOC) sosobadeoacddbooneo0 ice ah eeetta
Notes on East African Lizards collected 1920-1923,
with the description of two new Races of Ayana lionotus
Blgr.
SHOP He eee ewe mee eee ree eseeseeeesneseeseeeeereteresivenseresseas
Page
1085
1096
656
971
685
841
571
899
923
939
Notes on East African Insects collected 1915-1922 ... 1013
Bee Page
Lucas, N.8., M.B., B.Ch.
LOSI MMLOMTOM CB, TONGOUNS CYS soooodengeoobodsnononndasnpncsose 656
Mackintosn, N. A., B.Se.
The Chondrocranium of the Teleostean Fish Sebastes
frennoRowISs . ((ANepxes tae IIS) Goo boequncocadccsens vospunenboce d01
Mauutik, 8., B.A., .Z.8,
New Cryptosome Beetles. (‘Text-figures 1-8.) ......... 599
Mirenen., Dr. P. Cuaumens, C.B.K., M.A., D.Se., LL.D.,.
F.RAS. (Secretary to the Society).
Kixhibition of Photographs of Big Game from Choma,
DSICOT ee) UR LST IOUU CL Ta emai ata ors ena My val aa Ne 655
Report on Additions to the Society’s Menagerie during
NS) THORN One VANDEL, IG ENa) sre snocaondnedoduaosaduduadusesncanode Ol)
Report on Additions to the Society’s Menagerie during
Ne: MOM OT IM laii, SYS) oaooovsamaanbosnonoccsosv0debonocon se soo ODE
veport on Additions to the Society’s Menagerie during
the months of June, July, August, and September, 1923. 1093
Exhibition of photograph of a Ratel ...............0..00- 1096
Report on Additions to the Society’s Menagerie during
themmonthori@ctowers O23 menaseeeeneC ere asker renee Mee eG 1097
Exhibition of an autograph letter of Sir Stamford
Raftles, and of a collection of Autographs recently pre-
sented tosthe; Societyaee were nee ee ee eee see eee 1097
Monracu, Ivor G.58., F.Z.S.
On some Mammals from Jugoslavia © ...........ccseseeee 865
Monrracu, Lvor G.5., and Pickrory, Grace.
Onithe Guernsey Crocidiunasl acer eee eee se 1048
NeEAue, Mrs.
Exhibition of a living Tree-Hyrax (Dendrohyrax
LOU SCLUS) were i vvaes entice Ne tet ter rast easy Bt ak Sele saiseualsoeteenie 657
Norcsa, Dr. Francis, Baron.
lteversible and Irreversible Evolution; a Study based
on Neptilosss (lext-heuresy Orde) o\is ease ecenr nea 1045
xil
PickrorD, Grace. See Monvaau, Ivor G. 8.
Pocock, R. I., F.R.S., I°.Z.8.
The External Characters of the Pigmy Hippopotamus
(Cheropsis liberiensis) and of the Suidw and Camelide.
(Rext=fignnes/ S024 65a: aimee SU Satan ona seen dee
Procren, Joan B., F.Z.8., F.L.8. (Curator of Reptiles).
On New and Rare Reptiles from South America.
(Wextolpmreswl ds: 2.) hor. sata eee te ACen ee 5 Baar
On New and Rare Reptiles and Batrachians from the
Australian Region. (Text-figures 3 & 4.) .........cccceeee.
Rav, A. Suppa, B.A., M.Sc., F.R.M.S., and Jounsyron,
Heo Lele Ibe, IEMs IDWAIS)
Observations on the Development of the Sympathetic
Nervous System and Suprarenal Bodies in the Sparrow.
(Mext=npurestl = ZOM ese geen enna ace nae eran
Reean, C. Tarr, M.A., F.R.S., F.Z.8.
Exhibition of lantern-slides from Photographs of the
Spotted Basking Shark (2hinodon typicus) .........c6cee eee
Suita, D. Srru-, 1.2.8.
On the Display of the Magnificent Bird-of-Paradise,
Diphyllodes magnifica hunsteini. (ext-figuves 1-4.) ...
(1) Exhibition of photographs of Lion-cubs; and
(2) of rare varieties of Black Mangabey Monkey .........
Kixhibition of young Pigmy Hippopotamus. (‘Text-
PUBS HIE): Ss cteehe soc eee Re ee ees HRCA RNa IN eee
Smiru, ). Seru-,and Baur, P. 1H. Manson, M.D., F.R.C.P.,
D.5.0.
[Exhibition of photographs and sketches showing the
display of Hunstein’s Magnificent Bird-of-Paradise
(Diphyllodes maynificu hunsteiit)..........0.cecee eee ec ee eevee
Smirm, Mancoum A., M.R.CS., L.R.C.P., F.Z.8.
A Review of the Lizards of the Genus Zropidophorus
OH UNE ATAUO Miley halen) i posoocsonbsoaeg0a0 doeuopdogdnduecacoaas
Page
531
1061
1069
741
656
609
1095
1096
657
Xi
Sonnrae, Carus F., M.D., I.Z.8. (Anatomist to the
Society.)
The Comparative Anatomy of the Tongues of Mam-
malia.—IX. Edentata, Dermoptera, and Insectivora.
(Text-figures 50-57.)
On the Pelvic Muscles and Generative Organs in the
Cee e eter ress eeteoe
Male Chimpanzee. (‘lext-figures 58-64.)
Sraun, EH. A., F.Z.8.
Exhibition of specimens showing metamorphosis of
Axolotls induced by injection of pituitary gland anterior
HoWeexcGra ctr ya ie acetigoncesetaais «santa sce Sanbin sch Sivanuntayanale ne ara
Remarks on acceleration of metamorphoses of Frog-
tadpoles
Ce i ire ica]
Ti10oMASs, Or prie.D, I’. R.S8., F.Z.8.,and Hinton, Manrin A.C.,
I.Z.8.
On Maminals collected by Capt. Shortridge during the
Percy Sladen and Kaffvarian Hxpedition to the Orange
NIT (eu os) Sat ee i aeet pratt PRM oer Uti oA ure artanial aie Aer eaten Arn ra
Vivers, G. M., M.R.C.S., L.2.C.P., 1.2.5.
Exhibition of the parasitic Protozoon Balantidiwm colt .
Page
515
1001
656
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INDEX OF ILLUSTRATIONS.
Aluroguathus tigriceps, Fig. 16, p. 683.
Amphibolurus fionni, Fig. 4, p. 1075.
Amphishena mitchelli, Fig, 2, p. 1065,
Ancystropus ethiopicus, Figs. 7, 10,
pp. 980, 983.
(Meristaspis) calcaratus, Fig. 11,
p. 984.
9
2, 13,
— lateralis, Figs. 1
pp. 985, 986.
—— macroglossi, Figs. 8-10,
pp. 981-983.
zelebori, Fig. 6, p. 978.
Anthropopithecus troglodytes, Figs. 58-
64, pp. 1902-1008.
Anychus latus, Figs. 17, 18, pp. 992,
993. ; )
Alopomyia rothschildi, Vig. 11, p. 597.
Balenoptera musculus, Fig. 1, p. 1086.
Balioptera combinata, Vigs. 10, 19-28,
pp. 795, 814, 816-818.
Batrachiterpeton, Pig. 9, p. 1054.
Batrachosuchus, Wig. 9, p. 1054.
Burnetia mirabilis, Pigs. 9, 10, pp. 671,
672.
Callispa alinora, 599.
Camelus dromedarius, Vigs. 40, 41, 44-
46, pp. 543, 544, 547-549.
Camptosaurus, Fig. 9, p. 1054.
Cassida dimbakar, Vig. 8, p. GOS.
—— manipuiria, Fig. 7, p. 697.
Proc. Zoon. Soc.—1923.
Cassida rati, Fig. 6, p. G05.
Champsosaurus, Fig. 8, p. 1050.
Chelone, Fig. 8, p. 1050.
Chlovops teiopus, Figs. 1-18, pp. 785,
786, 789, 790, 792-795, 801, 808
809.
Cheropsis liberiensis, Figs. 80-33,
pp. 9382-534 ; Fig. 1, p. 1096.
Chrysochloris trevelyant, Vig. 57, p. 526.
Cynognathus, Big. 8, p. 1050.
>
Delphinosaurus, Fig. 8, p. 1050.
Demodex sciurinus, Fig. 19, p. 994.
Dermochelys, Figs. 8,9, pp. 1050, 1054.
Diadectes, Fig. 9, p. 1054.
Dicotyles pecari, F igs.
pp. 537, 538, 541.
Dicynodon andrewsi, Vig. 18, p. 680.
Dinarianus vanderbyli, Fig. 8, p. 670.
Dinophenens ingens, Figs, 4-7, pp. 666-
669.
Diphyllodes magnifica hunsteini,
Figs. 1-4, pp. 609-612.
Diplocaulus, Fig. 9, p. 1054.
Disparactus, Fig. 8, ». 1050.
Downesia sasthi, Fig. 2, p 601.
39, 36, 38,
Endothiodon angusticeps, Fig, 15,
p. 682. ,
“nobius strubent, Fig. 17, p. 684.
Erinaceus europeus, Fig. 55, p. 624.
Euparkeria, Fig, 9, p. 1054.
xvi
Galcopithecus volans, Fig, 54, p. 522. }
Heloderma, Fig. 9, p. 1054.
Ichthyosaurus, Vig. 8, p. 1050.
Kannemeyeria simocephalus, Vig. 14,
p. 681.
Kari brunnea; Figs.,
ce
i
wie PPs 603; ee
Labidosaurus, Fig. 8, p. 1050.
Lama glama, Figs. 39, 42, 46, pp. 542,
546, 549.
-— huanacus, Figs. 39, 42, 46, pp. 542,
546, 549.
—— vicugna, Figs. 39, 41, pp. 542, 544.
Ligidium Jragile, Fig. 5, p. 881.
-—— germanicum, Fig. 9, p. 836.
— gracile, Fig. 6, p. 838.
hypnorwm, Figs. 1-4, pp. 826, 827,
829, 830.
japonicum, Fig. 10, p. 837.
— latun, Figs. 7, 8, pp. 834, 835.
Liphistius malayanus, Pl. 1., Fig. 1,
pp- 769, 771.
Liponyssus gordonensis, Vigs. 2, 3,
pp. 973, 974.
Listraphoroides ethiopicus, Vig. 2+,
p. 1000.
Listrophorus bothe, Fig. 24, p. 10C0.
Lygosoma (Emoa) battersbyi, Fig. 38,
p. L070.
Manis, Vig. 52, p. 519.
Meromyza nigriventris, Figs, 10, 14-18,
pp. 795, 810, 812, 818.
Microgomphodon, Vig. 9, p. 1054.
Myrmecophaga jubata, Fig. 50, p. 516.
Neusticosaurus, Fig. 8, p. 1050.
Orycteropus capensis, Fig. 52, p. 519.
Oxybelis boulengeri, Fig. 1, p. 1063.
4
Pantophthalmus batesi, Fig. 2, p. 567.
conspicabilis, Wig. 5, p. 579.
INDEX OF ILLUSTRATIONS.
Pantophthalmus conspicuus, Figs. 6, 7,
pp. 582, 585.
hellerianus, Vig. 3, p. 570.
splendidus, Fig. 8, p. 090. .
—— talaninus, Big. 1, p. 597.
variegatus, Vig, 9, p. 593,
versicolor, Wig. +, p. 573.
Paratetranychus indicus, Fig. 16, p. 990.
Passer domesticus, Vigs. 1-20, pp. 743-
750, 762, 704-756, 739-762, THD,
| Peloncustes, Wig. 8, p. 1050.
philarchus, Vig. 16, p. 683.
Periglischrus interruptus, Wigs. 14, 15,
pp. 988, 989.
Phacocherus africanus, I
pp. 537, 41,
Placochelys, Fig. 9, p. 1054.
Placodus, Vig. 9, p. 1054,
Plagiosaurus, Fig. 9, p. 1004.
Tlaticarpus, Pigs. 8, 9, pp. 1050, L054.
Polyeotylus, Fig. 8, p. 1050.
Potamocherus porcus, Wigs. 35, 36,
pp. 537, 538.
| Procolophon, ig. 9, p. 1054.
Proneusticosaurus, ig. 8, p. 1050.
| Prosqualodon davidi, Wigs. 18-21,
| pp. 639-642.
_ Protacmon, Fig. 9, p. 1054.
Prozeuglodon atrox, Wigs, 8-11, 22,
| pp. 627-629, 619.
Pseudotarsonemoides spinitarsus,
Figs, 22, 28, pp. 997, 908.
_ Pleranodon, Fig. 8, 1050.
Ithaphiorhynchis planiventris, Fig. 10,
p. 595.
Rhinonyssus (Neonyssoides) nuci/rage,
Figs. 4, 5, pp. 976, 977.
Rhynchoeyou, Vig. 57, pp. 526.
| Sauranodon, Fig. 8, p. 1000.
| Seapanodon duplessisi, Wigs. 1-2,
pp- 663-665.
| Sebastes marinus, Figs. 1-9, pp. 502,
503, 505-508, 510, 511.
| Sus leucomystaz, Fig.234, p. 536.
| —— scrofa, Bigs. 34, 37, pp. 5386, 540.
INDEX OF [LLUSTRATIONS.
Talpa curepea, Vig. 56. p. 526.
Tamandua tetradactyla, Big. 50, p. 516.
Tanganyika Territory, Map of, 685.
Tarsonemus (Larsonemella) africanus,
igs. 20, 21, p. 996.
Tatusia, Wig, 51, p. 61%.
Testudo proctere, Pls, 1., 11., p. 928.
Titanosuchus cloctei, Wigs. 11, 12,
pp- $77, 678.
Toxochelys, Wigs. 8, 9, pp. 1050, 1054.
Trachytes austent, Pig. 1, p. 972.
Trematops, Wig. 8, p. 1050.
Tylosaurus, Fig. 8, p. 1050.
Varanus, Pig. 9, p. 1054.
Zeugiodon ellioismithit, Figs, 5—7
pp. 625, 626.
—— intermedius, Vies. 12-14
pp. 630, 631, 650.
XViL
?
99
9 ay
—— osiris, Figs. 15-17, 24, pp. 682,
633, 651.
622, 624.
sensitivus, Wigs. 1-4, pp. 619, 621,
He a ilk
Ninoy
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|
es
PROCEEDINGS:
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS |.
OF THE
ZOOLOGICAL SOCIETY
OE LONDON 6
1928.
PART III.
contaInine Paces 483 To 659, wirn 81 ‘Texr-ricuREs.
“ansotla lnséy;
oe
SOLD AT ITS HOUSE IN REGENT’S PARK,
LONDON :
MESSRS. LONGMANS, GREEN, AND CoO.,
PATERNOSTER ROW.
[Price Twelve Shillings. |
List OF CON THN Ws:
1923, Parr IIL (pp. 488-659).
EXHIBITIONS AND NOTICES.
‘Tun Secrerary. Exhibition of Photographs of Big Game from Choma, North Rhodesia.
Miss L. W. Cunmsmay, F.E.S., F.Z.8. Uxhibition of (1) living specimens of Peripatus
and (2) of section of nest of Stingless Bee from Australia .
Mr. F. Martin Duncan, F.R.M.S., F.Z.S. Account of his recent work on microscopic
structure of Mammalian hairs ..........
‘Tue Secrurary.. Report on Additions to the Society’s Menagerio during the month of
April O23 errata:
Dr. G. M. Vevrrs, F.Z.S. Txhibition of the parasitic Protozoon Balantidium coli
Prof. J, P. Hitt, F.R.S., Vice-President. Exhibition of Photographs of the Australian
IDLTEI (CARMA) Solsainocsobugomcassnoaqonodde
Dr. N.S. Lucas, F.Z.S. Exhibition of a mucous cyst ..
Mr. O. Tarn Reaan, M.A., F.R.S. Exhibition of lantern-slides from Photographs of the
Spotted Basking Shark (Rhinodon typicus)........... D ‘QonoooaHOLO OC OOOO COBO CE
Tne Secretary, Report on Additions to the Society’s Menagerie during the month of
MI IIS PBL S cine Nebisg 76 SMG Reco GMC OB GH bamamaten o.cema AS OHRGmA Hom Guede be
Mr, D. Seru-Sairn, F.Z.S., and Dr. P. H. Manson Bann, D.S8.0O., F.Z.8. Exhibition of
photographs and sketches showing the display of Hunstein’s Magnificent Bird-of-
Paradise (Diphyllodes magnifica hunsteini)..........065.
Miss Atysz Ounninanam, Exhibition of cinematograph records of the Gorilla “John” .
Mr. E. G, Boutunarr, F'.Z.S. qlixhibition of models for the Rockwork of the Socioty’s
NOW. AQUALIIMY fe ohele) ololaleslelolalelere. stove ehelessiulcisite olitolsiehetale -lteleayate/V~)-\01=) +l-/clrlelajelerei=
Mrs. Nratv, Exhibition of a living Tree-Hyrax (Dendrohyrax dorsalis) .......+.. «++
Dr. H. A. Bayuis, F.Z.S. Exhibition of a Nematode worm, Zoxascaris leonina .....
Mr. E, A. Sravt, F.Z.S. Exhibition of specimens showing metamorphosis of Axolotls
induced by injection of pituitary gland anterior lobe extract ............. ooo a0oG
Page
655
655
655
656
657
657
657
657
657
657
657
Contents continued on page 3 af Wrapper.
INDEX.
1923.—Pages 483-1097.
[New names in clarendon type. Systematic references in italics.
(z.8.L.) indicates additions to the Society’s Menagerie. |
Abdimia abdimi, 918.
Ablepharus, 861.
lineo-ocellatus var. adelaidensis,
1077.
—- wahlbergit, 963.
Acinonya jubatus raincyt, 721.
Acomys selousi, 704.
Acontias, 862.
meleagris, 964.
Acrocephalus griseldis, 906.
Actias mimose, 1080.
Aipyceros melampus suara, 735.
Atthosciurus byatii, 698.
Agama, 847.
—— atricollis, 946.
—- flavicauda, 946.
—— hispida var. distanti, 942.
—— lionotus var. dodome, var. n.,
944.
var. Mwanze, vi". n,, 945.
-—— mossambica, 9438.
Agapornis personatus, 911.
Alectoris greca kurdistanicus (2.8. w.),
1093.
Algiroides, 853.
—— alleni, 952.
Amnblystoma, 657.
Proc. Zoou. Soc.—1923.
Amphibolurus adelaidensis, 1076.
—— fionniy, sp. n., 1075.
imbricatus, 1074.
Amphisbena, 851.
—— mitchelli, sp. n., 1065.
Amphisbenula, 851.
Amplorhinus nototenia, 882.
Anaphe reticulata, 1032.
Anaplectes rubriceps, 908.
Anarosawrus, 1048,
Anastomus lamelligerus, 919.
Ancylorrhynchus, 1018.
Ancystropus xthiopicus, sp. n.?,
979.
(Meristaspis)
sp. n., 983.
— —— lateralis, 985.
—— —— macroglossi, sp. n., 981.
—— zelebori, 978.
Anthia striatopunctata, 1028.
Anithoscopus caroli, 901.
Anthropopithecus troglodytes (%.8.t.),
1001, 1093. ‘
Antidorcas euchore, 498.
Anychus latus, 991.
Aonyx capensis helios, 713.
calcearatus,
Aparallactus capensis, 889.
Cc
xx INDEX.
Aparallactus jacksonti, 889.
—— werneri, 889.
Apis mellifera adansoni, 1016, 1019.
Aporoscelis, 849,
Apus affinis, 908.
Ardea cinerea cinerea, 917.
—- goliath (z. 8.4.), 1093.
purpurea purpured, 917.
Ardeola ralloides, 917.
Astur? polyzoniodes, 916.
Atelerix hindei sotille, 696.
Atilaax paludinosus rubescens, 717.
Atopomyia, gen. n., 596.
rothschildt, 597.
Atractaspis rostrata, 897.
Avicanthis abyssinicus nuanse, 704,
—~ —— nairobe, 704.
—— —— newmanni, 704.
— — virescens, 704.
— tenebrosus, 705.
Balenoptera musculus, 1085.
Batrachiderpetwm, 1052.
Batrachosuchus, 1052.
Belonogaster griseus, 1023,
Bitis arietans, 894.
gabonica, 896.
Boodon lineatus, 876.
Bos caffer radcliffei, 732.
Brachytrypes membranaceus, 1036.
Bradypus cuculliger, 517.
—— tridactylus, 517.
Broinophila caffra, 1015.
Bubalis coket wembaerensis, 733.
—— lichtensteini, 733.
Bubo africanus africanus, 912,
—— lacteus, 912.
Bufo granulosus, 1061, 1064,
Bunocnemis, 845.
Burhinus capensis capensis, 921.
Burnetia mirabilis, gen. et sp. n.,
671.
Burnetiamorpha, subord. u., 673.
Buteo augur, 915.
Cacops, 1047.
Calamelaps polylepis, 889.
Callispa almora, sp. n., 599.
Camelus dromedarius, 543.
Camponotus (Orthonotomyrmex) sericeus,
1027,
Campothera sp., 911,
Camptosaurus, 1046, 1055.
Canis mesomelas, 489.
—— thous (z. 8. t.), 1094.
, 1094.
Caparinia tripilis, 1000.
~ Caprimulgus europeus imeridionalis,
909.
—— fossei mossambicus, 909.
Cardiocondyla emeryi, 1026.
Cardioderma cor, 693.
Cassida dimbakar, sp. n., 607.
——- manipuria, sp. n., 606.
— rati, sp. n., 605.
Causus defilippit, 894.
— lichtensteini, 894.
—— resimus, 893.
rhombratus, 893.
Centetes, 523.
Centropus superciliosus intermedius,
910.
-—— loande, 910. .
Cephalophus grinmi shirensis, 734.
melanorheus schusteri, 734.
Cerceris, sp., 1017.
Cerchneis naumannt, 918,
tinnunculus tinnunculus, 913.
Cercocebus albigena johnstont, 687.
—— aterrimus (2. 8. u.), 10938, 1095.
-—- congicus, 1095.
Cercopithecus albogularis rufilatus, 688.
—— doggetti, 687. .
—— pygerythrus, 486.
centralis, 688.
-—— —— johnstont, 688.
Cherephon emini, 699.
limbatus, 695.
Chalicotheroidea, 1005.
Chameleon biteniatus, 967.
—— dilepsis dilepsis, 966.
— hochnellii, 967.
—— yacksonit yar, varerescece, 267.
—— melleri, 968.
parvilobus, O64.
INDEX,
Chameleon senegalensis, 964.
Chamesaura, 849.
Chametorus aulicus, 882.
Chelone imbricata, 933.
mydas, 930.
Chirindia, 851.
Chlamydophorus truncatus, 517,
Chlorophis irregularis, 878.
— — neglectus, 878.
—— teniopus, 783.
Cheropsis liberiensis, 531, 1096.
(z. 8. t.), LO96.
Choiropotamus choiropotamus demonis,
737.
Cholepus didactylus, 517.
Chrysidide, 1018.
Chrysochioris, 524.
Cicindela brevicollis, 1028.
Cinixys belliana, 924, 931,
Cinnyris loveridget, 900.
senegalensis inestimata, 900.
Circaétus cinereus, 915.
JSasciolatus, 915.
Cireus macrourus, 916.
Civettictis civetta orientalis, 718,
Clonia wahlbergi, 1087.
Coluber hodgsoni (z. 8. u.), 1094.
Compsognathus, 1046. |
Condylura, 524.
Connochetes taurinus taurinus, 733.
Cophias flavescens, 1065.
Coracias garrulus garrulus, 909.
Coracina pectoralis, 905.
Coronella scheffleri, 880.
semiornata, 880.
Corvultur albicollis, 903.
Corvus scapulatus, 904.
Corythosaurus, 1046.
Cremastogaster castanca, 1025.
Cricetomys gambianus osgoodi, 703.
Cristicola nana, 906.
Crocidura bicolor elgonius, 698.
flavescens, 697.
--—— hindei, 698,
_—— hirta, 697.
—— martiensseni, 697.
—- russula cintre, 1043.
peta, subsp. n., 1044.
Proc, Zoou, Soc.—1923.
Crocidura russula pulchra, 1048.
—- russula, 1043.
Crocuta crocuta germinans, 721,
Cryptoprocta ferox (2. 8. u.), 1094.
Cuculus canorus gularis, 910.
Cynictis pennicillata, 488.
Custecalia absidata, 1038,
Dactylopteryx sp., 1039.
Dacus brevistylus, 1017.
pectoralis, 1017.
Damaliscus korrigum jimela, 733.
Dasypeltis scabra, 880.
Delphinosaurus, 1051.
Demodex sciurinus, sp. n., 995.
Dendraspis angusticeps, 893.
Dendrelaphis caudolineatus (2.8.1),
656.
Dendromus ochropus, 700.
—— pumilio, 700.
Dermanyssus (Allodermanyssus)
sanguineus, 975.
Dermochelys, 1048, 1051.
Desmodillus auricularis, 490.
Diadectes, 1047, 1053.
Dicotyles pecari, 536.
tajacu, 536. +
Dicrurus afer lugubris, 904.
Diemenia textilis, 1077.
Dinemellia bohm, 903.
Dinophoneus ingens, goui. et sp. 1.,
666.
XX1
Diphyllodes magnifica hunsteini, 609, .
657.
Diplocaulus, 1052.
Diplodactylus, 843.
tessellatus, 1074.
Dipoditlus luteus, 699.
Dispholidus typus, 888.
Dolichomutilla guineensis, 1024.
Dorylus helvolus, 1027.
Downesia sasthi, sp. n., 600.
Dreata (Jana) sp., 1034. meee
Eidolon heluum, 692.
Hlanus ceruleus, 913 °
xxii INDEX.
Flapechis guentheri, 889.
-— niger, 890.
Flaps marcgravii, 1067.
—— spixii, 1063, 1067.
Elasmodactylus, 847.
-— triedrus, 942.
Elasmosaurus, 1051.
Hlephantulus ocularis, 697.
—— pulcher, 697.
— renatus, 697.
—— rupestris, 487.
Hlephas africanus knockenhauert, 738.
—— maximus (4. 8..), 652.
Emberiza flaviventris, 902.
Enobius strubeni, gen. ct sp. n.,
683.
Enyaliopsis, 1037.
Ephippior hynchus senegalensis, 918.
Epomophor us labiatus, 692.
Lptesicus capensis, 486.
Equus quagga crawshayi, 738.
Eremias, 855.
-—— spekii, 953.
Evremomela flaviventris tardinata, 906.
Erinaceus, 523.
Eristalodes quinquemaculatus, 1016.
Eryops, 1047.
Tiwmenes dyschyr aides, 1015.
maxillosa, 1015.
Humerus sp., 1017.
Euparkeria, 1055.
Eupodotis wrabs on 1..), 657.
Eurytela hiarbas, 1029.
Eutolmaétus spilogaster, 914,
Evotomys glareolus sobrus,
subsp. n., 867.
—— gorka, sp. n., 867.
Taleo ruficollis, 918.
—— subbuteo subbuteo, 9138.
Felis capensis hindei, 780.
—— concolor (z.s.u.), 1093, 1094, -
1097.
—— leo (z.s.u.), 1094.
—— —— massaica, 722,
— ocreata cafra, 487.
Felis ocreata ugande, 731.
onca (4. 8.1..), 1097.
pardalis (4.8. ..), L094.
-— pardinoides (4. 8.1.), LOO.
pardus, 487.
suahelica, 727.
servwl (4%. 8.1.), 1097,
Feylinia, 863.
Galago panganiensis, 691.
sennaariensis, 691.
Galeopithecus volans, 522.
Gallinula angulata, 920.
chloropus brachyptera, 920.
Gastropholis, 854.
—— vittata, 9538.
Gazella thomsont, 736.
Genetta dongakana neumanni, 714.
—— felina, 488.
—— suahelica, 715.
stuhlmanni stuhlmanni, 716.
Geocalamus, 851.
acutus, 949. -
modestus, 949.
Geodipsas proctere, 881.
Geosciurus capensis namaquensis, 489,
Gerrhosaurus, 856.
—— major, 954.
nigrolineatus, 955,
Givaffa camelopardalis (4. s.%.), 657,
1094,
Glaucidium perlatum, 912.
—— capense scheffleri, 912. -
Glauconia albifrons, 1061.
distanti, 874.
—— emini, 874,
—— longivauda, 875.
—— merkeri, 874.
Glauconycteris argentata, 694.
variegata, 695.
Glis glis postus, subsp. n., 866.
Gonatodes, 843.
- Grammomys surdaster surdaster,, 701.
Grayia tholiont, 880.
Gymnodactylus lorie, 1069.
-—— lousiadensis, 1069.
Gymnura raffles, 523,
INDEX.
Hagedashia hagedash, 916.
Halcyon leweocephatus centratis, 910.
Harpolestes senegalus orientalis, 905.
Helicops polylepis, 1062. .
Heliophobius emini, 707.
Heliosciurus rufobrachiatus nyanse, 698.
undulatus undulatus, 698.
Heloderma, 1055,
Helogale undulata undulata, 719.
-— viclorina, 719.
Hemidactylus, 843.
-—— brookii, 940.
citernii, 939.
—— mabouia, 936.
-—— sguamiulutus, 939.
Herpestes (Calogale) flaviventris, 716.
—— —~ gracilis ibee, 717.
lademanni, 716.
—— —— granti, 716.
melanurus rufescens, 716. --
ichneumon funestus, 717.
ratlamuchi, 488.
Heleronota binoei, 1074.
Hieractus wahlbergi, 914.
Hippopotamus amphibius, 499, 534.
phacocherus, 1095.
Hipposideros caffer, 653.
—- marungensis, O94.
-—— ruber, 694.
Hippotragus equinus langheldi, 736.
—— niger roosevelt, 736,
ITirundo puella abyssinica, 907. -
smithii smithii, 907.
Holaspis, 856.
Homalosoma lutriz, 880.
Hoplocephatus bitorquatus, 1073.
Hyena hyena schillingsi, 720.
Hyla lesueurii, 1071.
Hyperechia bifasciata, 1014.
Hypsilophodon, 1046.
Hystrix africe-australis, 496.
——-- galeata, 706.
Tearia ambigua, 1016.
Ichneumia albicauda ibeana, 717.
Ichnotropis, 834,
Ictonyx striatus, 489.
XX111
Ietonyx striatus albescens,713.
Idolwm diabolicum, 1041.
Iguanodon, 1046.
Ilysia seytale, 1062.
Ischioden scutellare, 1018.:
Kari, gen. n., 602. —
-——— brunnea, ‘sp. n.; 602.
Kaupifalco monogrammica, 915.
Kabus ellipsiprymnus, 735, |
Kritosaurus, 1046.
Labidosaurus, 1047. -
Lacerta, 852.
jacksoni, 952. —
Lelaps ugandanus, sp. n., 971.
Lagonosticta senegalla, 902.
Lama glama (2. 8. U.), 1094.
huanacus, 548.
—— vicugna, 543. .
Lamarckiana, 1037.
Lanius cabanisi, 904.
collurio, 905.
Latastia, 854.
johnstont, 952.
—— longicaudata, 953.
Lavia frons rex, 693.
Leggada bella bella, 703. :
Lemniscomys albolineatus, 705.
—— griselda rosalia, 706.
maceulus macculus, 705.
Leptodira hotambeia, 882.
Lepus capensis crawshayi, 709.
-— saxatilis aurantii, subsp. u.,
497. th am
-— victoria, 408.
Lialis jicart, 1069.
| Ligidium bosniense, 837.
-— fragile, 831.
gracile, 832.
—— germanicum, 833.
-—— hypnorum, 829.
—— yaponicum, 836.
— latum, sp. n., 834,
longicaudatum, 838.
—— nodulosum, 837.
XXIV
Ligidium (Typholigidium) cacum, 838.
Limnodynastes fletchert, 1071.
Liogryllus bimaculatus, 1037,
. Liphistius malayanus, sp. n., 770.
Liponycteris nudiventris, 692, -
Liponyssus gordonensis, sp. n.,
973. '
—- sylviarum, 979. :
Listrophoroides, gen. n,, 999.
—— ethiopicus, sp. n., 999.
Listrophorus bottre, sp. n., 999.
Lophionys imhaust, 1091,
Lophoceros jacksontt, 910.
Lophuromys aquilus aquilus, 703.
Loxodoa africanus (z. 8, u.), 1094.
Lutra maculicollis, 489.
Lyhius albicauda albicauda, 911.
Lycaon pictus, 710.
Lycophidium acutirostre, 877.
capense, 878, 1080.
jacksonti, 878.
—— semiannulis, 878.
Lygodactylus, 845.
— grotei, 940,
picturatus, 941,
Jischeri scheffleri, 940.
Lygosoma, 859.
—— ferrandti, 962.
—— sundevalli (z. 8..u.), 962, 1095.
—— (Emoa) battersbyi, sp. n.,
1070.
—— (Hinulia) monotropis, 1072.
—— —— quoyi, 1072.
tenure, 1072.
—— (Homolepida) mpobergi, 1073.
-— (Liolepisina) challengeri, 1073,
entrecasteauxti, 1076.
—- (Rhodona) bipes, 1077.
bougainvilliz, 1077.
—— —— punctatovittatum, 1073.
Lynx caracal nubicus, 732.
Mabwia, 857.
—— comorensis, 956.
—— diesneri, 957.
—— hildebrandtii, 959.
INDEX.
Mabuia irregularis, 962.
—— maculilabris, 956.
—— obsti, 960.
planifrons, 956.
striata, 960.
—— varia, DDR.
yar. isselit, 950,
Malaconotus poliocephalus blanchoti, 905.
Manis javanica, 517.
= pentadactyla, 517.
tetradactyla, 517.
Mantis religiosa, 1041. -
Martes flavigula (z.s.1.), 1094.
Mazxera marchali, 1032.
Megachile sp., 1016.
pilicrus, 1018.
Melanoseps, 862.
ater var. longicauda, 963.
Melieraxanoru s metabates, 915.
poliopterus, 915.
Melipona braunst, 1019.
Melittophagus pusillus meridionalis, 909.
Mellivora capensis, 712.
ratel, 489.
Mephitis amazonicus (z. 8. u.), 1094.
Merops apiaster apiaster, 909.
Mesophoyx intermedius brachyrhynchus,
917,
Microdon sp., 1018.
Microgale, 523.
Microgomphodon, 1053.
Microparra capensis, 921.
Microtus agrestis punctus, subsp. n., 868.
arvalis arvalis, 868.
—— —— calypsus, subsp. n., 869.
- levis, 869.
Milvuus migrans parasitus, 913.
Mops osborni, 695,
Motacilla aguimp, $01.
Mungos mungo colonus, 719.
Mus musculus, 492, 703.
Mycetes caraya (2, 8.1.), 1094.
Muylabris dicineta, 102),
—- oculata var. tricolor, 1028.
Myogale, 524.
Myrmecophaga didactyla, 517.
—— guiata (4.8 1.), 517, L094,
INDEX:
Naia hate, 892.
nigricollis, 890.
Necrosyrtes monachus, 916.
Negritomyia maculipennis, 1016.
Nephele peneus peneus, 1030.
Neusticosaurus, 1048, 1051.
Nomia sp., 1017.
Nucras, 852.
emini, 949.
kilose, 951.
Numida mitrata, 920.
—— coronata reichenowi, 920.
Nycteris capensis, 692.
—— damarensis, 486.
— hispida, 698.
—-- luteola, 693.
marica, 693.
Nyctinomus bocaget, 486.
Odynerus tropicalis, 1016.
(cophylla smaragdina, 1027.
dura tryoni, 1071.
Omomantis zebrata, 1040.
Orectocera diabolus, 1016.
Oreotragus oreotraqus, 499.
schillingst, 734.
Ornithlestes, 1046.
Ornithorhynchus, 635, 643, 647.
Orycteropus capensis, 518.
Otocyon virgatus, 711.
Otomys angioniensis elassodon, 705.
—— nyille canescens, 705.
Otus leucotis? grantt, 912.
Ourcbia cottoni, 734.
Oxybelis boulengeri, sp. n., 1062.
Oxyrhopus trigeminus, 1066.
Pachydactylus, 846.
bibronit, 941.
—— boulengeri, 941.
Palophus greyt, 1039.
Paltothyreus tarsatus, 1025.
Paniscus epaculus, 1017.
Pantodactylus schreibersii, 1061.
Pantophthalmus argyropastus, 671.
* acy
Pantophthalmus batesi, sp. n.,
566.
-—— bellardii, 587.
— chuni, 576.
| —— conspicabilis, sp. n., 577.
—— conspicuus, sp. n., 577.
—— frauenfeldt, 565.
hellerianus, 569...
—— pictus, 575.
—— splendidus, sp. n., 589.
—— tabaninus, 562.
—— variegatus, sp. n., 592.
—— versicolor, sp. n., 572.
— vittatus, 586..
Papilio demodocus, 1029, 1030.
Papio cynocephalus, 689.
neumannt, 690.
tessellatum, 690.
Paratetranychus indicus, sp. ‘n.,
990.
Parasxerus ochraceus ochraceus, 699.
palliatus suahelicus, 698.
Parotomys (Liotomys) littledalei, 490.
Passer domesticus, 741.
Pedetes cafer, 496.
— — surdaster, 706. °
Pelomedusa galeata, 930.
Pelomys fallax fallax, 764.
Pelusios nigricans, 930.
Periglischrus interruptus, 987. -
Peripatus, 658.
Peropus variegatus, 1074.
Petrodromus matschie, 696. -
| —— nigriseta, 696.
Petromys typicus, 496.
Pheocherus ethiopicus massaicus, 737.
—-— africants (7%. 8. L.), 657.
Phascolomys miichelli (a. 8. U.), 655.
Phelswma, 846. ;
Philothamnus semivariegatus, 879.
Phyllastrephus cabanisi suecosus, 906.
Phylloperus africanus, 920. Rare
Phymateus viridipes, 1038.
Physocephala sy., 1015.
Pipistrellus kuhlii fuscatus, 694,
—— nanus; 694.
Pison sp., 1017.
Placochelys, 1054.
¥Xvl
Placodus, 1053.
Plagiolepis (Anoplolepis) Ennion,
1026.
Plagtosaurus, 1052.
Plagiostenopterina submetallica, 1016.
Platalea,alba, 917,
leucorodia leucorodia, 917.
Platecarpus, 1051, 1055.
Platypholis, 846.
Platyura sp., 1017.
Plecticus elongatus, 10147.
Podiceps vuficollis capensis, 919, °
Polyborides typicus, 916.
Polychrus marmoratus, 1061.
Polyspilota eruginosa, 1039.
Popitlia ligulata, 1028.
Poreula, 537.
salvania, 539.
Potamocherus hassama (2. 8. L.), 65.
——- porcus, 536.
_ Potamogale velox, 524.
Procavia brucei matschiei, ‘738.
prittwitzt, 739.
capensis, 499.
terricola schusteri, 739.
Procolophon, 1047, 1053.
Procompsognathus, 1045.
Proteles cristatus termes, 720.
Proneusticosaurus, 1051.
Pronolagus crassicaudatus rupestris, 498.
Prosqualodon davidi, 638.
Prozeuglodon atrox, 627, 649.
Psammochares sp., 1014, 1015, 1016.
venans, 1023.
Psammophis angolensis, 887.
- biseriatus, 887.
sibilans, 886.
subteniatus, 884.
Psuedechis australis, 1073.
Pseudocreobotra wahlbergi, 1041.
Pseudorhynchus pungens, 1037.
Pseudotarsonemoides spinitar-
sus, sp. n., 997.
Pycnonotus dodsoni, 906.
tricolor micrus, 905,
Pyrosymna anbigua, 880.
Pytelia afra cinerigula, 903.
_ Python sebe, 875.
——
INDEX.
Quelea sanguinirostris intermedia, 903.
Raphiceros campestris neumanni, 734.
Raphiorhynchus planiventris, 596.
Rattus rattus alewandrinus, 701.
—— (Atthomys) walambe pedester, 101.
chrysophilus singide, 702.
—— (Mastomys) coucha, 492.
— —— -— hildebrandtii, 703.
microdon, FO2.
_—— (Praomys) delectorum, 702.
—— ——- namaquensis, 491.
Redunca redunca toki, 785.
Ithabdomys pumitlio bechuane, 492.
diminutus, 705.
Rhamnophis jacksonit, 879.
Rhamphiophis ovyrhynchus, 883.
Iehinocalumus meleagris, 889.
Rhinodon typicus, 656.
thinolophus lobatus, 693.
Rhinonyssus (Neonyssoides)
nucifrage, sp. n., 975.
thinoptilus cinctus cinctus, 921.
Rhodogastria vitrea, 1031.
Rhynchiwm sp., 1018.
Rhynchocyon, 524.
——— petersi petersi, 696.
—-— swynnertoni, 696,
Rhynehotragus kirki nye, (35.
Riparia fuligula rufigula, 907.
Rousettus leachi, 692.
Saccostomus hilde, 495.
Sarcophilus harvist (4.8. u.), 655.
Sargus sp., 1017.
Sarkidiornis melanotius afr icanis, 919,
Scalops, 524,
Scapanodon duplessisi, 663.
Scapanus, 524.
Scelotes, 862.
Schistocera yreguria, 1088.
Scolecoseps, 862.
Scopus umbretta bannermanne, 918.
Sebastes marinus, 501.
Sepsina, 862.
Serinus sulphur atus shelleyi, 902.
INDEX.
Serpentarius serpentarius gambiensis
(z. 8. L.), 657. a8
Seymouria, 1047. ;
Stimocephalus nyasse, 878.
Simoides crassipes, 1016.
Sisyrnodytes sp., 1018.
Solenodon paradocurus, 23,
Sorex, 524.
Sphenophryne variabilis, 1071.
Sphodromantis viridis, 1040.
Spizattus bellicosus, 914.
Spreo superbus (7:8. u.), 655.
Steatomys loveridget, 701.
—-muanse, 701.
Slenodactylus, 845.
Sternotherus derbianus, 932.
nigricans, 932.
sinuatus, 932.
Streptopelia semitorquata semitorquata,
OU.
Struthio camelus massaicus, 921.
syriacus (z.8. L.), 1093.
Struthiomimus, 1046.
Stylogaster sp., 1017.
Sus cristatus, 542.
——leucomystax, 512.
scrofa, 535, 542.
Synagris estuans, 1023.
Synagris carinata var, albonotata, 1015.
Tachyoryctes demon, 700.
ibeanus, 700.
Talpa, 524.
ceca ceca, 866.
Tamandua tetradactyla, 517.
Taphozous mauritianus, 692.
Tapinocephatia, subord. n., 671.
Tarbophis semiannulatus, 881.
Tarsonemella, subgen. n., 995.
Tarsonemus translucens, 995.
(Tarsonemella) africanus,
sp. n., 995.
Taterona miliaria siclle, 490.
—— muanse, 700.
-—— swaythlingi, 700.
vicina vicina, 699.
Teracolus casta, 1030.
Testudo loveridgii, 928,
XXV1i
Testudo pardallis, 925.
proctere, sp. n., 928.
torniert, 931.
Tetranychus crategi, 991.
Thalassochelys carctta, 933.
Thallomys nitela, sp. n., 493.
—— scotti, sp. n., 494.
-—- shortridgei, sp. n., 492.
Thelotornis kirtlandii, 887.
Theropitheeus gelada (2.8. L.), 1094,
Thescelesaurus, 1046,
Thos adustus notatus, 709.
—— mesomelas memitlant, 710.
Thrasops rothschildi, 879,
Threskiornis ethiopica, 916.
Lhrinarodon, 1058.
Thryonomys swinderianus
708.
Titanosuchia, subord. n., 671.
Titanosuchus cloetci, 661.
Toxascaris leonina, 657.
Toxochelys, 1051, 1053.
Trachyphonus margaritatus (2.8.1.),
655.
Trachytes? austeni, sp. n., 971.
Tragelaphus scriptus massaicus, 736.
Trematops, 1049.
Treron delalandit granti, 911.
Trimerorhinus triteniatus, 882.
Trionyx triunguis, 938.
Tropidonotus olivaceus, 876.
Tropidophorus assamensis, 779.
—— berdmorei, 776.
cocincinensis, 780,
—— hainanus, sp. n., 779.
—— laotus, sp. n., 777.
—— microlepis, 781.
—— queenslandie, 1073.
robinsont, 778.
sinicus, 780.
thai, 781.
Tylosaurus, 1051.
Tympanocryptis cephalus, 1076,
Typhlops, 1079.
—— dinga, 873.
excentricus, 864.
—— mandensis, 873.
variegatus,
“= gacriiro var, hunbo,, 873.
Tyto alba affinis, 911,
XXVIili
Ureginthus niassensis, 902.
Uranocentredon, 1049.
Ursus anglieus, L097.
Varanus, 850, 105d.
niloticus, 948,
—— weellatus, 948.
var, belli, 1072.
Vulpes chama, 488.
Walierinnesia egyptia 1081.
Nenopeltis unicolor (4%. 8. L.), 656.
Nylocopa caffra, 1014, 1018.
INDEX.
| Xylocopa inconstans, 1014.
nigrita, 1014.
.—— torrida, 1019.
Zamensis arenarius (%. 8. %.), LOL.
Zeuglodon elliotsmithi, sp.n., 625.
— intermedius, sp. n., (29, 649,
652.
osiris, 682, 648, 652.
—— sensitivus, sp. n., 618.
zitteli, 627.
Zonocerus elegans, 1038.
Zonurus, 849.
—— tropidosternum, 947.
PRINTHD BY TAYLOR AND FRANCIS, RED LION COURT, FLEBT STREET,
PROCEEDINGS
OF TIE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS
OF TH
ZOOLOGICAL SOCIETY OF LONDON.
————E
PAPERS.
26. On Mammals collected by Captain Shortridge during
the Perey Sladen and Kaffrarian Museum Expedition
to the Orange River. By O.Lprizetp Tomas, F.R.S.,
F.Z.8., and Martin A. C. Hinton, F.Z.8.
[Received May 15, 1923: Read June 12, 1923. ]
(Published by permission of the Trustees of the British Museum.)
Thanks to the generosity of the Percy Sladen Trustees, and
the kind co-operation of the authorities of the King William’s
Town Museum, Capt. Guy Shortridge, already so well known as
a collector in Burma and elsewhere, has been enabled to make
a collecting expedition to the Middle Orange River, a region
whence the British Museum has received but very few mammals,
and one whose faunal characteristics much needed investigation.
The only considerable collections that have been received from
this direction were one made in Little Namaqualand by Mr. C.
H. B. Grant for the Rudd Exploration *, a later one formed by
Messrs. Woosnam & Dent in Bechuanaland ft, and finally that
obtained on Mr. Woosnam’s expedition to Lake Ngami and
described by Capt. Dollman ¢.
The present collection was obtained in the neighbourhood of
Upington, on the Middle Orange River, and forms a very
important addition to our materials for the study of the mammal
fauna of South-West Africa. It consists of 378 specimens
* See Thomas & Schwann, P. Z.S. 1904, vol. i. p. 171.
+ Schwann, P. Z.S. 1906, vol. i. p. 101.
t Ann. & Mag. N.H. (8) vol. vi. p. 888, 1910.
Proc. Zoou. Soc.—1923, Ne. XXXII, 32
484 MESSRS, OLDFIELD THOMAS AND M. A. O. HINTON ON
representing 34 species, and as these were all obtained in the
course of two months, it forms a striking tribute to Capt. Short-
ridge’s energy and collecting ability.
We have had occasion to describe five species and sub-species
as new, partly from the collection itself, and partly from other
collections which the study of the fresh material has enabled us
more fully to understand. Such more accurate discrimination of
earlier material is by no means the least of the benefits to science
rendered by the collection of such series as the present.
Half. of the collection, including all the types, is, presented to
the National Museum by the Percy Sladen ‘l'rustees, and half is
returned to the Kaffrarian Museum, King William’s Town, of
which Oapt. Shortridge is Curator, He has sent us the following
report on this expedition, and as it gives a good account of the
natural characteristics of the region, we haye thought it well to
publish it nearly in eaxtenso:— ites
“Through the kindness of the Administrator, Sir Frederick
de Waal, special shooting facilities: were obtained; while by
arrangement with Dr. L. Péringuey, Director of the South African
Museum, considerable rail fare reductions were secured.
“During the first six weeks I was joined by Mr. A. White, who
was sent by Dr. Péringuey to gain experience in trapping and
other field work, in which he proved himself a keen and useful
assistant. awa
“Tater, during my visit to and below the Augrabies Falls
(which by crossing a number of side streams were seen while in
flood) I was accompanied by Mr. W. Louw, who acted as
assistant, and Dutch interpreter.
“The country generally, with its sparse rainfall, but remarkable
strip of rich irrigable soil that varies from a width of under a
hundred yards to over two miles on either side of the river, is
in some ways quite like Egypt on a small scale, and should be
capable of great development, although the river itself, with its
dense belt of low trees, and muddy water, is more reminiscent of
the Jordan. |
“This belt is by far the most striking feature in the district.
The trees grow only on the low ground that is periodically flooded,
and consist largely of Willows, Mimosa and other shade trees.
There is little or no real undergrowth, and prior to cultivation
the soil between the trees is bare and loose. An extraordinary
variety of birds collect and breed here.
‘On either side of this alluvial belt the country. is bare and
arid, sparsely covered with very low scrub with here and there a
few isolated kopjes, which are supplemented to the westward by
considerable ranges of rocky hills that ‘commence about halfway
between Upington and Kakamas.
‘In several places on the north bank strips of typical red
Kalahari sand extend to the river, which roughly forms the
southern boundary of the Kalahari. The river itself should not
otherwise prove an important barrier between any two faunistic
MAMMALS COLLECTED ON THE ORANGE RIVER. 485
regions, as it is intersected for nearly its whole length between
Upington and the Falls by innumerable islands which are often
only divided by narrow channels, most of which cease running or
become entirely dry when the river is low ; while again the tree
belt is broken in several places for short distances by rocky hill
ranges that run astride of the river.
“The first three camps (altitude 2,600 ft. approximately) were
made at Louisvale, a new but very progressive irrigation settle-
ment on the south bank of the river, at points between ten
and fifteen miles west from Upington; but as Louisvale is not
marked on small-scale maps, all specimens obtained in that
district were labelled: ‘Near Upington —south bank of Orange
River.’
“These camps, although within a few miles of each other,
enabled me to make collections among kopjes and rocky areas ;
salt-bush and typical open country ; the river tree belt; and near
cultivation.
“The fourth camp (altitude 2,000 ft. approximately) was
made on the north bank of the river, close to the little known
Orange River (Augrabies) Falls, about ninety miles west of
Upington.
“The river here splits up into several streams, the main and
two subsidiary Falls descending into a narrow gorge, said to be
about 500 feet in depth. ‘T'wo others—one almost equal in volume
to the main fall—enter several miles lower down.
‘Except for a few squatters, who are chiefly engaged in
looking after herds of donkeys and goats, there is no population
in the neighbourhood.
“A fifth camp (altitude 1,500 ft. approximately) was made
also on the north bank, about twelve miles below the Falls, a few
miles above where the dry Molopo joins the Orange River, which
nas widened out again into large pools that resemble a string
of Jakes, shut in by high rocky hills and thinly fringed with
trees.
“Except for a few wandering Hottentots the country here is
entirely uninhabited.
“It may be noted that neither Chrysochloris nor Cryptomys
were met with. I found no signs of molehills and was assured
by residents that they did not exist in the region.
- “ Hedgehogs, if not absent, must be extremely rare, as they
appear to be practically unknown along this section of the river.
Shrews may occur, but there is little doubt that they are generally
scarce thr oughout the dry South-Western area.
““Antbears, judging by the Colne ane scarcity of their burrows,
are not plentiful.
“* At Upington I saw part of an old dry skin of a ee that
had been obtained in that district.
‘“‘ A few residents spoke vaguely of Wild Pig from some of the
islands, but their present occurrence is exceedingly doubtful.
a Among the larger game reported to occur within from 50-100
32*
486 MESSRS, OLDFIELD THOMAS AND M, A, C. HINTON ON
miles of Upington in the direction of the Kalahari Desert may be
included :—Red Hartebeest, Blue Wildebeest, Black Wildebeest
(said to exist in a wild state), Gemsbuck, Kudu (a single straggler
was recently reported from near the Falls), Springbuck, Duiker,
Zebra, Lion, Leopard, Wild Dog, Spotted and Brown Hyrena,
Proteles, and Cynelurus.”
1, CERCOPITHECUS PYGERYTHRUS Cuv.
3. 228. 2. 7,195. Louisvale, near Upington. 2600’.
¢. 343. Augrabies Falls, 90 miles W. of Upington. 2000’.
‘ Plentiful among the trees that fringe the banks of the Orange
River, probably rarely if ever leaving the wooded course of the
river, along which it is said to extend as far as its mouth,
although, as there is an extensive break in the wooded area below
the Falls, this remains to be confirmed.
Like the Burmese and Malay Crab-eating Monkeys they take
readily to water when hunted. Often doing a considerable
amount of damage among crops and fruit; their consequent
persecution has rendered them rather shy.
Baboons were observed among the rocky hills at and below the
Falls; but nowhere further east.”
2. NYcrERIS DAMARENSIS Pet.
377, 378. Louisvale.
The specific distinction of damarensis from capensis depends
mainly on the greater size of the ears, but there is some variation
in the size of these organs, and every probability that the two
forms will be found to intergrade. Dr. Andersen, however, has
recognised damarensis asa species, and we therefore provisionally
follow his example.
3, Eprestcus cAPENSIS A. Sm.
_Six males, thirty-one females, in skin, and seven in spirit.
Louisvale.
‘By far the most plentiful bat in the district. Appearing well
before dusk, frequently collecting in large numbers over water.
Most of the specimens obtained were found roosting in hollow
trees along the banks of the Orange River; but also hiding by day
under the roofs of houses.”
4, Nycrinomus Bocacrl Seabra.
3. 245, 250, 251, 314, 379. 9. 267, 315.
“Obtained in hollow trees; singly or in pairs; often in
company with Hptesicus. Seldom observed in flight, but possibly
a late flyer.
A Fruit-bat, considered to be somewhat rare, occurs around
Upington, having possibly found its way there since the culti-
vation of fruit was started.”
MAMMALS COLLECTED ON THE ORANGE RIVER. 487
5. ELEPHANTULUS RUPESTRIS A. Sm.
3. 24, 60, 86, 89, 126, 133, 146, 153, 226. 9. 52, 69, 114,
123, 124, 132, 145. Louisvale.
“ A rock Elephant-shrew, the only species observed, although
an individual described from open country near Upington on the
north bank of the river may prove to be a species of Aacro-
scelides.
There is a narrow naked gland, most conspicuous in adult males,
and exuding moisture when fresh, on the underside of the tail,
about half an inch from its base, which is easily noticeable during
life owing to the slight swelling of that portion, and has a distinct
although inoffensive smell of musk.
Elephant-shrews are amongst the most attractive of all small
mammals. When caught alive they are quiet and seemingly not
very timid; never attempting to bite even if a finger is placed
inside their mouths. When handled, the snout—which is very
sensitive—is moved about in a circular manner to avoid any.
contact. Extremely active; their movements are Jerboa-like,
and when hopping among rocks curiously resemble a small ball
bouncing about. Although normally diurnal, during the hot
season they are more frequently to be seen at dusk, and even on
moonlight nights. This appears to be the case with a large
number of small desert mammals, many of which may become
diurnal during the cold weather. Although without doubt chiefly
insectivorous, they may occasionally be taken in traps baited with
mealies. No Elephant-shrews were observed in the rocky country
around the Augrabies Falls.”
6. Fevis parpus L.
3. 374. Kakamas, 60 miles W. of Upington. 2300’.
“ Rarealong the Orange River between Kakamasand Upington,
although apparently becoming rather more numerous in the rocky
hill country in the neighbourhood of the Augrabies Falls.
Said also, together with Cynelwrus, to become more plentiful
northwards in the Kalahari area.”
7. FELIS OCREATA CAFRA Desm.
@. 206. Louisvale. 2600’.
354. Skull only. Upington district.
“Fairly plentiful, and, as usual, very destructive to poultry and
small game. Very numerous further north, judging by the
number of Karosses sent down.
A cat that is undoubtedly 7. nigripes has been described to
me from the Zwart Modder district, some 100 miles north of this
river.
Karosses made from the skins of J’. serval and J. caracal are
488 MESSRS. OLDFIELD THOMAS AND M. A. C. HINTON ON
frequently obtained from. Reitfontein and elsewhere in the
neighbourhood of the Kalahari.”
8. Generra FELINA Thunb.
d. 350. Louisvale. 2600’.
364. Skull only. Upington district.
363. Skull only. Augrabies Falls.
“Said to be fairly plentiful, and evidently becoming decidedly
more numerous further north, judging from the number of
Karosses sent down from Reitfontein and elsewhere.”
9. HERPESTES RATLAMUCHI A. Smith,
go. 19. @. 8,26. Louisvale. 2600’.
365, 366, 367. Skulls only. Upington district.
Hitherto the type of this species—B.M. No. 46.6, 1.14—has
been the only example of it in the British Museum, so that these
‘specimens form a valuable accession. That type, which is also
the type of Smith’s badius, was obtained “ between Latakoo and
the tropic” —probably therefore on the Molopo River.
“Not plentiful. In this district at any rate, like the Vervet
and local tree-rat, they seem to be strictly confined to the belt of
trees on either side of the Orange River.
Said never to occur among kopjes or in open country. Although
occasionally living in burrows excavated either by themselves 01,
Zorillas they are very largely arboreal, and are active tree-climbers,
which would be necessary in an area so frequently inundated.
To a certain extent diurnal.
They are said occasionally to take to a vegetarian diet, and to
be fond of digging up and eating ground-nuts.. One specimen
was caught in a trap baited with mealies.
A grey mongoose—probably HW. pulveraulentus
observed in rocky country away from the river.”
was twice
10. Cynroris penicityata G. Cuv.
3. 14, 22, 213, 244. 9. 20, 189.
Probably referable to C. ». pallidior Thos. and Schw., of which
they would appear to be bleached summer examples.
“ Plentiful, diurnal, gregarious.
These meerkats were nearly always found living with or close
to colonies of Xerus capensis, and they do not appear to interfere
with each other. In any case, it is impossible to distinguish
their. warrens,
I never observed Suricata in this district. Wherever it is
plentiful it is very conspicuous and easily obtained, frequenting
the same localities as Cynictis and Geosciwrus; so that, even if
according to a few residents it actually does occur locally, it can
be by no means numerous. It is reported from the country
round Kenhart.”
MAMMALS COLLECTED ON THE ORANGE RIVER. 489
11. Canis MESOMELAS Schr.
@. 344. Augrabies Falls. 2000’.
353. Skull only.
“ Fairly numerous, especially in the neighbourhood of hills or
rocky country. As with other small carnivora, judging from
local Karosses, becoming considerably more abundant further
north.
African jackals are far less noisy than the Indian species, which
may be heard during the night at all times.”
12. VubpEs cHamaA A, Sm.
356-8. Skulls only. Upington district.
These skulls were purchased. The exact locality where they
were obtained is unknown, except that it was in the vicinity of
the Orange River.
‘As Karosses made from fox skins from Reitfontein are
scarce and rather expensive, it appears that they are either com-
paratively rare or difficult to procure. Said to exist close to the
Augrabies Falls, but I could obtain no other information regarding
their occurrence near the river.”
13. MELLIVoRA RATEL Sparrm.
352. Skull only. Upington district.
“Widely distributed, but not plentiful. Entirely nocturnal
and seldom seen. ‘T'racks were observed near the Augrabies
Falls, while among rocks a deserted bees’-nest was found that had
evidently been disturbed by one of these animals.”
14. Icronyx srriarus Perry.
3d. 72, 121, 159, 202, 248. 9. 18, 119, 120, 188, 249.
Louisvale.
368. Skull ong. Upington district.
“Plentiful, nocturnal. Very easy to trap. It is easily tamed,
even if caught when nearly or entirely full-grown, when it will
seldom, if ever, make use of its offensive scent.”
15. Lurra mAcuticotiis Licht.
@. 322. Young, sex unknown, 321. Louisvale. 2600’.
‘Not plentiful. If, as is reported, Aonyx capensis also occurs
in this district, this species is probably the more numerous. Very
few otter tracks were observed any where along the river.”
16. GrosCcIURUS CAPENSIS NAMAQUENSIS Tight!
gd. 18, 21, 201, 311. ©. 12,190. Louisvale, near Upington,
S. bank of river. 2600’.
490 MESSRS. OLDFIELD THOMAS AND M. A. C. HIN'TON ON
These specimens are quite unlike those of G. capensis pre-
viously in the Museum, and show, by their pale cinnamon-bufly
colour, that the Ground Squirrel of the Kalahari and Nama-
qualand should be subspecifically distinguished from the ordinary
more brownish form of the rest of the colony.
Although Lichtenstein’s description of his Sciwrus namaquensis*
is curiously inapplicable to Geosciurus— corpore supra nigro ;
subtus brunneo,” yet the locality, the presence of white lateral
lines, and the universal assignation of his name to the Cape
Ground Squirrel, seem to render it certain that it really was
referable to this animal, and we can only suppose that he had a
very dirty specimen in the old Holthuisen collection. This
squirrel is also the Sciwrus levaillanti of Kuhl.
‘‘ Plentiful, diurnal, gregarious.
During the hot weather their coats become particularly short
and ragged.
There is no doubt that Geosciurus, Cynictis, and also Suricata
where it occurs, frequently inhabit the same warrens, as traps
often catch them alternately.
Probably the thickness of their skins, in addition to their
unusually great muscular strength, protect them to a great extent
from the smaller carnivores. These squirrels are savage, and in
captivity do not make good pets, as they are uncertain in temper.”
17. TATERONA MILIARIA STELLA Wr.
3. 49, 69, 200, 240, 243, 246, 247, 263. 9. 50, 51, 158,
212, 215, 286, 312.
These specimens average paler than the original series from
Kuruman, but this would seem to be due to seasonal bleaching,
as Mr. Woosnam’s skins were collected in April and May, and
the present set in full summer—November to January.
“Plentiful. Social, rather than gregarious. Attracted by
cultivation. Nocturnal. Away from settlements chiefly occurring
along the beds of dry water-courses.
Their burrows, which are easy to find, are excavated among the
thick scrubby bushes that grow in such localities.”
18. DEsMopILLUS AURICULARIS A, Sm.
Glo Will, sie, CH, Bhs, Ore ake) We). BOO OA Bile a iO,
178, 179, 205; 209, 216, 217, 232/°233°)23722298) 2995 301, al9e
Louisvale. 2600’.
3. 349. 2. 338. Augrabies Falls.
These specimens vary very considerably in colour, many being
as pale as the Kalahari D. a. pudicus Dollm., while others closely
match the true D. auricularis. It would seem therefore that we
are here on the meeting ground of the two forms.
There seeins a tendency for the darker coloured animals to have
* Cat. Rer. Nat. p. 2, 1793.
MAMMALS COLLECTED ON THE ORANGE RIVER. 491
larger bull; but many more specimens are needed before this
can be definitely asserted.
“* Plentiful; occurring in flat open country.
Their burrows, which are excavated in open sandy patches, are
circular, small for the size of the animal, and generally more or
less perpendicular, resembling those of an open country elephant-
shrew (Alacroscelides). Nocturnal. Attracted by cultivation, and
said occasionally to appear in large numbers.
Individuals from the same locality varied considerably in
colour.”
19. Paroromys (Liotomys) LITTLEDALEI Thos.
40 specimens. Louisvale.
Quite agree with the specimens from Bushman Land (Kenhart)
obtained and presented by Maj. Littledale. This striking animal
must be very restricted in range as it was only described in 1918
in spite of its evident local abundance.
There is a good deal of variation in the general colour, but the
more brightly toned specimens just match the type from Tuin,
Kenhart.
“Local, but extremely plentiful where they occur; their
_ presence coinciding with the Jarge but rather infrequent patches
of a succulent species of salt-bush, which they appear almost
exclusively to feed on.
Diurnal, not attracted by cultivation, Their burrows, which
are excavated among the roots of thick masses of salt-bush, are
very conspicuous, being interpolated above ground with thick
networks of sticks, and the amount of this work possibly makes
them appear to be even more numerous and more gregarious
than they really are. .
Not readily trapped, this genus being probably less omnivorous
than most other small rodents.”
20. Rarrus (PRAoMYS) NAMAQUENSIS A. Sm.
36 specimens. Louisvale.
3. 325, 326, 333, 329. 9. 327, 331, 334, 336. Augrabies
Falls, 90 miles W. of Upington. Mamme 1-2=6.
‘A rock rat. Plentiful on either side of the Orange River
between Upington and the Augrabies Falls ; but strictly confined
to kopjes and rocky country. Appearing to become rather less
plentiful among the hill ranges below the Falls. This may,
however, be only a local scarcity owing to a shortage of their
favourite food, or on account of frequenting similar situations it
may compete with Petromys.
The occurrence of this rat is easily ascertained, as it closes up
the entrances of rock crevices with large masses of small sticks
and grass, probably as a protection against enemies, these
492 MESSRS, OLDFIELD THOMAS AND M. A. C. HINTON ON
structures being perforated by tunnels just large enough for them
to pass through.
Nocturnal, Feeds largely on the seeds of Kamel-thorn and
other leguminous bushes. Not attracted by cultivation.”
21. Rarrus (Masromys) coucua A. Sm.
3. 57, 151, 191-194 (young), 266, 296. 9. 191, 192 (young),
313, 316. Louisvale.
“Fairly plentiful, but far outnumbered by habdomys pumilio,
Tatera, Desmodillus, etc. Nocturnal: attracted by cultivation,
and sometimes entering houses. Immature individuals are very
different in colour from adults, being dark bluish slate.”
22. Mus muscuuus L.
6. 185, 235, 241. 9. 105, 113, 303, 373. Louisvale.
“Fairly plentiful in stores and houses, but, as elsewhere, has
not taken to an outdoor existence as it has done in Australia ;
possibly finding it difficult to compete in this country with the
large number of small indigenous rodents.
Imported house rats have not yet found their way to settle-
ments along this part of the river, probably on account of the too
recent extension of the railway.”
23. RHABDOMYS PUMILIO BECHUAN#& Thos.
42 specimens. Louisvale. 2600’.
The deep grey Namaqua form grisews is readily distinguishable
from any of these specimens, but the paler deserti of the Kalahari
is more doubtfully separable.
Several examples from Louisvale have the hind foot recorded
as 27 mm. in length, the same as in the original type, while
others have the foot considerably shorter. In colour also there
is marked variation in the general tone.
The skulls have unusually heavy supraorbital ridges, and their
bulle are larger than in most forms of the group—about 7:0 mm.
in length.
“Diurnal. Particularly plentiful in the vicinity of cultivation,
where it probably outnumbered all other small rodents,”
24, THALLOMYS SHORTRIDGET, sp. n.
g- 169, 170 (young). 9. 1,180,181. Louisvale.
“Shot in. tree, Arboreal, not occurring away from the
wooded area along the river bank. Not plentiful. Mammz
0-2=4.’—G. C.8.
A brown species with very slightly developed face markings.
Size about as in nigricauda, or slightly smaller. General colour
above brown—near “ Brussels brown ”—not so greyish as in other
species. Under surface white, but the bases of the hairs are
MAMMALS COLLECTED ON THE ORANGE RIVER. 493
mostly slaty ; in the type of nigricauda they are slaty laterally,
white-based centrally, and in kalaharicus and the Mossamedes
species (nitela, see below) they are wholly white. Face grey, but
the brown of the crown reaches as far forward as between the eyes.
Dark facial markings scarcely developed at all, the darker edges of
the orbit quite narrow and inconspicuous, and not extending
forwards or backwards as an ocular streak. Ears about of the
colour of the head; hairs behind them buffy brown. Hands and
feet white, with darker patches on the metapodials. Tail long,
black, scarcely so thickly hairy as in nigricauda.
Skull with well-marked supraorbital ridges, long palatal fora-
mina, and rather small bulle, much smaller than in Aalaharicus,
rather smaller than in nitela, those of nigricauda not known.
Dimensions of the type, measured in the flesh :-—
Head and body 150 mm.; tail 189; hind foot 29; ear 22.
Skull: greatest length 35-5; condylo-incisive length 34; zygo-
matic breadth 18:5; interorbital breadth 5; palatal foramina 86;
bulla 6°9 ; upper molar series wor n) 5°6,
Type. Old female. B.M. No. 23. 6, 9.156. Original number
180. Collected 2 December, 1922.
This species is distinguishable from 7’. nigricauda by its slaty-
based belly hairs, its larger feet, and the marked reduction in the
development of the black facial markings.
Capt. Shortridge made special mention of this striking species
as being quite new to him, and we have much pleasure in naming
it in his honour.
[In working out Thallomys shortridgei we have found that the
genus would appear to contain six species, which might be
arranged as follows :—
A. Black ZEliomys-like face markings strongly
developed.
a. Bulle comparatively gual about 7 mm. in
length. Angola.. 1. 7. nitela, sp. n.
b. oe pales over 8 mm, in “length. “Hast
aul ve tans sete nee Rem ean ccomaeeee eee 2. T. loringi Hell.
B. Black face Spee less developed.
ce. Face markings medium.
a*, Hairs of chest wholly white, of belly
slaty at base. Namaqualand ...... 3. TZ. longicauda Thos.
62, Hairs of whole underside white to
base. Kalahari .........0..0:0000, 4, DZ. kalaharicus Dollm. :
d. Face markings practically absent.
c®, Belly hairs slaty based. Orange River. 6. 1. shortridge: T. & Ti. '
d*, Belly hairs white at base. Ii. Africa. 6. Z. scotti, sp. n.
The teeth of all are of about the same size—5°6-5°9 for the
upper series—with the exception of 7’. longicauda, in which this
measurement is only about 5:1 mm.
Details of the new species :—
T. nitela, sp. n. ait:
‘Size comparatively large. Central dorsal area buffy brown.
Sides and flanks markedly greyer. Under surface white, the hairs
white to their bases. Forehead grey. A strong black facial line
running from the side of the muzzle through the eye nearly to
494 MESSRS. OLDFIELD THOMAS AND M. A. C. HINTON ON
the ear. Cheeks grey, but a distinct whitish collar running up
from the throat towards the ear. Hands and feet white, with
small darker metapodial patches. ‘Tail strongly pencilled, deep
black.
Skull with bulle smaller than in loringi, larger than in short-
ridget.
Dimensions of the type, measured in the flesh :—
Head and body 160 mm.; tail 191; hind foot 30; ear 22.
Skull: front of incisors to back of m* 18:8; palatal foramina
8:7 ; upper molar series 5:9.
Hab. Mossamedes. ‘Type from Bomboné, altitude 3200’; other
specimens from Ponangkuma 3300’.
Type. Adult male. B.M. No. 9.10.1.49. Original number
17. Collected 11 March, 1906, by Dr. W. J. Ansorge. Six
specimens.
A very well-marked species, with its black facial lines, white
belly, whitish collar, and bushy black tail.
TL’. scotti, sp. n.
General colour pale greyish buffy becoming stronger buffy on
the rump, sides greyer buffy ; under surface sharply defined pure
white, the hairs white to their bases. Head grey; facial streaks
present, in the form of a blackish clouding round the eyes, but
far less developed than in 7’. nitela. ars with fine buffy hairs
and a buffy tuft at their anterior base. An indistinct whitish
collar perceptible below ears. Hands and feet white, the latter
wholly white, the former with small dark metatarsal patches.
Tail as usual greyish brown basally, black for the greater part of
its length, not very bushy.
Dimensions of the type, apparently not fully adult :—
Head and body 140 mm.; tail 150; hind foot 24.
Hab. British East Africa; type from the Yata Plains, two
days’ march East of the Thika River where it joins the ‘lana.
Altitude 4000’.
Type. Immature female. B.M. No. 12.5.19.17. Collected
19 November, 1911 and presented by R. L. Scott, Esq.
By its reduced facial markings and wholly white belly hairs
this species is readily distinguishable from the other East African
form, 7’. loringi, while its paler general colour is also characteristic.
We have named it in honour of its collector Mr. R. L. Scott,
to whom the National Museum owes many important donations. |
‘“‘ Entirely arboreal, and confined to the tree belt that fringes
the Orange River. This belt is liable to partial or complete
inundation during the frequent rising of the river, -being
occasionally under water to a depth of from six to ten feet, at
which times these rats must live entirely among the topmost
branches of the trees.
Shy, and a very active tree climber. Apparently mainly
nocturnal, although occasionally to be seen high up among the
branches of trees in the early evenings before sunset, so that
during the cold season they may become partly diurnal. Breeding
MAMMALS COLLECTED ON 'THE ORANGE RIVER. A495
in hollow branches, the entrances to which are guarded by large
conspicuous structures of sticks, one of these, which was about
fifteen feet from the ground, measuring over six feet in length ;
while another was built under the roof of a seldom-used Kaffir
hut. I did not find this species plentiful, although it is said at
times to occur in comparatively large numbers.”
25. SAccosroMUS HILD& Schw.
6. 59, 161,182. ¢. 98,127. Louisvale.
These specimens agree closely with the original series from
Kuruman, but our attention has been drawn to the set from
Molopo and Lehutitung, in the real Kalahari, determined by
Capt. Dollman as S. anderssoni de Wint.*, which he distinguishes
from hild@ by the slightly lighter colour of the adults and much
lighter colour of the young, specimens of a similar age being
present in both sets.
In this separation we quite agree with him, but not as to the
reference of the Kalahari form to the Damaraland species, the
difference in colour being in our opinion sufficient for the dis-
tinction of the two. We therefore now describe
Saccostomus pager, sp. N.
General colour above very light, nearly matching Ridgway’s
“ drab-grey,” still paler on the sides, much paler and less buffy
than in S. anderssoni, rather paler than in S. halde. Half-grown
specimens like the adults or even paler, while in S. hilde these
are markedly darker than their parents.
Other characters as in S. hilde.
Dimensions of the type, measured in the flesh :—
Head and body 124 mm.; tail 53; hind foot 19; ear 18.
Skull: greatest length 33; condylo-incisive length 31; upper
molar series 48.
Hab. Kalahari and Northern Bechuanaland. Type from
Lehutitung, 3300’; other specimens from the Molopo River.
3000’.
Type. Adult male. B.M. No. 10. 6.3.54. Original number
18. Collected 18 May, 1909, by R. B. Woosnam. Nine
specimens.
The striking difference in the colour of the young is the main
reason for distinguishing this animal from S. hilde, In S. ander-
ssoni the adult is decidedly darker.
We have named this Kalahari species after Mr. John Page,
one of the Kaffrarian Museum Trustees, to whom Capt. Shortridge
has been very much indebted for help in many ways both in
regard to his King William’s Town work, and especially in relation
to his collecting trip to the Kalahari region.
“ Fairly plentiful, but probably owing to the fact that other
gmall rodents occurred in greater numbers in similar localities,
not often trapped.
Nocturnal ; attracted by cultivation.
* Ann. & Mag. N. H. (8) vi. p. 398, 1910.
496 MESSRS. OLDFIELD THOMAS AND M, A. C. HINTON ON
This is one of the species that is said occasionally to occur in
large quantities. This is a peculiarity of many of the small
Karroo rodents, which may in some cases be a migratory move-
ment, but it is equally probable that at times a favourable
season, or temporary absence of some plague or enemy, enables
them to breed in abnormal numbers.”
26. PEDETES CAFER Pall.
do. 122, 208, 220, 222, 223, 320. 9. 41, 165, 174, 207, 218,
219, 231. Louisvale.
“Nocturnal. Plentiful around Louisvale, especially near
cultivation. Rather stupid and clumsy, and, although somewhat
similar to, possessing little of the agility of a Wallaby or even the
Australian Kangaroo Rat (Bettongia), which perhaps they more
resemble in their movements and habits.
Kasy to shoot with a lantern at night. Normally, when
hunted; they make for the nearest burrow. Their large burrows,
which occasionally have an emergency outlet, are usually excavated
in open sandy patches.”
97. Hystrix AFRICH-AUSTRALIS Pet.:
351. Skull only. Upington district.
Apparently rather rare generally, although probably occurring
wherever there are large enough areas of broken rocky country
to afford them plenty of cover.
Tracks, shed quills, and old burrows were observed near and
below the Augrabies Falls.
28. Perromys rypicus A. Sm.
6.330, 3387. 9. 341. Augrabies Falls, Orange River, 90
miles W. of Upington. 2000’.
@. 346, 347. Below Augrabies Falls, 100 miles W. of
Upington. 1500’. 5
' Comparing this series with the set obtained by Mr. Grant
at Klipfontein, Namaqualand, there seems at first sight a
good deal of difference in colour, the present series being
paler. ‘This difference, however, proves to be due to season,
those from Klipfontein having been collected in the winter
and those from Augrabies Falls in the summer. Moreover,
some of the latter show new hair, as strongly coloured as in
the Klipfontein specimens, pushing through the old blackened
fur.
“First ‘obtained among hills and rocks at the Augrabies
Falls, afterwards in similar localities twelve miles below.
Although: it was easy to ascertain where they occurred, they
did not appear to be particularly plentiful, and were not readily
trapped. This may be the eastern limit of their range,
although possibly extending as far as a line of rocky hills that
MAMMALS COLLECTED ON THE ORANGE RIVER. 497
runs’ astride of ‘the’ river ‘about halfway between Kakamas
and Upington. Though previously considered to be entirely
diurnal, during the hot season at any rate they become most
active towards sunset, while several were caught in traps set
overnight.
They creep rather than jump among the rocks, and did not
appear, to be particularly active. Their habit of filling up the
entrances of their hiding places between crevices with a network
of sticks resembles that of Rattus namaquensis, except that
larger and stouter sticks of succulent plants are largely used.
Petr omys, like Thryonomys, is a most difficult animal to prepare,
the skin being of the consistency of wet blotting-paper. It may
be noted that a large area of broken hilly country is often like a
forest in that small mammals. are generally more numerous
among the outside hills and rocks, in the same way that there is
more life near the edge of a forest. A female examined contained
one young.”
29. LePus SAXATILIS AURANTH, subsp. n.
3. 72,73. 9. 74,118. Louisvale. 2600’.
“ Not at all confined to rocky country—-plentiful around
cultivation.”—G. C. 8.
General characters of ordinary saxatilis, the ears not especially
lengthened as in the Namaqua subspecies megalotis. Colour
above, where’ unbleached, rather paler grey than in true saxatilis,
but bleaching in summer to a much greater extent, the dif-
ference in summer specimens of both very striking. Type with
its unbleached hairs “light buff” tipped with black, its bleached
areas wholly dull sandy. Crown bleached dull brownish. Throat-
band pale greyish buffy.
Skull with ver y large bull, markedly larger than in the longer-
eared megalotis.
Dimensions of the type, measur ed in the flesh :—
Head and body 540 mm.; tail 117; hind foot 126; ear 137.
Skull: greatest length 101 ; condylo-incisive length 89; length
of bulla 14,
Type. Old femate. B.M. No. 23.5.9.173. Original number
118. Collected 23 November, 1922.
_ While December specimens of ZL. saxatilis from Central Cape
Colony are little paler or more bleached than winter ones, these
November examples are all strikingly paler than any specimens
of the group that we have seen, whatever time of year they were
killed. Such patches of fresh fur as are present are also some-
what paler than occurs in true saxatilis. The ears of this form
are only of the same length as in true sagaéilis, not of the
extraordinary dimensions of those pf L. s. megalotis, while on the
other hand its bull are larger than those of any others of
the group. . :
498 MESSRS. OLDFIELD THOMAS AND M. A. 0. HINTON ON
‘Rather plentiful. Attracted by cultivation. Like Pedetes
easily shot in cultivated ground by night with a lantern, at which
times they become dazzled by the light and unusually sluggish in
their movements. Sometimes put up by day among thick salt-bush.
Kohl-haas or Rhebok-haas of the Dutch.
Another hare, known locally as Vlack-haas, without doubt a
local form of Lepus capensis, is reported from this district. It is
said to occur in open country and not to be attracted by cultivation.
Described as being shy, solitary, and rather rare.”
30. PrRoNOLAGUS GRASSICAUDATUS RUPESTRIS A. Sm.
3. 15, 142, 149, 150,155. @. 154. Louisvale. 2600’.
9. 332. Augrabies Falls. 2000’.
We are now quite convinced that Smith’s immature type of
Lepus rupestris should be referred to the Namaqualand form, to
which in 1904 Thomas and Schwann assigned the later name of
Lepus melanurus Riipp., the latter being therefore a synonym
of the former.
These specimens are more suffused with rufous than most of
the examples from Namaqualand, but the difference would
appear to be one of season.
‘* Plentiful where they occur, the range coinciding with that
of rocky kopjes and hill ranges, where they are frequently to be
found in considerable numbers ; otherwise not gregarious to the
same degree as a European rabbit. Shy, and rarely appearing
before dusk, and even then seldom wandering far from cover.
Occasionally the entrances of their hiding places under rocks are
protected by an entanglement of sticks and brushwood, but this
may indicate breeding places and afford protection for their
young. When in the open they are comparatively slow as
compared with typical hares, relying more on quickness in dodging
behind rocks or under crevices. Often frequenting the same
localities as Procavia capensis, although many places where they
may be numerous will not afford suitable cover for Dassies, which
prefer kopjes and other situations where there are abundance of |
large loose boulders. These rabbits are difficult to prepare, the
skin being exceedingly tender, especially on the tail. The fur
comes out very easily, and when shot the cloud of fur readily
indicates a hit. They are said to pull out large quantities for
the purpose of making nests for their young.
Klip-haas or Rooi-haas of the Dutch.”
31. ANTIDORCAS EUCHORE Zimm.,
2. 375. Worns only. Upington district.
-. “ Springbuck seldom wander close to the Orange River nowa-
days in this district, but are said to be still reasonably plentiful
in the direction of the Kalahari and elsewhere inland.”.
MAMMALS COLLECTED ON THE ORANGE RIVER. A499
32. OREOTRAGUS OREOTRAGUS Zimm.
3. 340. @. 345. Augrabies Falls.
@. 355. Skull only.
“ Klipspringers exist in small numbers among the rocky hills
and ravines in the vicinity of the Augrabies Falls, and may
become more numerous in the hill country still further west.
A pair were put up on one of the islands near the Falls, which
crossed over a shallow branch of the river to the mainland when
hunted.
I have heard of no other locality eastwards along the river
between the Falls and Upington where they occur, although they
are said to reappear near Prieska.
The hair of Klipspringers was formerly much in demand among
the Boers for stuffing saddles.
Apparently—apart from stragglers—-the only other resident
buck occurring near the river is the Steinbok. A single indi-
vidual was observed about twelve miles south of Louisvale.
They are well known locally, although not considered plentiful.”
33. HipropoTAMUS AMPHIBIUS L.
204. Incisor picked up from bed of river near Upington.
“The portion of a tusk sent was picked up in the bed of the
Orange River near Louisvale, and presented by Mr. D. Biggs.
Although Hippopotami have long been extinct above the
Augrabies Falls, fragments of tusks ave not infrequently picked
up in or near the river bed.
A small school of Hippo are said to exist in the lower reaches
of the Orange River, but as this is a very uninhabited region few
authentic reports are available about them.
Cornell, in his fairly recent book, ‘The Glamour of Prospecting,’
mentions having seen them, so that there is every reason to believe
they are still there.”
34, ProcAviA CAPENSIS Pall.
3. 329, 335. 9. 328, 342. Augrabies Falls. 2000’.
359, 360. Skulls only. Upington district.
361. Skull. Augrabies Falls.
203. Skull. Kopje 12 miles from Louisvale.
‘‘Plentiful among rocks and hills around and below the
Augrabies Falls. There are few suitable localities for Rock Dassies
near Upington; although a skull was picked up on a large isolated
kopje about twelve miles to the south of Louisvale.
Diurnal; but during the hot weather seldom showing them-
selves except during the early mornings and evenings.”
Proc. Zoot. Soc.—1923, No. XX XITI. 33
THE CHONDROCRANIUM OF SEBASTES MARINUS. 501
27. The Chondrocranium of the Teleostean Fish Sebastes
marinus. By N. A. Macxintosn, B.Sc., Demonstrator
in Zoology, Imperial College of Science, South Ken-
sington.*
[Received May 8, 1923: Read June 12, 1923.]
(Text-figures 1—9.)
ConTENTs.
Page
The Minh@e EON scc.ccanoanes casos vsouecone sekooooanedeanoonncen GOL
2. Material and Method ..0.........ccccccccecsccceeesee ove 502
3. The Chondrocranium...............cec000s00eee eee esses. 608
a. Stage 1. The unhatched embryo ......... 503
b. Stage 2. The 4°5 mm. larva.................. 504
e. Stage 3. The 5°5 mm. larva.................. 505
d. Stage 4. The 25 mm. stage ...........0...... 507
Als (SUMMALY Hee ase ce eos nce ee Meee ne LL NRE)
BapAth breviationsieeseaiey ee cere eee TS TS
GyuTbiteralbures. cs ccee cae ete cee LC Lees
1. Inrropucrion.
The work of Parker and subsequently that of Gaupp, Boker, and
others, on the skull of Salmo, constitutes the basis of our knowledge
of the cranial development of the Teleostomi. Further contri-
butions have been made by Stéhr, McMurrich, Allis, Winslow,
Swinnerton, and Pehrson amongst others. In 1910 Allis pub-
lished a paper dealing in some detail with the cranial anatomy of
the mail-cheeked fishes. This paper dealt with the osteology,
and to some extent the latero-sensory canals and muscles of the
adult cranium, but, so far as J am aware, no work has so far been
(lone on the development of the cartilaginous skull of any of the
Scorpenide. It is hoped that the present paper will take a step
towards meeting this deficiency in that it describes the chondro-
cranium of a fairly representative member of the group.
The following description is not an exhaustive comparative
account, and few generalizations will be made ; it deals with the
development of the chondrocranium from a very early stage up
to the time when the cartilaginous elements of the head are
fully developed. Certain comparisons with other forms will be
made where points of special interest or peculiarity arise.
The work was undertaken at the suggestion of Prof. MacBride
and { wish to take the opportunity of thanking him and
Mr. H. G. Cannon for many helpful suggestions.
* Communicated by Prof. E. W. MacBripg, F-.R.S., F.Z.S.
33*
502 MR. N. A. MACKINTOSII ON THE CILONDROCRANIUM
2. Mavertat AND MErHop.
Some difficulty in obtaining all the desirable material has
rendered the series of developmental stages a little incomplete,
for I was unable to procure any specimens between the 5-5 mm.
and 25 mm. stages. his leaves rather a large gap during which
a considerable development of cartilage takes place. It is not
difficult, however, to reconstruct the more important processes
which occur during this period.
T had at my disposal large quantities of specimens from the early
segmentation stages up to the 5-5 mm. stage, and three or four
specimens ranging between 25 and 30 mms. ‘The abundance
early stages compensates to some extent for the lack of some of
of the later stages.
Text-figure 1.
Stage 3. External features.
ft has been found most convenient to divide the following
account into four parts :—
Stagel. The unhatched embryo.
2. The 4:5 mm. larva.
» 3 The 5°5 mm. larva.
,, 4. The 25 mm. post-larval stage.
99
Stages 2 and 3 are close together, but the differences between
them are sufficient to justify their separate consideration. The
specimens for stages 1-3 were supplied from Lowestoft and those
for the 25 mm. stage were supplied by the Board of Fisheries for
Scotland at Aberdeen. Fixation was in every case in formalin.
Consequently it was not easy to follow much of the histological
details of the formation of cartilage, though preservation was
otherwise satisfactory. The principal stains used were thionin
and orange G, Mallory’s triple stain and picronigrosin. For the
first stage specimens were removed from the egg capsule for
sectioning. he diameter of the curled up embryo is at this
stage about 1 mm. ‘The eyes can already be seen with the naked
eye and serve as a useful guide for orientation. The 4°5 mm. larva
has just escaped from the egg capsule and still retains a large
yolk sac. In most respects it closely resembles the 55 mm. larva.
In the latter (text-fig. 1) the yolk sac is considerably reduced
though it is still a fairly substantial body. The principal features
of this larva are the very long tail, the large eyes, and the distinct
cranial flexure. The mouth is still in a more or less ventral
OF THE TELEOSTEAN FISH SEBASTES MARINUS. 503
position, but has shifted further forward than in the 4:5 mm.
stage. The auditory vesicles are fairly well advanced and can be
seen clearly in whole mounts. Large pigment cells are to be
seen over the posterior part of the gut, and there is a row along
the ventral side of the tail. The 25 mm. specimens are of a post-
Jarval stage and exhibit roughly the form of the adult.
Reconstructions were in each case made from series of transverse
sections cut at 10. These were examined with the aid of a
. Squared eyepiece micrometer, and lateral and dorsal views were
plotted directly on to squared paper. The details of this method
are described in a recent paper by Wells (1922) on the chondro-
cranium of Clupea. As it is necessary in this method to have a
Text-figure 2.
Stage 1. Lateral view of chondrocranium.
base line from which all measurements are made it is usual to
assume that some element in the head, generally the notochord
or some piece of cartilage, is straight. I have been able to avoid
this assumption by plotting an accurate outline of the head
before sectioning it, and then taking all measurements from the
outline. Other methods of reconstruction are described by Norman
(1923) in a paper dealing specially with the subject.
3.. THE CHONDROCRANIUM
a, Stage 1. The unhatched embryo.
This is the earliest stage it was found profitable to investigate
(text-fig. 2). The head skeleton is in an extremely elementary
condition, and is represented principally by tracts of procartilage
and connective-tissue cells which are becoming concentrated into
the positions in which cartilage is subsequently to be laid down.
It is always a matter of some difficulty to draw a definite line
between procartilage and cartilage proper. The trabecule at
this stage appear to be in an intermediate condition. ‘Che outline
is quite definite, but the cells have not become thoroughly welded
together to form a solid matrix. The trabecule are very short
504 MR. N. A. MACKINVOSIT ON 'THE CHONDROCRANIUM
paired rods, ending freely at both ends and lying just below the
apex of the notochord and behind the forebrain. There is a
marked cranial flexure. At present there is no sign of the para-
chordals.
The visceral skeleton is represented by a relatively extensive,
but barely differentiated mass of procartilage. In this it is
possible to distinguish on each side an element running down-
wards from the auditory vesicle. This element ultimately gives
rise to the jaw suspension apparatus. At the foot of it the
procartilage runs forwards for a short distance indicating the
position in which Meckel’s cartilage is to be laid down. This
cartilage appears later as a condensation on each side which grows
forwards as the cranial flexure is eased and the mouth moves
forwards. At the present stage the mouth has not appeared.
The rest of the mass of procartilage extends upwards and back-
wards and constitutes the rudiments of the branchial bars.
b. Stage 2. The 4:5 mm. larva.
The trabeculw have at this stage lengthened considerably
(text-fig. 3) and form a roof to the mouth, which has now made
its appearance. The parachordals are now established and are
continuous with the trabeculw, but where they form the floor of
the auditory capsule they are in a procartilaginous condition.
They cannot, however, be distinguished as anterior and posterior
plates as in the case of Salmo, Gasterosteus, ete. The trabeculie
are closely approximated along the whole of their length and end
freely anteriorly, though there is at their extremity a condensation
of connective tissue representing the ethmoid plate. The noto-
chord reaches well forward and turns down under the rudiment
jof the infundibulum.
In the visceral skeleton Meckel’s cartilage has made its appear-
ance, its two components being joined anteriorly by a tract of
procartilage. A substantial bar of cartil age articulates with the
lower jaw near its posterior end and with the | lower surface of the
auditory capsule. ‘This element, which appears to be a piece of
continuous cartilage, later gives rise to the hyomandibular,
symplectic, and quadrate cartilages. Pouchet (1878) employed the
term “ primordial temporal” to a somewhat similar element which
he found in certain forms. As “temporal” is now obsolete
as an alternative name for the hyomandibular, and as I am
unaware of any name at present in use which could rightly be
applied to the bar which, in Sebastes, joins the lower jaw to the
auditory capsule, I propose for convenience to call it the temporal
cartilage. From the posterior side of this temporal cartilage a
branch is given off which turns inwards and forwards as the
ceratohyal to join the anterior end of a mass of cartilage from
which the branchial bars are beginning to be differentiated. The
point at which the ceratohyal branches off is composed of rather
immature cartilage which later becomes more compact and forms
the stylohyal.
OF THE TELEOSTEAN FISH SEBASTES MARINUS. 505
A remarkable feature which has become evident at this stage is
the precocious development of the visceral skeleton. The lower
jaw, temporal, ceratohyal, and to seme extent the first two
branchial bars are al] distinguishable, while the cranium consists
only of the simple parallel trabeculz and the barely established
parachordals. It is interesting at this point to note in connection
with the form assumed by the parachordals that Filatoff (1916)
has shown that if the auditory vesicle of Bufo is transplanted to a
position overlying the trabecule a cartilaginous capsule tends to
Text-figure 3.
Stage 2. Lateral view of chondrocranium.
form round it, while the parachordals from which the vesicle has
been removed do not form a capsule. It is pointed out that
the formation of the cartilaginous capsule is not an inherent
property of the parachordals, but depends upon the presence of
the vesicle.
From now onwards the mouth tends to move forwards and
upwards, and Meckel’s cartilage follows it by growth at the
anterior end.
c. Stage 3. The 5:5 mm. larva.
The trabeculz have still further lengthened and are now joined
anteriorly in an extensive but very thin ethmoid plate (text-fig. 5)
which as yet shows no sign of any of the upgrowths which later
form the investment of the olfactory organs. According to
Gaupp it has been shown that in Salmo salar the cartilage of the
ethmoid plate is formed from the epithelium lining the roof of the
mouth, and is therefore ectodermal in origin. In Sebastes at this
stage it can still be seen from the structure of the ethmoid plate
that it is formed from a single layer of cells. But a study of the
4-5 mm. stage shows that these cells are derived from a local
condensation of connective tissue.
506 MR. N. A. MACKINTOSH ON 'TH1) CHONDROCRANIUM
The notochord still reaches well forward and maintains a
marked flexure (text-fig. 4). Another peculiarity to be mentioned
here is that there is no sign of a fenestra hypophyseos which 1s
normally present in a tropibasic chondrocranium at this stage.
The trabeculw are still closely approximated along the whole of
their length. The parachordals are now well developed and form
broad, saucer-shaped bases to the auditory vesicles.
The visceral skeleton has maintained its advance over the
eranial elements. ‘lhe lower jaw has now attained a considerable
length, but there is still no vestige of an upper jaw. In this
peculiarity the development resembles that of the larval Clupea
Text-figure 4.
Stage 3. Lateral view of chondrocranium.
(Wells, 1922) in which the upper jaw does not appear until the 13
mm, stage. The temporal cartilage (text-fig. 4) has lengthened,
but has otherwise undergone no change except for the fact that
the hyomandibular portion is better defined. ‘here is at present
no foramen for the hyomandibular branch of the VIIth nerve.
It appears that the nerve becomes enclosed in the cartilage later
on. ‘The cartilage joining the ceratohyal to the temporal is now
constricted to a narrow stylohyal. Considerable development has
taken place in the branchial bars (text-fig. 6). There is a median
copula communis to which the ceratohyal and the first three
branchial bars are attached. The fourth bar is free and the fifth
OF THE TELEOSTEAN FISH SEBASTES MARINUS. 507
has not yet appeared. The branchial bars occupy a rather
advanced position. They form a kind of ridge on the floor of the
mouth, the copula communis being at a higher level than the gill-
Text-figure 5.
'Text-figure 6.
Stage 3. Dorsal view of
branchial bars.
Stage 3. Dorsal view of
chondrocranium.
bars which slope downwards and backwards in relation to it.
The interior of the mouth is thus in the form of an inverted V
when viewed in transverse section.
d. Stage 4. The 25 mm. Stage.
The chondrocranium may now be looked on as fairly complete
(text-figs. 7, 8, and 9). A certain amount of ossification has
taken place. As already mentioned I was unable to procure any
specimens between this and the 5°5 mm. stage, and at first glance
it would seem that the gap is so large that it would be a matter
of difficulty to connect the two stages with any profit. However,
by a careful study of the complete chondrocranium and by
analogy with the changes which take place in other forms, it is
possible to reconstruct from a general point of view the processes
which bring about the conditions found in the present stage.
The ethmoid region of the mature chondrocranium of Sebastes
(text-fig. 7) bears a strong resemblance to that of Salmo, and
there can be little doubt that it arises in much the same manner.
In the 25 mm. stage it consists of a substantial mass of cartilage
roughly triangular in shape when viewed from the side. The
508 MR. N. A. MACKINTOSH ON THE CILONDROCRANIUM
base of this triangle consists of a thickened plate representing the
originally thin ethmoid plate. Above and continuous with this
lies the nasal septum which expands posteriorly to form a vertical
pillar, the antorbital planum. The upper part of the antorbital
planum turns backwards in the form of two lateral horns, the
teniw marginales (text-fig. 8) approaching the extremities of the
epiphysial bar and postorbital processes presently to be described.
The antorbital planum is pierced on either side by the foramen
for the olfactory nerve. The ethmoid structure has evidently
arisen from a median ridge and posterior upgrowth on the
ethmoid plate, the former giving rise to the nasal septum and the
latter to the antorbital planum. It is probable that at a slightly
Text-figure 7.
RAP
Stage 4. Lateral view of chondrocranium.
earlier stage the’ teniz marginales were connected with the
epiphysial bar, and afterwards became separated as is the case in
Gasterosteus (Swinnerton, 1902). ‘This point will be referred to
again later.
The fused trabecule are now hardly recognisable as such.
They are represented by two median pieces of cartilage, one of
which is attached to the ethmoid region while the other is a small
piece of free cartilage level with the posterior border of the eye.
The former remnant of the trabecule consists of a thickened base
resulting from the fusion of the two originally separate elements,
bearing a plate-like upgrowth which constitutes an interorbital
OF 'THE TELEOSTEAN FISH SEBASTES MARINUS. 509
septum. ‘This fragmentation of the trabeculae is not uncommon
in Teleosts. A somewhat similar process takes place in Cyclo-
pterus (Uhlmann, 1921).
Above the nasal septum les a slightly elongated rostral
cartilage. It shows no connection either with the ethmoid or
palatine cartilages.
‘The parachordals have given rise to a substantial investment
of the auditory capsules and posterior part of the brain. The
notochord has retreated considerably relative to the auditory
capsules, and is now bridged over by cartilage. Below and in
front of the articulation of the hyomandibular with the auditory
capsule a process projects forwards on either side. These probably
represent the anterior part of the parachordals. ‘The trigeminal
nerve passes out through a notch above them. ‘The auditory
capsules are now completely enclosed and the cartilage is continued
as a roof over the hinder part of the brain. From the side wall
of the brain there runs forward on each side a postorbital process
which joins the ends of a transverse bridge, the epiphysial bar.
This bar (text-fig. 8) cuts off the posterior dorsal fontanelle, and
the fontanelle is divided into lateral halves by a tania tectum
medialis, a thin band of cartilage joining the epiphysial bar with
the posterior covering of the cranial cavity. In Salmo, Amia
(Pehrson, 1922), and others the epiphysial bar, being connected
also with the tenis marginales, forms in addition an ‘anterior
dorsal fontanelle which gradually fills up. This anterior roofing
in does not oceur in Sebastes. In Amiurus (Kindred, 1919) a
permanent anterior fontanelle is formed which, however, does not
close up, while in G'asterosteus the fontanelle is formed, but the
epiphysial cartilage, as is probably the case in Sebastes, becomes
disconnected from the ethmoid region.
It is now necessary to consider how the cranial elements
developed from the parachordals have arisen. It is obvious that
there hag been a vigorous upward growth from the edges of the
cartilage which in the 5:5 mm. stage formed the base of the audi-
tory capsule. ‘This growth has extended so far as to roof over the
posterior part of the cranial cavity. ‘The postorbital processes
have evidently grown forwards from the lateral walls of the
cranium, but it is difficult to say whether the epiphysial bar has
arisen independently or whether it has been formed by the joining
together of processes growing upward from the ends of the post-
orbital processes. Independent formation is the more common
condition among Teleosts. The tania tectum medialis is probably
a backward growth from the epiphysial bar. This is how it is
formed in Cyclopterus, and in Sebastes the anterior part of the
band is broad and strong while the posterior part is very thin and
frail.
The changes undergone by the jaw arches and visceral skeleton
are almost as great as ; these displayed by thecranium. The lower
jaw isa long, rod-like cartilage bearing coronoid and retro-articular
processes at the point of its articulation with the quadrate. In
510 MR. N. A. MACKINTOSH ON THE CHONDROCRANIUM
this respect it resembles the lower jaw of Amia. In place of the
temporal cartilage we have now the’ clearly differentiated
hyomandibular, symplectic, and quadrate. The hyomandibular
presents a rather unusual appearance. It is club-shaped and
very long. At its point of articulation with the auditory capsule
it has a broad plate-like head which is continued downwards as a
comparatively slender stem merging into the symplectic. ‘The
latter is also rather longer than usual and runs down close to
the lower margin of the quadrate. This isa very large plate of
cartilage extended upward and backward as a metapterygoid
process. Posteriorly the quadrate is in intimate association with
Text-figure 8.
Stage 4. Dorsal view of chondrocranium.
the symplectic but is not continuous with it. The upper jaw,
consisting of the palato-pterygoid cartilage, is now established and
runs from the upper anterior angle of the quadrate up to a point
level with the anterior end of the rostral cartilage. The posterior
end of the palato-pterygoid is very slender. It passes close to the
margin of the ethmoid plate but is not articulated with it. At
its extremity it turns outwards (text-fig. 8), reaching almost to
the surface of the snout.
The fact that the upper jaw is connected with the quadrate
only by a very slender piece of cartilage suggests that it may have
arisen independently and grown back to meet the dorso-anterior
OF THE TELEOSTEAN FISH SEBASTES MARINUS. 511
angle of the quadrate. This supposition is strengthened by a -
comparison with the conditions found in the chondrocranium of
the larval Clupea, in which the upper jaw also arises at a late stage
and grows back to join the quadrate in a slender connection. A
somewhat similar process occurs in Amiuwrus and Syngnathus, in
each of which the palatine arises separately.
It appears that the quadrate, symplectic, and hyomandibular
have all arisen from the original temporal cartilage. In such a
case it might be expected that the three elements were formed
separately in the earliest stages, then became fused to form the
temporal and finally separated again. In studying the earliest
stages in which the temporal can be distinguished I have not
been able to trace any division. But in these early stages the
Text-figure 9.
Stage 4. Dorsal view of branchial bars.
cartilage is in an immature condition, and it is very hard always
to be certain whether it is continuous or not. It would also be
difficult: to reconstruct the details of the process by which the
differentiation of the hyomandibular, symplectic, and quadrate
has come about. It is worth noting that in the 5°5 mm. stage the
hyomandibular part of the temporal is very short and that it has
become greatly lengthened in the intervening period. The long
hyomandibular is a feature correlated with the rather unusual
depth of the head in the adult Sebastes.
The stylohyal is now a small cartilage articulated with the
hyomandibular and ceratohyal. The latter is in the form of a
very large plate bearing a posterior process for the articulation
with the stylohyal. The anterior part of the ceratohyal tends to
512 MR. N. A. MAOKIN'TOSIL ON THE CILONDROCRANIUM
become more rod-like. It is possible that the expansive form of
the ceratohyal is correlated with the fact that the gill-bars reach
far forward, in that its position suggests that it serves as a
protection to the anterior parts of the bars, the posterior parts
being of course protected later by the development of the
opercular bones. The anterior junction of the ceratohyals is
surmounted by a short tongue of cartilage, and there passes back
a median bar, the basi-branchial, corresponding to the copula
communis (text-fig. 9). ‘To this bar are attached the first four
branchials. The fifth pair are free but articulate with one
another at their bases. The third pair is peculiar in that the
base of each is produced forwards in two downwardly directed
processes. The ceratobranchials are very long and _ fairly
stout near their articulation with the basibranchial; further
back they become very slender, but before meeting the epibran-
chials they become thicker again. Perhaps the most important
development of the branchial elements lies in the appear-
ance of the epibranchials. These are possessed by the first
four bars but not by the fifth. They are short in comparison
with the ceratobranchials and of a slightly irregular, twisted
shape.
Here again it is difficult to fill in any details during the
preceding period of development. This, however, is not of great
importance as the branchial bars have not undergone any striking
changes beyond the addition of the fifth bar, the epibranchials,
and a few subsidiary elements.
4, SuMMARY.
. There is a precocious development of the visceral skeleton.
2. In at least the earlier stages there is no sign of a fenestra
hypophyseos.
3. The cranial flexure is retained for a considerable period after
hatching.
4. In the 5°55 mm. stage the hyomandibular, symplectic, and
quadrate are represented by a simple temporal cartilage.
This appears also to be a single piece of cartilage in the
earlier stages.
5. The upper jaw does not appear until a comparatively late stage,
and there is evidence that it arises independently and grows
back to meet the quadrate.
6. The anterior part of the trabecule gives rise to an interorbital
septum of rather limited area. .
7. There is a free rostral cartilage situated above the nasal
septum.
8, The lower jaw is Jong and exhibits prominent coronoid and
. retro-articular processes.
9. The hyomandibular is club-shaped and very long. Its length
is correlated with the special depth of the head,
OF THE TELEOSTEAN FISH SEBASTES MARINUS. 513
5. ABBREVIATIONS.
Aud.c....... Auditory capsule. ION, G50600 Notochord.
BEB ae eae: Basibranchial. INES Foaeeeens Nasal septum.
Br. ......... Branchial bars. - Par. ......... Parachordals.
C.Br. ...... Ceratobranchials. IPERS Savenes Palato-pterygoid.
C.Com. .... Copula communis. Po.P. ...... Postorbital process.
Cane Ceratohyal. Qin: Quadrate.
C.P. ......... Coronoid process. RAP. ...... Retro-articular process.
Ep.B. ...... Epiphysial bar. RC. ......... Rostral cartilage.
Ep.Br. ...... Epibranchials. Sty. ......... Stylohyal.
Beth. ......... Ethmoid plate. Sy. ......... Symplectic.
F.0........., Olfactory foramen. T'mp. ...... Temporal.
Hym. ...... Hyomandibular. TDM. sacs Tonia marginalis.
TESS ee: Interorbital septum. TV.M. ... Tenia tectum medialis.
Mk. ......... Meckel’s cartilage. Tr. ......... Trabeculee.
6. LrrERATURE.
Auuis, E. P.—The Cranial Anatomy of the Mail-cheeked Fishes.
Zoologica, Stuttgart, Heft 57, 1909.
Boxer, H.—Der Schiidel von Salmo salar. Anat. Hefte, Abt. i.,
Bd. xlix.
Fitarorr, ).—'The Removal and 'Lransplantation of the Auditory
Vesicle of the Embryo of Bufo. Revue Zoologique Russe,
Tome 1, 1916.
Gaupp, E.—Die Entwickelung des Kopfskeletts. Handbuch der
Vergleichenden und Experimentellen Entwickelungslebre
der Wirbeltieren von Oscar Hertwig, Bd. 111.
\ Kinprup, J. E—The Skull of Amiurus. Illinois Biological
Monographs, vol. v. No. 1, 1919.
\ Kinprep, J. E.--The Chondrocranium of Syngnathus fuscus.
Journ. Morph. vol. xxxv. No. 2, 1921.
McMorricu, J. P.—On the Osteology and Development of
Syngnathus peckianus. Q.J.M.S. vol. xxiii. 1883.
Norman, J- R.—Methods and Technique of Reconstruction.
Journ. Roy. Micro. Soc. Pt. 1, 1923.
Parker, W. K.—On the Structure and Development of the Skull
in the Salmon. Phil. rans. Roy. Soc. 1873.
\, Peurson, T.—'The Cranial Development of leleostomian F ishes.
Acta Zoologica, 1922.
Poucurr, F. A.—Le Développement du Squelette des Poissons.
Journ. de l’Anat. et de la Phys. 1878.
\ Sréur, P.— Zur Entwicklungeschichte des Kopfskelets der Teleo-
stier, in Festschrift Feier der 300-jahrigen Bestehens des
Jul.—Max-Universitiits Wurzburg. Leipsig, 1882.
Swinnerron, H. H.— Morphology of the Teleostean Head
Skeleton. Q.J.M.S. 1902.
‘ Unimann, E.—Studien zur Kentniss des Schiidels von Cyclopterus.
Zeitschr. f. Naturwiss. 1921.
Winstow, G. M.—The Chondrocranium of the Ichthyopsida.
Tufts College Studies, vol. 1. 1897.
aia
ee
THE ANATOMY OF THE TONGUES OF THE MAMMALIA. 515:
28. The Comparative Anatomy of the Tongues of the
Mammalia.—IX. HEdentata, Dermoptera, and Insec-
tivora. By Cnuartus F. Sonytac, M.D., FV.Z.8.,
Anatomist to the Society and Demonstrator of
Anatomy, University College.
[Received May 4, 1923: Read May 29, 1923.]
(Text-figures 50-57.)
ConTENTS.
Page
Ordershdentataymsecn-coce cee nooner es eer OLD,
Han eDermopterayrszcletene chsssseaseob i tsiseelsaesesneen| Oem
i ie MNISCCELVOLA pase ay seats cea veessene teabinn aeiseneeacue 523
Wonelusions! 4s -Asecae cee eee Dae
IBibliooraphiya caeenae te eee ee dee seeere | ODT
Order EDENTATA.
The tongues .of the Edentata are characterised by a high
degree of specialisation of their extrinsic muscles, and by a
slight or moderate development of their glands and gustatory
organs. ‘Their mobility is greater than that of all other
Mammalian tongues except those of Zaglossus and Acanthoglossus.
In the Myrmecophagide and Manide the mobility is designed
mainly for the purposes of prehension; but in other Edentates
prehension is combined with a mechanical action of the food.
Form, Apea and Lateral Borders:—The tongue is long,
vermiform and not flattened anteriorly in the Myrmecophagide
(text-fig. 50); it is cylindrical posteriorly and flattened anteriorly
in the Manide (text-fig. 52); it is long, flat and triangular in
the Dasypodide and Orycteropus (text-fig. 52); and it is short,
with the usual Mammalian form in the Bradypodida*.
The apex is rounded in the Bradypodide; it is pointed in
Orycteropus ; and it bears peculiar globular or pointed organs
in the other Edentates. These take the form of globular
expansions of the whole apex in the Myrmecophagide and
Dasypodide, or of a button-like structure attached to the centre
of the apex in the Manide (text-figs. 50 and 52). No trace
of this terminal swelling is present in the Bradypodide and
Orycteropus. I was unable to examine the swellings micro-
scopically to settle their nature, as the tongues are preserved
in the Museum of the Royal College of Surgeons. Some authors
regard them as sense organs, and Mayer (14) described the
swelling in Myrmecophaga as gustatory in function. Perhaps
* See text-fig. 10 in my paper on the anatomy of Bradypus (26).
Proc. Zoou. Soc.—1923, No. XXXIV. 34
516 DR. C. F. SONNTAG ON THE ANATOMY
they are allied to the marginal lobes in Galeopithecus, the
Pinnipedia and Cetacea.
Mayer (14) described two small sharp processes, with their
ends directed forwards and inwards, beneath the apex of the
tongue of Dasypus peba, D. decem-, D. novem-, and D, octocinctus.
They receive prolongations of the lingual musculature. Their
function is probably to capture and grip food, and Mayer
thought they might open and close and be useful for killing
insects.
Text-figure 50.
The tongues of the Myrmecophagide. A: Tamandua tetradactyla; B.C.D: the
front, middle, and back of the dorsum in Myrmecophaga jubata; E: side
view of the tongue in Myrmecophaga showing the sheath (SH.) for the
tongue.
The lateral borders are not distinguishable on those tongues
which are cylindrical throughout (Myrmecophaga, Tamandua,
Cyclothurus). In Manis the flattened anterior part has a
pronounced edge, but the posterior cylindrical part has no
distinct edge. In the Dasypodide and Orycteropus the lateral
borders are pronounced and moderately thick, but the Brady-
podide have borders of considerable vertical depth. Lateral
organs are only found in some specimens of Dasypus.
Sulei and Ridges:—Median dorsal sulci are found in Manis,
OF THE TONGUES OF THE MAMMALIA. 517
Myrmecophaga and Orycteropus. In Manis (text-fig. 52) there
is a wide, shallow groove on the flattened anterior part of the
tongue. In Orycteropus (text-fig. 52) a fine groove runs along
the greater part of the dorsum. And in Myrmecophaga (text-
fig. 50) there is a shallow groove behind the circumvallate papille ;
when it is traced forwards it merges into a central strip of the
tongue in which the mucosa is thrown into transverse folds.
Numerous fine transverse sulci are found in these animals and in
Tamandua and Cyclothurus.
Median ridges are present on the inferior surface of the
tongue in Orycteropus and Zatusia. In the former (text-fig. 52)
there are three ridges and many transverse sulci; in the latter
there is a single ridge.
In the case of Edentates with long, vermiform, protrusible
tongues there are sheaths into which they can be retracted.
In text-fig. 52 is shown a corrugated cuff-like sheath in Manis.
Circunwallate Papille :—The following list summarises the
observations made by myself and those recorded by others :—
Family MyrmMEcoPHAGID#£.
Myrmecophaga jubata: 2 vallate papille (Flower, 8; and
self).
a didactyla: 2 vallate papille (Carus and
Otto, 4).
Tamandua tetradactyla: 2 vallate papille (Mayer, 14;
Miinch, 16)
Cyclothurus : 2 vallate papille.
Family DAsypopips.
Podwisotzky (19) showed how all species have two vallate
papille, and this has been confirmed by observations of many
authors on all known species of Dasypus, Tatusia and Tolypeutes.
The records have been collected by Oppel (17).
Family BrapypopDID&.
Bradypus tridactylus : 2 vallate papille (Cuvier, 6 ;
Sonntag, 25).
oe a : 3 vallate papille (Mayer, 14).
an es : 2 vallate papille (Mayer, 14).
8 cuculliger: 2 vallate papillae (Briicher, 2).
Cholepus didactylus : 2 vallate papille.
Chlamydophorus truncatus: 2 vallate papille (Tuckerman, 21).
Family Manip.
Manis pentadactyla: 3 vallate papille in a_ triangle
(Carus and Otto, 4).
,» tetradactyla: 3 vallatepapilleina triangle (Mayer, 14).
a4 re : 2 vallate papillae (Carus and Otto, 4).
5» javanica: 3 vallate papille in a triangle.
Hs » : 3 vallate papille in a triangle (Oppel).
34*
518 DR. O. F. SONNTAG ON THE ANATOMY
‘Family OnycrEropopip&.
Orycteropus capensis: 3 vallate papilla in a triangle.
53 » + 3 vallate papille (Rapp, 20).
It is thus evident that with very few exceptions the American
Edentates have a pair of circumvallate papilla, whereas the
African forms have three papille disposed in a triangle with the
apex directed backwards.
The papille are frequently found retracted within a deep fossa,
but those in some of the Dasypodide stand up very prominently.
In the specimen of Zatusia shown in text-fig. 51 they are very
prominent, and it is at first sight difficult to distinguish them
from the larger fungiform papille on the posterior part of the
oral division of the dorsum.
The vallate papilla are close to the epiglottis in the Brady-
podidee, Dasypodidze, Myrmecophagide and Orycteropus, but they
lie far from it in Manis, thus making the basal part of the
tongue long.
In Tamandua (text-fig. 50) there are two small pin-hole-like
pits in front of the vallate papille, but I was unable to make a
histological examination to ascertain whether or not they receive
the secretions of glands,
The vallate papille are circular on plan, and cylindrical or
conical on elevation, with the apex of the cone attached to the
tongue. The surface is smooth or granular, and a histological
examination shows that the granular effect is produced by
several secondary papille. The fossa is well marked, and the
vallum frequently overlaps the edge of the papillae. Oppel (17)
has published an illustration of a section through a papilla in
Manis javanica, showing how the long axes of the taste-buds
run downwards and outwards towards the fossa which passes
under the base of the cone-like papilla.
Tuckerman (21) pointed out that the papille in Dasypus peha
resemble those of the higher Mammalia, but those of Chlamy-
dophorus truncatus are very similar to those of the Marsupialia ;
their resemblanee to the anterior papille in Belideus and
Phalangista is very marked indeed.
Fungiform Papille :—I did not observe any trace of fungiform
papilla in Ayrmecophaga, Tamandua, Cyclothurus or Mais, nor
has any other anatomist described them. ‘They are present in
all other Edentata, but they are never very numerous. No
apical cluster exists, but they have the usual arrangement in rows
of varying degrees of obliquity behind that in the Dasypodide ;
and they stretch right across the dorsuin (text-fig. 51), In the
Bradypodidee * the apical] cluster varies in size, but the papille
do not cross the middle line. In Orycteropus there is no apical
cluster, and the papillae only form a dorsal bounding zone.
Tuckerman (21) points out that some of the papille are sunk in
* See foot-note on page 615.
OF THE TONGUES OF THE MAMMALIA, 519
little hollows in Dasypus peba; and those lying on the posterior
part of the tongue have well-developed taste-buds. No papills
fungiformes are present on the lateral borders.
Lateral Organs :—There appears to be considerable variation
in the lateral organs, for the views of various authors are
conflicting. I observed no trace of organs in Myrmecophaga
jubata, Tamandua tetradactyla, Cyclothurus didactylus, Manis
(sp. %), Orycteropus capensis, Bradypus tridactylus, Z'atusia (sp. *)
and Dasypus villosus. Mayer (14) observed them in J/yrmeco-
phaga and the Dasypodide. Boulart (1) and Tuckerman (21)
Text-figure 52.
Text-figure 51.
The tongue of Tatusia.
ay
The tongues of the Mande. A.B: front and back
of the tongue of Manis; C: under surface of
the apex of the tongue of Manis; D: the
tongue of Orycteropus capensis.
found them in Zatusia, and the latter author found them taking
the form of minute openings in several Armadillos; taste-buds
and serous glands were connected to them. Gmelin (9) found
no trace of organs in Orycteropus, the Armadillos and Sloths, and
Podwisotzky (19) saw them in Dasypus seacinctus. So it is
evident that the organs are variable in the Dasypodidex, but
usually absent in the other Edentates.
Conical Papille:—These papille are well-marked in the
Bradypodide, Dasypodidm and Orycteropus, but I found no trace
of them on naked-eye examination in the Myrmecophagide or
520 DR, C, F. SONNTAG ON THE ANATOMY
Manis. In the species in which I observed them they have the
usual arrangement in an apical cluster, and in rows of varying
degrees of obliquity behind that. They are also present on the
lateral borders, but they are restricted to a narrow bounding
zone on the inferior surface of the tongue. They stretch back
to the neighbourhood of the epiglottis, but those on the base of
the tongue are not very large. In Orycteropus (text-fig. 52) the
individual papille are easily distinguished ; in the Brady podidee
they can only be detected through a lens; and in the Dasypodide
they vary in prominence to such a degree that the dorsum
appears granular (Dasypus) or almost shaggy (Tatusia). Some of
the types of conical papille in the Dasypodide and Brady podide
are shown in text-fig. 53.
Oppel (17) described scanty conical papille in Manis, and
Tuckerman (21) stated that those in Dasypus villosus are inter-
mediate in character between the coronate and fasciculate
papille in Marsupials and the mechanical papille in higher
Mammalia. Secondary papille are found on many of the
main ones.
Text-figure 53.
ee We Sa
WW tYoen
The papille in the Edentata, a-c: conical papille ; d-: cireumvallate
papilla: cut vertically ; i-4: fungiform papilla.
Glands :—Oppel (17) and Podwisotzky (19) mapped out the
serous and mucous glands of the tongue in Alanis javanica.
They showed that the mucous glands form a cluster on the base
of the tongue, whilst the patch of serous glands lies far forwards
round the vallate papilla. Owing to the large size of the
salivary glands the mucous glands are not very numerous. I did
not observe any patulous glandular orifices in the Myrmeco-
phagide or Orycteropodide. In the specimen of A/anis at my
disposal the base of the tongue was concealed to » considerable
extent by the sheath. But in Bradypus I observed a number of
orifices and lymphoid nodules. No apical gland of Nuhn is
present.
The serous glands vary in the Dasypodide ; they are numerous
in Dasypus villosus, but scanty in D. peba.
Lytta:—The only Edentates which possess a lytta are Manis
gigas and M, javanica, It has been described and figured by
Oppel (17) and Rapp (20); and Mayer (14) peints out that it is
OF THE TONGUES OF THE MAMMALTA. 521
large aS in many Carnivora. I was unable to cut into the
tongues of Manis, Myrmecophaga or Orycteropus, but I found no
trace of it in Zamandua, Bradypus, Dasypus or Tatusia.
The frenwm is short in all Edentates.
There is no trace of plice fimbriate or frenal lamella. The
musculature and elastic tissue have already been fully described
by Owen (18) and others. No intermolar eminence is present.
Sir William Flower (27) came to the following conclusions as
regards the affinities of the Edentates:—“ All the American
Hdentates at present known, however diversified in form and
habits, belong to a common stock. The Bradypodide, Mega-
theriide, and Myrmecophagide are closely allied, the modifications
seen in existing families relating to food and manner of life.
The ancestral forms may have been omnivorous, like the present
Armadillos, and gradually separated into the purely vegetable
and purely animal feeders; from the former are developed the
modern Sloths, from the latter the Anteaters. The Armadillos
are another modification of the same type, retaining some more
generalised features, as those of the alimentary organs, but in other
respects, as their defensive armature, remarkably specialised.
“The two Old-World forms Manide and Orycteropide are so
essentially distinct from all the American families, that it may
even be considered doubtful whether they are derived from the
same primary branch of mammals, or whether they may not be
offsets from some other branch, the remaining members of which
have been lost to knowledge.”
It was shown above that the tongues of the American
Edentates have two circumvallate papille, whereas those of the
Old-World forms have three; and it is only in rare specimens
that this arrangement is departed from.
If the Armadillos were like the ancestral forms, as Flower
believed them to be, it is necessary to see whether their tongues
could have been transformed into those of the Myrmecophagide
and Bradypodide. The tongue of any Armadillo is long and
triangular, with a slightly bulbous apex and a good supply of
fungiform and conical papille; and in some species there are
lateral organs. If the tongue be simply shortened, with loss by
absorption of the apical swelling, it would become indistinguish-
able from that of Bradypus. Conversely, enormous elongation
of the tongue, with retention and specialisation of the apical
swelling, produces a tongue like that of Ziamandua or Myrmeco-
phaga; and as the tongue in these insectivorous animals is merely
a glutinous finger, the conical and tungiform papille on the
dorsum would undergo disuse atrophy. So it is evident that the
mere changes in form, with or without disuse atrophy, would
convert the tongue of an Armadillo into that of a Sloth or an
Anteater.
The tongue of Manis resembles those of the Myrmecophagide
in its general form, and in the specialisation of the apex to form
a sensory organ; but the likeness between them stops there.
522 DR. OC. F, SONNTAG ON THE ANASOMY
These resemblances are the result of convergence and are,
consequently, no test of affinity. There is, moreover, no resem-
blance between the tongues of Manis and Orycteropus, beyond
the characters of the circumvallate papillae. If they have arisen
from a common form, the great contrast between them is the
result of adaptation to a particular diet and mode of feeding. And
there appears to be no affinity between the tongues of the African
and American Edentates. So it is evident that the characters of
the tongues lend additional support to Flower’s views on the
mutual affinities of the Edentata.
Order DERMOPTERA.
The tongue of Galeopithecus volans (text-fig. 54) is long,
narrow and thick, and its free part is of considerable extent.
The upper surface is chocolate-coloured, but the lower surface is
pale. The apex is rounded, devoid of a notch, and divided into
lobules by fissures, which pass backwards and inwards for a
considerable distance on the inferior surface. The lateral borders
are full and rounded, and have well-marked lateral organs at
their posterior extremities. A well-marked median dorsal sulcus
runs back along the greater part of the oral division of the
Text-figure 5A,
The tongue of Galeopithecus volans. A: dorsum; B: under surface;
C: vallate papillz of a second specimen.
dorsum, but no transverse fissures diverge from it. ‘Che median
ventral sulcus is equally well-marked, and transverse sulci diverge
from its posterior part. There is no trace of a foramen cecum
or plice fimbriate, and the frenum is very short. No frenal
lamella is present.
Papille:—The whole of the dorsum from the apex back to the
epiglottis is covered with thickly-set, sharp, hard conical papilla
which have the usual arrangement in clusters and rows; and
they increase in size from within outwards, and from before
OF THE TONGUES OF THE MAMMALIA. 523
backwards. They also cover the upper part of the lateral borders,
but they are not present on the inferior surface of the tongue.
They are of a simple character. In one tongue I was unable to
detect any circumvallate or fungiform papille; but these were
probably buried under the conical papillae. In a second tongue
the conical papillze were short, the fungiform papille were scanty,
but had the usual arrangement; and there were two large
vallate papille, with a minute one in between.
Lateral Organs :—Owing to the great length of the oral part of
the tongue, the lateral organs appear to be placed far back on the
lateral borders. They are as well marked as those in Primates :
and each consists of eleven laminz separated by deep sulci.
Sublingua :—Gregory (10) pointed out the Galeopithecus has a
sublingua similar to that in Z’upaia, but I was unable to detect
either a sublingua or plice fimbriate in three tongues.
No lytta is present, and no gland orifices are visible on the
base of the tongue.
Order INSECTIVORA.
The tongue, like other anatomical characters, is very simple in
the Insectivora, and it is possible to show how the tongues of
animals belonging to other Orders can be derived from them by
specialisation of certain structures or areas. In the present
paper the structural characters alone are dealt with, the phylo-
genetic considerations being postponed till all the systematic
papers of this series have been completed.
The tongue is long and narrow in all species, and its free part
is of considerable length. It usually thickens gradually from
before backwards; but its thickness is not greater in proportion
to its length in Lrinaceus (text-fig. 55). The oral part of the
dorsum is long, and the pharyngeal part is short in all species.
The apex is rounded, truncated or pointed, and is usually devoid
of a notch. The lateral borders are rounded, but they only
possess lateral organs in Hrinaceus. .
Median dorsal sulci are absent, but transverse ridges and sulci
may be produced by the impression of the palatal rugw on the
tongue. Median ventral sulci are absent, but median ventral
crests are present in Z'alpa (text-fig. 56) and Rhynchocyon (text-
fig. 57); they are blunt in the former, and sharp in the latter.
Circumvallate Papille:—In the Insectivora there are two
large papillae, or three papille in a triangle with the apex turned
backwards. And the following list contains the observations of
myself and others :—_ a
Family Erinaceide :—Two papille in Gymnura rafflesii.
| Three papille in all species of Hrinaceus.
Family Centetidee:—Three papille in Centetes, Hriculus and
Microgale.
Family Solenodontidz :—-Three .papille in Solenodon para-
doxurus.
524 DR. O. F. SONNTAG ON THE ANATOMY
Family Potamogalids :—Three papillse in Potamogale velox.
Family Chrysochloride :—Three papille in all species of
Chrysochloris.
Family Talpide:—Two papille in all species of Zalpa, Myogale,
Condylura, Scapunus and Scalops.
Family Soricidz :—Two papille in all species of Sores.
Three papille in Rhynchocyon.
The papilla are round or oval on plan, and cylindrical or
conical on elevation, The surface is smooth or granular, and
there may be a small central depression. The fossa and vallum
are well-marked. Carlier (3) showed that the papille in Hrinaceus
europeus contain serous glands and ganglion cells, and their
connective-tissue cores are very cellular. The ducts of these
serous glands open along with those of the serous glands within
- Text-figure 55.
B
The tongue of Hrinaceus ewropeus. A: dorsum; B: under surface;
C-E:; vallate papille.
the base of the tongue, The only other Mammals, so far as I am
aware, in which there are glands within the vallate papille are
the Chiroptera.
Lungiform Papille :—These papille are never very numerous,
but more are present in Hrinaceus and Chrysochloris (text-fig. 55)
than in any other genus. It may be difficult to detect them, for
they are usually very small. Dobson (7) points out, however,
that they are prominent in Gymnura rafflesti and Chrysochloris
villosa, They may be composed entirely of epithelial cells, as in
EHrinaceus, or they consist of both epithelial and connective-tissue
elements.
Conical Papille:—In all species the conical papillw are
numerous and closely packed. They are usually very small on
the oral part of the dorsum; and they are usually small on the
OF THE TONGUES OF THE MAMMALIA. 525
base as well. The basal papille are large in Centetes and Chryso-
chloris.
When examined through a hand lens they are seen to be
simple in character, having one, two or three points. In Chyso-
chloris scoop-shaped forms are present. In many species they are
composed entirely of epithelium, which is strongly cornified, the
component cells being formed of the strata from the surface down
to the Malpighian layer. The different forms are met with in
genera of the same family; thus Gymmnura has bifid papille, but
Erinaceus has them divided into three points. So it is evident:
that the forms are simpler than in animals belonging to all the
Orders hitherto considered in this series of papers.
Text-figure 56.
The tongue of Talpa europea. A: dorsum; B: under surface;
C: cross-section showing the oval lytta.
Lateral Organs:—Many authors have described the lateral
organs in Erinaceus, and some have stated that these structures
are absent in all other genera. With both these remarks I am
thoroughly in accord.
In Hrinaceus europeus, according to Mayer (14) and Gmelin (9),
the organ consists of two large folds well provided with taste-buds
and fat. In the animal examined by myself (text-fig. 55) the
organ varied on both sides, On the right side a fissure, with
slightly curved ends, much as in Wydrocharus capybarw; on the
526 DR. C. F. SONNTAG ON THE ANATOMY
left side the fissure undercut part of the dorsum, thereby
producing an overhanging lobe.
Glands :—I agree with Carlier (3), Oppel (17) and Podwisotzky
(19) that the serous glands are very highly developed, and the
muvous glands ‘are scanty in Hrinaceus. As was mentioned
above, serous glands are present even within the vallate papillee in
EL. europeus. No apical gland of Nuhn exists in any Insectivore.
The frenwm linguwe is short in all Insectivora except Z'upaca.
And it is tine in Hrinaceus, but thick in Talpa, Rhynchocyon, etc.
In no case is there any trace of frenal lamellze as in the Lemuroid
Primates.
Sublingua:—It has been shown by Owen (18) and Garrod that
there is a sublingua in Z'upaia; Owen described it as a fimbriate
plica, but Garrod described it as resembling that in Chiromys.
Vogt and Yung (24) recorded the presence of a sublingua in
Text-figure 57.
The tongues of Chrysochloris trevelyant (A and B) and Rhynchocyon (C and D).
In both cases a third papilla is concealed in the linear groove on the back
of the dorsum.
some Insectivora. I found no trace of the sublingua in EHrinaceus,
Centetes and Chrysochloris, but it may be represented by the
median ventral ridge in Talpaand Rhynchocyon. That ridge may
correspond to the median crest which is found on the ventral
surface of the sublingua in the Marsupialia and Primates. In
that case the remainder of this organ has been absorbed into the
tongue.
Lytta:—It has been shown by Nussbaum (28), Oppel (17),
Hesse (11), and Ludwig Ferdinand, Prince of Bavaria (13), that
the lytta is well-developed in Hrinaceus europwus, Sorex fodiens
and Z'ulpa europea. It is composed of muscle fibres, fat and
fibrillar connective tissue, but no cartilaginous elements are
present. The histological details are given fully in the papers
enumerated above. I observed a very pronounced lytta in
OF THE TONGUES OF THE MAMMALIA. 527
Khynchocyon. The lytta in the Insectivora is in many ways
similar to that in the Lemuroid Primates.
Musculature :—This has been fully described by Carlier (3) and
Oppel (17).
CONCLUSIONS.
1. The tongucs of the Edentata and Insectivora are of particular
interest, for they afford clear illustrations of some of the principles
which should guide us in settling the blood-relationships of
animals. ‘Thus they exhibit primitive, convergent and adaptive
characters.
2. The tongues of all the American Edentata can be derived
from Armadillo-like forms.
3. The tongues of the Myrmecophagide are similar in many ~
ways to that of Manis, but the resemblances have been produced
by convergence.
4, All American Edentata have two circumvallate papilla, but
the African forms have three in a triangle. My
5. The differences between the tongues of Manis and Oryc-
teropus are the result of adaptive modifications. |
6. It is a well-established principle of phylogeny that obser-
vations should be drawn from all parts of the body; and
conclusions as to aflinities should not be based on the examination
of asingle organ. The truth of this can be seen in a study of the
Kdentata; for an examination of the tongue alone would make
one agree that JMJanis and the Myrmecophagidw are closely
related ; an examination of the entire anatomy of these animals
shows that that is not true. °
7. The tongues of the Edentata are more mobile than those of
all other Mammals except Acanthoglossus and Zaglossus.
8. The tongue of Galeopithecus has affinities with those of the
Insectivora. The characters of its apex, however, are unique.
9. The tongues of the Insectivora are very primitive, for their
mechanical papille are simple, their gustatory papille are not
numerous and they have traces or complete examples of the lytta
and sublingua. Many of their characters are similar to these of
the Lemuroid Primates.
BIBLIOGRAPHY.
a Bourarr and Piturer.—Note sur l’organe folié de la langue
des Mammiféres. Journ. de l’anat. et de la physiol.
1885, pp. 337-345.
2. Briicurr, C.—Deutsch. Zeitschr. f. Tiermed. u. vergl. Pathol.
. (Inaug.-Diss. Tiibingen), 1884.
3. Cartier, EH. W.—Contributions to the histology of the
Hedgehog. Journal of Anatomy and Physiology, 1893
(several papers). Ae ae
528
16.
17.
DR. C. F. SONNTAG ON THE ANATOMY
. Carus and Orro.—Erliuterungstaf. zur vergl. Anatomie,
vol. iv. Leipzig, 1835.
. Cuatin, J.—Les organes des sens dans la série animale.
Legons d’anat., ete. Paris, 1880.
. Cuvier, G.—Lecons d’anatomie comparée. Paris, 1835.
Trans. by Duvernoy, Stuttgart, 1837.
. Doxsson, G. E.—A Monograph of the Insectivora, Systematic
and Anatomical, London, 1882-1890.
. Firowrr, W. H.—Lectures on the comparative anatomy of
the Mammalia. Medical Times and Gazette, 1872
(several papers).
GmeE.in, F.—Zur Morphologie der Papilla vallata und foliata.
Arch. f. mikr., Anat. 1892, pp. 1-28.
. Grecory, W. K.—The Orders of Mammals. Bull. Amer.
Mus. Nat. Hist. xxvii. 1910.
. Hfesszr, F.—Uber die Muskeln der menschlichen Zunge.
Zeitschr. f. Anat. u. Entw. 1876, pp. 80-106.
. Honiascumrep, J.—Kleine Beitrige zur Verteilung der
Geschmacksknospen. Zeitschr. f. wiss. Zool. 1877,
_ pp. 255-262,
. Lupwic Ferpinanp.—Zur Anatomie der Zunge. Eine
vergleichend-anatomische Studie. Anstalt, 1884.
. Maver, F. J. ©.—Uber die Zunge der Vermilinguia.
Froriep’s Notizen, 1842, pp. 289-290.
. Mayrr, F, J. C.—Uber die Zunge als Geschmacksorgan.
Nov. Act. Acad. Leop.-Carol. Nat. Cur. 1844, pp. 721-
748.
Miincu, F.—Die Topographie der Papillen der Zunge des
“Menschen und der Siugethiere. Morphol. Arbeit.
1896, pp. 605-690.
Opret, A.—Lehrbueh der vergleichenden Mikroskopischen
Anatomie, 1900, vol. ii.
. Owen, R.—On the Anatomy of Vertebrates, 1868, vol. i1i.—
Mammals.
. Popwisorzky, V.—Anat. Untersuch. tber die Zungendriisen.
Diss. Dorpat, 1878.
. Rarp, W. yv.—Anatomische Untersuchungen uber die
Edentaten. ‘Tubingen, 1843.
. Tuckerman, F.—On the Gustatory Organs of some Hdentata.
Internat. Monatsschr. f. Anat. u. Physiol. 1890,
pp. 335-339.
. Tuckerman, F.—On the Gustatory Organs of some of the
Mammalia. Proc. Boston Soc. Nat. Hist. 1890, p. 470.
23.
24.
25.
26.
27.
28.
OF THE TONGUES OF THE MAMMALIA. 529
Tuckerman, F.—On the Gustatory Organs of some of the
Mammalia. Journal of Morphology, 1890, pp. 152-193.
Voer and Yune.—Lehrbuch der praktischen vergleichenden
Anatomie, vol. ii. 1894.
Wyss, H. v.—Die becherférmigen Organe der Zunge.
Arch. f. mikr. Anat. 1870, pp. 237-258.
Sonnraa, C. F.—Contributions to the Anatomy of Bradypus
tridactylus. P.Z.S. 1921, pp. 157-177.
Fuowsr, W. H.—On the Mutual Affinities of the Animals
comprising the Order Edentata. P.Z.S. 1882, pp. 358-
367.
Nusspaum, M.—SB. d. Niederrhein. Ges. f. Nat. u. Heilk.,
Bonn, 1881, pp. 181-183.
EXTERNAL CHARACTERS OF THE PIGMY HIPPOPOTAMUS, 531
29. The xternal Characters of the Pigmy Hippopotamus
(Cheropsis liberiensis) and of the Suide and Camelide.
By R. I. Pocoox, F.R.8., F.Z.S8.
{Received May 28, 1923: Read June 12, 1923.]
(Text-figures 30 to 46.)
ConTENTS.
Page
The Muzzle and Ears of Cheropsis liberiensis.............. 632
The Feet of Choropsis liheriensis .......00...s00ceceeeseneeeese 633
The Tail and Penis of Cheropsis liberiensis.................. 636
The Rhinarium of the Suidse ...........0..0.eccecee eee ceeeeeeee 635
Nhe MacialVibrissag ces ee tare cess sone oies Redes eee ee DSS
Mem Ma Ie ie vceccons coset eae oe tate leet eatetek eaen eine ae eat SO)
Mer Beet ss ceaes shkence ieee ehacncis oa emma eee san aetea ec ecneet a DAO)
The Muzzle in the Camelida ..............0..cccccce cee eeeeeeee 642
The Occipital Gland of Camelus ..........00c.ccccceceeeseeeeeee. 644,
The Feet and Metatarsal Gland of Lama ..............0...... 646
The pWect\ of Camelusney ccaccccciscsen cock Moianinnt eee BAT
The Penis of Lama and Camelus.......0...ccccsceceeceverseeeeee B48
The fresh material upon which the observations contained in
this paper are based was examined in the Society’s Prosectorium
immediately after the death of the specimens. In the case of
some of the Suide, like Porcula and Hylocherus, I have been
compelled to rely upon dried skins in the British Museum.
The only rare species examined was the Pigmy Hippopotamus,
of which a single old male example, presented by the Duke of
Bedford on December 16, 1913, died February 11, 1919.
Section SUINA.
Family HiPPOPOTAMIDA,
CuGROPSIs LIBERIENSIS.
Owing to its smaller head, much shorter body, and relatively
longer legs, the Pigmy Hippopotamus differs markedly in
external appearance from its larger ally. It was generically
Proc. Zoot. Soc.—1923, No. XX XV. 35
532 : MR. R. I. POCOCK ON THE EXTERNAL
separated from the latter, owing to the loss of a pair of incisor
teeth in the lower jaw.
The®muzele is provided with short scattered bristles, but, apart
Text-figure 30.
Hind foot of Cheeropsis liberiensis trom above.
‘Text-figure 31.
. The same from below.
from these, the facial vibrisse characteristic of the normal terres-
trial mammalia appear to be unrepresented. The nostrils are
widely separated, oblique, valvular slits, capable of being tightly
CHARACTERS OF THE PIGMY HIPPOPOTAMUS. 533
closed, and look forwards and upwards from the summit of the
muzzle. There is no differentiated rhinarium.
The ears are small, simple, and, like the nostrils, capable of
being tightly closed. The hollow of the outer side is furnished
with three soft, ridge-like thickenings which, when the ear is
folded, ave pressed together so as to block the orifice. (Text-
fig. 33, A.)
Text-figure 32.
A. Right fore foot of Cheropsis liberiensis from below.
B&C. The same foot from the outer and inner sides.
The feet are of the most primitive type found in the Artio-
dactyla. They are symmetrical or nearly so; the tips of the
four toes rest on the ground, and with the help of a well-
developed plantar pad support the weight of the animal when
walking or standing.
The two main digits, morphologically the third and fourth,
ave united for the greater part of their length by integument,
but they are not tightly and compactly welded together. The
35*
534 MR. R. I, POCOCK ON THE EXTERNAL
depression between them on the upper side is tolerably deep ;
they can be separated to a considerable extent, and their distal
ends carrying the hoofs are free. The hoofs are small and
bluntly pointed, and there is no clear line of demarcation
laterally between their edges and the area of flat horny integu-
ment constituting the sole of the toe.
The lateral digits are much smaller, are free from webbing, and
the small hoofs encircle their tips.
The sole of the foot is covered with naked wrinkled skin. On
the area between the plantar pad and the edge of the webbing
uniting the two larger digits there is a shallow depression which
may be glandular, but I was unable to investigate its nature by
sections, ‘lhe plantar pad is irregularly heart-shaped, being
narrow and more or less truncated in front, where it stands up as
Text-figure 33.
A. Kar of Cheropsis liberiensis.
B &C. Dorsal and lateral views of the tail of the same.
a well-defined horny cushion, and widely rounded behind, where
it is ill-defined and gradually blends with the integument above
or behind it. (Text-figs. 30-32.)
There is no marked difference in size or structure between
the fore and hind foot, except that in the latter the two
smaller digits are alike in size, whereas in the former the
outer digit is considerably thicker and larger than the inner.
This is the only particular in which there is distinct asym-
metry in the feet.
The feet of living examples of Hippopotamus amphibius
that I have seen differ from those of Cheropsis liberiensis in
being much more compactly built. There is no great difference
in size between the four digits, and since they are much more
closely united by integument, the feet are incapable of the
CHARACTERS OF THE SUIDA. 535
expansion observable in Chwropsis. In the greater freedom of
the digits the latter genus stands nearer to the Suide *
The tail is short. Its base is broad with convex edges, but
its distal half is thin, parallel-sided, strongly compressed, and
provided with stiff short bristles. It serves apparently merely
as a cover to the anus, and in the female of the genital orifice as
well. (Iext-fig. 33, B, C.)
The penis is “quite simple, cylindrical for the greater part of its
length, and somewhat abruptly attenuated at the apex, the
orifice being terminal. The penis is like that of Ss scrofa, but
has no spiral twist when retracted.
Family Surpa#.
The Rhinariun.
The general character of the rhinarium in the Suide is well
known. It is a movable disk, the upper and lateral edges of
which project to a greater or less extent beyond the skin of the
muzzle. Its anterior moist surface is nearly flat, its upper
portion being nearly naked, and the lower covered with
short, stiff, sparse hairs. ‘The area below the nostrils is usually
wider than the area above them, and the edges are, generally
speaking, convex, although the lower is less curved than the
upper. The nostrils look straight forwards, and are set on each
side of the middle line of the disk some distance apart. They
do not extend to its lateral edge, the lateral narial slit being
absent or so short as to be practically negligible. Their inner and
upper edges are sharply defined, but the outer blends gradually
with the adjacent surface of the rhinarium.
In most of the genera of the family— Sas, Potumocherus, and
Dicotyles} (Tayassu and Pecari)—the variation in shape and
structure is comparatively slight, and the differences shown in
the sketches may be individual and not specific or generic. In
all these cases the median height is less than the gr catest width
hy about one-fourth or less, and the sides from nites widest points
. little below the nostrils are inclined inwards and upwards in
a nearly straight line to the curved summit, and the distance
between the nostrils is more than one-third but less than one-half
the median height.
In an example of Sus scrofa the widest part is just below the
inferior edge of the nostrils, and the infero-lateral margin is
rather markedly convex, and the sides from the level of the
* In the mounted specimen of Hippopotamus amphibius in the Natural History
Museum, the characters of the feet here mentioned are not so apparent. The
approximate equality between the four digits is seen, but there is a deep depression
between the second and third, making the feet look more like those of Cheeropsis.
This, however, is probably due to the shrinkage of the interdigital tissue with
drying.
+ For convenience I retain in this paper the old familiar name Dicotyles for the
American Suide.
536 MR. R. 1. POCOCK ON THE EXTERNAL
nostrils slope upwards and inwards with a slightly sinuous curve
to the narrowed, rounded summit. In S. leucomystaax the
greatest width is about midway between the nostrils and the
lower edge, which is straighter than in S. scrofa, and the height,
as compared with the width, is considerably greater than in the
latter. (‘lext-fig. 34.)
In Potamocherus porcus the upper edge is much more widely
rounded than in the two species of Sus, and the infero-lateral
margins at the widest part of the rhinarium below the level of
the nostrils is not so widely rounded, the edge from that point
inclining inwards and upwards, being very lightly concave.
(Text-fig. 35, B.)
Text-figure 34,
A, B. Front and side views of rhinarium of Sus leucomystax.
C, D. The same of Sus scrofa.
In two species of Peccary, the collared (D. tajacu=torquatus)
and white-lipped (D. pecari=labiatus), the rhinarium is as wide
on a level with the, upper edge of the nostril as below that
point, which is not the case in Sus or Potamocharus. (‘Text-
fig. 35, C.)
In Phacocherus the vhinarium differs from that of the genera
described above in several particulars. It is about twice as wide
as high, has a lightly convex upper edge, the lateral margins
strongly convex round the nostrils, and sinuously concave below
where they form a definite angle with the lower edge, which is
lightly concave, especially in the middle line. (Text fig. 35 A.)
It is not possible to determine the exact shape of the rhinaria
CHARACTERS OF THE SUID. 537
on dried skins; but so far as I can judge from material in the
Natural History Museum, the rhinarium of Hylocherus is rela-
tively much larger than in Potamocherus, and approaches in
relative width that of Phacocherus.
Porcula, as might be expected, appears to resemble Sus in the
shape of this organ.
Text-figure 30.
A. Front view of rhinarium of Phacocherus africanus.
B. The same of Potamocherus porcus.
C. The same of Dicotyles pecari.
The following table of measurements in mm. will show the
principal distinctive features of the rhinarium of Phacocherus as
compared with that of the other genera :—
Height. ] Width. | Distanen between
JSUSSCHOI Cisse cee oe eiitnce Stas OSen eg SO 23
Sus leucomystad ............0.. 55) 68 18
Potamocherus porcus ......... DI 163 19
Dicotyles pecari (labiatus) ..... 43 | 57 | 18
Phacocherus africanus ...... enaaer ren eae) | 53
538 MR, R. I. POCOCK ON THE EXTERNAL
The Facial Vibrisse.
The full complement of facial vibrissee characteristic of the
Mammalia can sometimes be detected in the Suidx, but I have
failed frequently to trace the genal tufts, owing to their being
either suppressed or indistinguishable in the thick clothing of
coarse hair which generally covers the cheek. The mystacials
Text-figure 36.
gO ee x Ss :
‘eae
A. Side view of head of Potamocherus porcus showing facial vibrisse.
B. The same of Dicotyles pecari.
C. Base of the ear of P. porcus.
are relatively short and scattered, and apparently of less import-
ance than the others, probably on account of the prominence and
sensitiveness of the specialized rhinarium. The lower lip is
always provided with scattered submentals approximately as long
as the mystacials; but the superciliaries and suboculars are
always long and plentiful as if the protection of the eye was of
importance. ‘Typically the interramal tuft is composed of about
half a dozen moderately long vibrisse.
CHARACTERS OF THE SUID. 539
In the two species of Peccary (D. tajacu and pecari) I have
found all the tufts developed as described above, and, in addition,
the two genal tufts, the lower situated in a line with the slit of
the mouth and beneath the anterior angle of the eye, the upper
higher up and beneath and behind the posterior angle of the eye.
(Text-fig. 36, B.)
In Potamocherus porcus the ocular and interramal vibrisse are
well developed, and I detected two genal tufts, the upper situated
as in Dicotyles, the lower set very far back beneath the base of
the ear on the angle of the lower jaw. The upper tuft, however,
was only distinguishable from the normal coarse hairs of the
cheek by rising from a low integumental swelling, and the
homology of the lower tuft with that of Dicotyles is rendered a
little doubtful by its abnormally backward position. ('Text-
fig. 36, A.)
In dried skins of Porcula salvania the buccal, ocular, and
interramal vibrissee are normally developed, but the genals are
indistinguishable.
In Phacocherus the buccal and ocular vibrisse alone seem to
be constant. In one ease I detected a few bristles set about
one inch below the subocular wart, which I believe to represent
the upper genal tuft; but I could find no trace of the lower genal
tuft, unless a short row of black bristles beneath the white fringe
on the cheek is to be referred to it. The interramal is repre-
sented at most by one or two long bristles.
In dried skins of Hylocherus the buccal and ocular vibrissze
are as in Phacocherus, but the genals are not traceable in the
long hairs of the cheek. I may here add that the fringe of pale
hair on the cheek in Hylocherus seems to overlie an area of skin
with special glandular activity.
The Kar.
Boas (Die Ohrknorpel . . . der Saugethiere, 1912) describes and
figures the ears of Sus scrofa and Dicotyles tajacu. Hach is
provided with three supporting ridges, the anterior of which is
curved backwards at the base, where it arches over a deep groove
bounded below by an oblique thickening or ridge, which is itself
defined by a deep groove from the basal thickening which descends
to the auditory orifice. The anterior edge of the pinna is turned
backwards in both genera, and the posterior edge as well in
Dicotyles. At the base of the anterior edge there is a small
excrescence.
In Potamocherus porcus the auterior edge is folded backwards
at the base, but there are no definite tragal or antitragal
thickenings. There is a feebly-developed anterior and posterior
supporting ridge, and the former rises inferiorly close to the
basal thickening, which is divided into an upper and a lower
part by a deep groove or cleft. The part above this groove forms
an abbreviated longitudinal thickened ridge, well defined in
540 MR, R. I, POCOCK ON THE EXTRRNAL
front, but blending behind with the integument of the ear. It
suggestively resembles the supratragus (plicaw principalis) of the
ear of typical mammals, the retention of which in the Suide
would be of interest, but Boas does not identify it with that
ridge. The part below the groove descends into the tubular
part of the ear, and is pierced inferiorly by the auditory orifice.
Its upper edge, just beneath the supratragal ridge, is raised
posteriorly into a rounded excrescence. (Text-fig. 36, C.)
The ears vary in size and shape according to the genera. In Sus
scrofa and Potamocherus porecus the posterior edge is lightly con-
cave above and widely rounded below, the apex being more elon-
gated in the latter than in the former genus. In Phacocherus the
posterior border is not widely rounded inferiorly, but is emarginate,
and above the emargination there is an angular lappet.
The Feet.
The feet of the Suidee differ from those of the Hippopotamide
in that progression is bidigital, the weight ‘of the body resting
Text-figure Bie
A. Lower view of foot of Sus scrofa.
B. Section of foot of the same.
upon the hoofs of the second and third digits, those of the first
and fourth at most touching the ground with their tips. There
isno median plantar pad, but the third and fourth digits are
CHARACTERS OF THE SUIDA- 541
provided beneath with two horny pads behind the nail of the
hoofs, constituting a pair of heels. ‘Che nails of these digits are
narrowed apically, flattened on the inner side, and fit together
like those of typical ruminant Artiodactyles.
Except for the well-known variation in the case of Dicotyles,
where the fourth or outer digit of the hind foot is absent, the
inner being retained and used for scratching, the feet of the
different genera and species of Pigs are very much alike. The
interdigital depression is. shallow, sparsely hairy, and not glan-
dular, and the back of the pasterns and fetlock to a point
just above the lateral hoofs is usually naked, although in
Text-figure 38.
A. Lower view of left hind foot of Dicotyles.
B. The same from the side.
C. Lower view of foot of Phacocherus, showing the fusion of the heels.
an example of Sus scrofa there was a median line of hair extend-
ing almost down to the heels. The foot of Phacocherus differs,
however, from that of Dicotyles, Sus, Porcula, and Potamocherus
in having the heels united; but the degree of fusion is, I think,
variable, although in some cases it is very marked and practically
complete. Judging from dried skins, Zylocherus has Seerare’
heels like Potamocharus. (Text-figs. 37, 38.)
I have found no specialized glands in the feet either of
Dicotyles or Phacocherus, and with regard to the incidence of
the glands that have been recorded in other genera, further
information is required before definite conclusions can be reached.
S
542 MR. R. I, POCOOK ON THE EXTERNAL
‘Those on the carpus of the common pig are well known, and are
said to occur in both boars and sows. I figured and described
them as seen in a wild boar, Sus scrofa, but I subsequently failed
to find a trace of them in a sow of this species and also in a sow
of the Japanese species, Sws leucomystax, and I similarly failed
in the case of a boar of the Indian species, Sus cristatus.
T have had no opportunity of studying further the peculiar
glands in the feet of the male of Potamocherus porcus, which I
described a few years ago (Proc. Zool. Soc. 1916, p. 747), and
can add nothing to the original account.
Section TYLOPODA.
Family CAMELID&*,
The Muzzle.
The facial vibrisse in this family are so poorly developed as to be
practically negligible.
Text-figure 39.
Z Vie
A. Metatarsal gland of the Alpaca (Lama glama).
B. Muzzle of Lama huanacus with the lips spread and the nostrils
dilated.
C. The same of LZ. vieugna with the lips only partially spread and
the nostrils nearly closed.
The muzzle has the upper lip completely cleft, the two halves
being freely movable and separated by a philtrum of naked skin,
which extends down to the premaxillary gum-pad against which
* Tn all the examples of Zama that I have dissected, and in the one example of
Camelus dromedarius, I found the so-called “ water-cells” of the rumen packed
with food and not filled with water. They are no doubt primarily “ food-cells,”
and secondarily become filled with water in Camelus when food is unobtainable.
CHARACTERS OF THE CAMELIDA. 543
the lower incisors bite. The two halves of the upper lip hang
down on each side of this pad. The upper surface of the nose
and the area all round the nostrils, which are elongated dilatable
slits, are covered with fine short hairs.
In Lama vicugna and huanacus* the internarial septum is
hairy, and the philtrum is a short area about twice as long as
wide when expanded. (Text-fig. 89, B, C.)
The muzzle of Camelus dromedarius closely resembles that of
Text-figure 40.’
AW i.
Wir)
A. Muzzle of Camelus dromedarius with nostrils nearly closed.
B. Nostrils of the same dilated.
Lama, but is much deeper below the nostrils and has the phil-
trum very much longer. Inferiorly, moreover, the philtrum is
continued on each side as a narrow strip along the inner edge of
the lips, while it is slightly expanded above and abuts against the
inner ends of the nostrils. The nostrils are long and valvular,
* In Lydekker’s Catalogue of Ungulates, iv. p. 302, the Hunnnco is cited as
Lama glama huanacus, on the assumption that it is the wild form from which the
domesticated Llama and Alpaca were derived. The differences between the wild
and the tame animals are, however, too well marked to warrant the unqualified
acceptance of that opinion. The Llama, indeed, may represent a species wholly
reclaimed from the wild state.
544 MR. R. I. POCOCK ON THE EXTERNAL
and are lined for some distance inside with short hair. (‘T'ext-
fig. 40, A, B.)
The complete cleavage of the upper lip by a philtrum, and the
mobility of its two halves inferiorly where they project below and
on each side of the premaxillary gum-pad, are characters of the
Tylopoda as remarkable as any that are usually cited as diagnostic
of that section of Artiodactyla. .
The Occipital Gland of Camelus.
In my paper on the Specialised cutaneous scent-glands of
Ruminants (Proc. Zool. Soc. 1910, p. 973), I described the
occipital gland of the two species of Camelus as observable on
the living animals. I there stated that there are a pair of such
glands. ‘That was an error due to the interpretation of the gland
by touch. ‘here ig a single gland, about as wide as long and
Text-figure 41.
A. Transverso section of the occipital gland of Camelus dromedarius.
B. Longitudinal section of the same.
C. Baso of ear of Lama vieugna.
D. The samo of Camelus dromedarius.
covered with hair. It is composed of thickened skin, which
gradually thins out marginally where if runs into the normal
skin at the back of the head. Its lower half is rather thicker
than the upper, and in the middle there is a wide and tolerably
deep longitudinal depression, and it was the resulting biconvexity
of the gland which deceived me into thinking there were two.
(Text-fig. 41, A, B.)
CHARACTERS OF THE CAMELIDA. 545
The Ears. &
The ears of Lama are long, narrow, and pointed*. On their
inner surface they are strengthened by two longitudinal ridges. °
The posterior of these is slightly oblique, and does not descend
inferiorly into the posterior portion of the ear. The anterior is
straighter, and inferiorly rises close to the basal thickening,
which forms a rounded prominence. This thickening is pierced
below by the auditory orifice. ‘These ears are remarkably like
thoxe of the typical Ruminant Artiodactyla. (Text-fig. 41, C.)
The ear of Camelus dromedarius is much shorter and wider
and less freely movable on the head than that of Zama. The two
internal ridges are soft, low, and short, and probably little more
than functionless vestiges of those present in Lama. The basal
prominence is tolerably similar to that of Zama, but at its upper
extremity there is a deep transverse groove cutting off a ridge,
the position and structure of which suggest its homology with
the supratragus of normal mammals. The posterior extremity of
the ridge turns upwards behind the base of the anterior of the
two soft ridges above mentioned, -(Text-fig. 41, D.)
The Feet.
In Lama the feet are long and narrow, each of the digits
being provided with an elongated cushion-like pad, rounded
behind and narrowed in front, where it runs into the small,
somewhat nail-like hoof. Distally the digits are capable of being
widely separated by a deep cleft between them. Behind this cleft
the soles are united, but the extent of their fusion varies to a
certain degree. In an example of LZ. glama the fusion was so
complete as to obliterate superficially all trace of the line of
junction. But in LZ. vicugna there was a tolerably deep, wide,
and Jong groove between them. A similar but shallower and
narrower groove separated the heels in an Alpaca; and in this
animal the cleft between the digits extended half-way along
the soles, whereas in L. glama and L. vicugna the interdigital
cleft was shorter. On the upper side of the foot there is
a long, deep, interdigital depression lined with glandular integu-
ment, secreting a waxy substance smelling like the urine of
Mus nusculus. In L, vicugna and L. glama the floor and sides
of this depression were naked, but in an Alpaca they were clothed
with hair for the most part, only the distal edge of the depression
being naked on both fore and hind foot. On the latter the
proximal deepest portion, where the skin is most active, was also
naked, whereas on the fore foot the corresponding spot was
clothed with short radiating hairs. (Text-figs. 42, 43.)
In the Proceedings of this Society for 1916, p. 748, I described
and figured the two metatarsal glands of Lama vicugna. The
* In the domesticated races, but not in the wild species, the tips of the ears are,
typically at all events, curved slightly forwards. ay :
MQ. R. I. POCOCK ON THE EX'TERNAL
546
Text-figure 42.
A, B. Upper and lower views of foot of Zama glama (Alpaca variety).
C. Section of foot of Lama glama (common variety).
Text-figure 43.
=
A, B. Upper and lower views of foot of Lama huanacus.
CHARACTERS OF THE CAMELIDA. BAT
naked, depressed glandular area, overlapped by hair, was about
four times as long as wide, and gradually narrowed at its distal
end. In a male Alpaca the glandular area was more exposed
than in the Vicufia, was not depressed but flat, and was only
overlapped by hair at the extreme margin above distally. It was
also shorter and differently shaped, its length being 13 inches and
its breadth ? of an inch; the lower border was sinuously convex
and the upper had a submedian bulge; its proximal and distal
ends were approximately equal in width, the former being
7z inches from the hock, the latter 5 inches from the fetlock.
(Text-fig. 39, A.)
The feet of Camelus dromedarius are more specialized than
those of Lama. The two digits are fused nearly up to the hoofs,
so that their under sides form a continuous sole, about as wide as
long, with at most a shallow, irregular, median crease to mark
Text-figure 44.
Upper view of foot of Camelus dromedarius.
the line of union. At the digital end of the sole there is a
tolerably deep angular emargination where the digits are free
and diverge from one another. The fore foot is both wider and
longer than the hind foot, and in the specimen examined was
narrowed posteriorly, the hinder edge of the sole, or heel, showing
a shallow angular emargination. In the hind foot, on the con-
trary, the posterior border of the sole was convexly rounded, and
formed a continuous curve with the lateral margins, which, like
those of the fore foot, were tolerably evenly but lightly convex.
This hind foot was asymmetrical, the inner digit being larger
than the outer, and the proximal cleft between them was twisted
externally at its deepest part. On the upper side of both fore
and hind foot there is a deep, long interdigital depression lined
throughout with hair, and glandular, the secretion having a
strong odour like the urine of Jus musculus. The skin of the
Proc. Zoou. Soc.—1923, No. XXXVI. 36
548 MR. R, I. POCOCK ON THE EXTERNAL
sole is not very thick at the heel, but it thickens towards the
hoofs, forming a strong junction at this point between the two
digits. The inner surface of each digit between this junction and
the hoof is naked. (Text-figs. 44, 45.)
The Penis.
Athough the penis in the Camelidw has been previously
described, this organ in the Mammalia generally is so important
from the systematic standpoint that I venture to draw attention
to the profound differences it exhibits in this family from that of
the rest of the Artiodactyla. In the latter the glans is always
Text-figure 45.
A, B. Lower view of fore and hind foot of C. dromedarius.
C. Section of foot of the same.
soft and flexible. Considering its length, it seems remarkable
that in no case has an os penis or baculum been developed. In
the Camelide also there is no baculum, but the tip of the glans
instead of being soft is rigid, hard, and apparently cartilaginous.
The apex, moreover, is bent sharply to the left, like a stout hook,
underlying and apparently acting as a guard to the slender,
straight, forwardly directed process, at the tip of which the genito-
urinary orifice opens.
In the Alpaca and in Lama huanacus the hook is simple and
CHARACTERS OF THE CAMELIDA. 5A
nearly straight. In the latter it is slender, and is bent at right
angles to the axis of the glans; whereas in the Alpaca it is
stouter, and is bent at an obtuse angle. In Camelus dromedarius
the hook forms an almost rectangular bend, but is slightly curved
and has a lightly convex anterior and a concave posterior edge.
Text-figure 46.
B
A. Tip of glans penis of Zama huanacus from above.
B, C. Lower and upper views of the same of Z. glama (Alpaca).
D. The same of Camelus dromedarius trom above.
Externally on the right side it forms a shoulder-like excrescence,
defined posteriorly by a deep concavity from the main part of the
glans; and on the left side there is a soft projecting lappet on the
glans behind and beneath the base of the little process carrying
the genito-urinary orifice. (lext-fig. 46.)
36*
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ON TIE DIPTEROUS FAMILY PANLOPHTHALMIDA, 551
30. A Revision of the Family PanropnrHaLmipaz [ Diptera],
with Descriptions of New Species and a new Genus.
By Major E. H. Ausren, D.S8.0., F.Z.S.
[Received May 29, 1923: Read June 12, 1923.]
(Text-figures 1-11.)
TABLE oF ConTENTS.
: Page
(a) Introduction and Lae havens cagosunsonpacsamd Lait
(GO) MBibliograp hives eceeeentesesesseac eens. dstacnen us uamooesenton Oe
(c) Geographical DRC IEEE. pe AAPA ARIA cece RIS eRe EN SO)
(d) Taxonomic Position ..... .... 653
(e) Bodily Dimensions in Dargaaieemnea aonpensll viol
those of Exceptionally ave Diptera belonging to
other Families.—Giantism.. popGobcos ooHAdesGodeco hop Beane | ULI)
(f) Life-history and Bionomics ...............s000eseeseeseeeeees. 556
(g) MBseudosParasitesinc:-ssstiiere see ceeds eee DS
(®uSystematicyRoutioniee esas eee ee eee OD
(a) Introduction and Acknowledgments.
In spite of, as compared with the vast majority of Diptera,
imposing size and striking coloration—characters likely to attract
the attention of collectors in countries in which the flies forming
the subject of this paper are found—Pantophthalmide (formerly
known as Acanthomeride) are singularly rare in collections. It
is difficult to suggest even a possible reason for this, unless it is
to be found in the habits of the adult insects, which probably
spend most of their time in the depths of Neotropical forests.
At the present moment the series of these great Diptera in the |
British Museum (Natural History), though doubtless the most
extensive in any collection in Europe, consists of only sixty
specimens. ‘These, however, include no fewer than twenty-four
examples from the collection of the late M. J. M. F. Bigot,
which Mr. J. EB. Collin, F.E.S., has most generously presented to
the National Collection during the preparation of this paper.
The fact that among M. Bigot’s series of specimens are the
types of ‘five species, described | by himself, Maequart or Bellardi,
renders this addition peculiarly valuable, and its timely acqni-
sition has enabled the writer in the course of subsequent pages
to elucidate for the first time a considerable amount of synonymy.
It is therefore a pleasant task hereby to express to Mr. Collin, on
behalf of the Museum, sincere thanks for his public-spirited ana
most welcome gift.
Acknowledgments are also due to Professor E. B. Poulton,
¥.RS., F.Z.8., for the kind loan of the Pantophthalmid material,
552 : MAJOR E. E. AUSTEN ON THE
amounting to twenty specimens, in the Hope Department of the
Oxford University Museum; and to Dr. Hugh Scott, F.E.8.,
for similarly allowing the author to examine a female of Pant-
ophthalmus chuni Knderl. and a larva (represented in text-fig. 1,
on p. 557) of P. tabaninus Thunb., both of which are the property
of the University Museum of Zoology, Cambridge.
It only remains to add that, unless otherwise stated, all speci-
mens mentioned in the following pages, including the types of
new species, are in the British Museum (Natural History).
(b) Bibliography. |
Apart from descriptions of genera and species already pub-
lished, references to which will be found in the ‘Systematic
Portion” (h) of this paper, the more important items comprised
in the scanty literature of Pantophthalmide are the following :—
Braver, Fr.—Description of a dried larva of Pantophthalmus
(Acanthomera) frauenfeldi Schin, Denkschr. K. Akad.
Wiss., Math.-Naturw. Cl., Bd. xlvii. p. 25, Taf. i.
figs. 25 a-25 e (Wien, 1883).
Osten Sackrn, Baron C. R.—‘“‘ Fam. Acanthomeride.” Biol.
Centr.-Amer., Diptera, vol. i. pp. 63-68 (1886).
Fiesrie, K.—‘‘ Kine morphologisch und biologisch interessante
Dipterenlarve aus Paraguay (Acanthomera teretruncum
sp. n. Fiebrig).” Zeitschr. f. wiss. Insektenbiologie,
Bd. ii. pp. 316-323, 344-347, figs. 1-19 (1906).
Wiuutston, 8. W.—“ Family Acanthomeride.” Manual of N.
Amer, Diptera, 3rd Ed., pp. 173-175, figs. 2, 61, 62
(London: Wm. Wesley & Son, 28 Hssex Street, Strand,
1908).
Enperirin, G.—“ Die Dipteren-Familie Pantophthalmidx.” Zool.
Anzeiger, Bd. xli. pp. 97-118, figs. 1-15 (1912).
ENDERLEIN, G.--~‘‘ Weitere Beitrige zur Kenntnis der Pant-
ophthalmiden.” Zool. Anzeiger, Bd. xliv. no. 13, pp. 577-
586 (July 28, 1914).
Hermann, F.—‘ Hin neuer Pantophthalmus nebst kvitischen
Bemerkungen tiber die Systematik der Pantophthalmiden
(Dipt.).” Deutsche Ent. Zeitschrift, Jahrg. 1916, pp. 43-
49, Abt. 1, 2 (1916).
(c) Geographical Distribution.
Pantophthalmide are confined to the Neotropical Region,
where, however, their range is wide, extending from Mexico to
Paraguay, and including at least a portion of the West Indies.
It is true that, as pointed out by Osten Sacken (Joc. cit. p. 66),
Thunberg’s statement (cf. infra, p. 562) as to the provenance of
the type of Pantophthalmus tabaninus—the first species of this
family to he described—“ still requires confirmation.” Neverthe-
DIPTEROUS FAMILY PANTOPHTHALMID. 553
less, considering that some eleven years ago the same species (sub-
sequently regarded by Knab as new, and described by him under
the name Pantophthalmus fastuosus) was found by Mr. F. W. Urich
breeding in Trinidad; that, as is well known, the Dipterous
fauna of the West Indies generally is “ essentially a common one,
with a strong South American facies” *; and that the islands of
the Lesser Antilles are only separated one from another by short
intervals of sea—there can be no real reason to doubt the accu-
racy of Thunberg’s century-old assertion that the type of this
species came from St. Barthélemy I.
(d) Taxonomic Position.
Williston (‘ North American Diptera,’ 3rd Kd., p.175, 1908), who
places the “ Family Acanthomeride ” between Stratiomyiide and
Tabanide, writes:—‘‘ The relationships of the family are very
close indeed to the Stratiomyide, and the families might, very
properly, be united.” Kertész, however, in vol. ili. of his ‘ Cata-
logus Dipterorum,’ published in the same year, gives the families
embraced by the volume in the following sequence: Stratio-
mylide; Hrinnide (Xylophagide); Ccnomyiide; Tabanide ;
Pantophthalmide ; Rhagionide (Leptide olim). Hnderlein, on
the other hand (Zool. Auz., Bd. xli. p. 97, 1912), considers that,
from a phylogenetic standpoint, the Pantophthalmide are extra-
ordinarily closely allied to the Xylophagide and Cenomyiide,
and that the morphological relations between all three are so pro-
nounced that they might be regarded as groups belonging to a
single family, namely the Xylophagide. The latter would then
be divided into the subfamilies Xylophagine, Coenomyiine, and
Pantophthalmine. Remarking that the presence or absence of
spines on the scutellum, the relative size of the head and differences
in bodily shape, whether slender or thickset, furnish insufficient
grounds for the separation of families, Enderlein proceeds to
show with the help of a diagram ‘that in venation also the dif-
ferences between Pantophthalmide, Xylophagide, and Cceno-
myiide are unimportant. Hermann, in his paper published four
years later, while criticising Enderlein somewhat severely in con-
nection with other details, nevertheless (Deutsche Ent. Zeitschr.,
Jahrg. 1916, p. 47, 1916) expresses his complete agreement with
him (and disagreement with Kertész) in respect of the closeness
of the relationship between the Pantophthalmide, regarded as a
family, and the Xylophagide and Ceenomyiide.
There can be no doubt that the views of Enderlein and:
Hermann as to the systematic position of the Pantophthalmidse
are correct, and indeed, in order to be satisfied upon this point,
it is only necessary to compare two representative species, such
_as Pantophthalmus pictus Wied. and Xylophagus ruficeps Lw.
In view of the life-history of Pantophthalmide (vide infra (f)),
* Williston, Trans. Ent. Soc. Lond. 1896, p. 445 (1896).
554 MAJOR E. E. AUSTEN ON THE
it is perhaps worth while to recall that the preliminary stages of
Xylophagids and Ccenomyiidew are also associated with trees,
albeit in a dead or mouldering condition, the larve, which in
the case of the two latter families are carnivorous and predaceous,
living in decaying wood and under bark.
(e) Bodily Dimensions in Pantophthalmide compared with those
of Haceptionally Large Diptera belonging to other Famalies.—
Giantism.
Although, as will be seen directly, in wing-expanse, and of
course in length of leg, Pantophthalmide are surpassed by
certain Tipulids of abnormal size, while length and wing-expanse
in the largest species are about the same as or somewhat below
the corresponding dimensions of the largest representatives of the
Mydaide, the members of the present family as a whole are
undoubtedly by far the bulkiest, and therefore the largest of all
Diptera.
Among slenderly built Diptera, the largest forms are to be
found in the Vipulide, wherein, as regards actual size, certain
species of the genus Céenacroscelis Enderlein are probably with-
out existing rivals, Thus, a male belonging to a variety of
Ctenacroscelis brobdignagius Westw., from China (N.-W. Sze
Chuen), just over 39 mm. (1 inch) in length, has a wing-expanse
of 104 mm. (rather more than 4 inches), while the outstretched
legs cover an area at least 67 inches in length by 5 inches in
breadth.
Although the family Mydaidw includes a number of species,
the representatives of which are of moderate dimensions, certain
members of the genus Mydas, which are among the largest
of Diptera, are relatively gigantic. By way of illustration,
mention may be made of Mydas pregrandis Austen, a male of
which from Brazil has a wing-expanse of 85 mm, (between 37 and
34 inches), while the length of this insect, exclusive of the long
and prominent antenne, is 48 mm. (between 1? and 2 inches)*.
The dimensions of this species, in fact, as represented at any rate
by the type and paratype (both males) in the National Collection,
are but slightly larger than those of the somewhat aberrant re-
presentative of the Pantophthalmide originally described by the
* Writing with reference to what is almost certainly the female of this species,
of which he gives a life-size illustration derived from a photograph, Williston
(N. American Diptera, pp. 16, 17, fig. 1, 1908) furnishes considerably higher measure-
ments, remarking :—‘ The largest specimen of a fly of which I have knowledge is
‘that figured herewith natural size, pertaining to an indeterminable species of Mydas
from South America. ‘he length of this specimen from the tip of the antenna to
the extremity of the abdomen, is sixty-seven millimeters, or, omitting the antenne,
fifty-two millimeters; the expanse of wings one hundred and seventeen millimeters,
or a little more than four and one-half inches. The smallest dipteron that I have ever
observed in the examination of many thousand specimens and five or six thousand
species, is a cecidomyid measuring a trifle less than one-half millimeter, also omitting
the antennse. In other words, the Mydas is more than one million times the size
of the cecidomyid. Possibly there are still greater discrepancies between the largest
and smallest specimens of the order, but in all probability not much,”
DIPTEROUS FAMILY PANTOPHTHALMIDE. 555.
author as Lhaphiorhynchus rothschildi, for which a new genus is
erected in the present paper (cf. p. 596).
Turning to the Asilidz, we find in this enormous family a wide
range in size, and a considerable number of species which, though
smaller than the larger examples of the genus Mydas such as
that just mentioned, are yet of exceptional dimensions, some of the
largest forms being met with, for example, in the genera P70-
agonistes, Alcimus, Phellus, Blepharotes, Hyperechia, and Procta-
canthus. A male belonging to an undescribed species of
Proagonistes from Madagascar, 41 mm. (between 14 and 1? inch)
in length, has a wing-expranse of 80 mm. (between 3 and
32 inches); while, owing to the extraordinary length of the hind
legs, the space from the tips of these, when outstretched, to those
of the similarly extended front legs measures at least 82°5 mm.
(37 inches). The slender-bodied Alcimus brevipennis Ric., from
Natal, has in the female sex a length of 44 mm. (just under
12? inch), thongh its wing-expanse amounts only to 55 mm.
(between 2 and 24 inches). Among the much more stoutly built
representatives of the Australian genus Phellus, a female from
South Queensland, belonging to a species as yet undescribed,
measures 50 mm. (just under 2 inches) in length inclusive of the
ovipositor, and has a wing-expanse of 81 mm. (a little under
32 inches). The female of Blepharotes coriarius Wied., from the
same region, may measure 42 mm. (between 14 and 1# inch) in
length, and have a wing-expanse of 76 mm. (3 inches). Several
of the species of Wyperechia are bulky-bodied flies of large size, one
of the largest being H. (Laphria) consimilis Wood (syn. Dasyllis
usambare Lichtwardt), of which the type (a female from Natal)
is 33 mm. (over 1? inch) long, and has a wing-expanse of 62°5 mm.
(very nearly 24 inches). Among narrow-bodied Asilidee (sub-
family Asilinw), an exceptionally large species is Proctacanthus
(Asilus) penultimus Walk., the type of which—a male from India
—-is 40 mm. (over 14 inch) long, and has a wing-expanse of
60 mm. (between 2} and 22 inches).
Lastly, as regards Pantophthalmide other than the species
already mentioned, the wing-expanse in the case of females
belonging to the largest forms such as Pantophthalmus bellardiu
Bell. may amount to 84:5 mm. (a little over 33 inches), and the
length over all (z.e. including the ovipositor but not the antenne)
to 54 mm. (over 2 inches); while the broad, flat abdomen may
measure 22 mm. (considerably more than ? inch) in width at its
widest point. In the female sex of certain other species, notably
Pantophthalmus tabaninus Thunb., dimensions are often little
below those just given. Species of Pantophthalmide however
exist, in which, so far as it is at present possible to judge from
the limited amount of material available for examination, the
same standard of bodily development is not attained; and on
the other hand there are indications that, in some cases at any
rate (as, e.g. in P. tabaninus), the average size of the males is
considerably less than that of the females.
556 MAJOR I. E. AUSTEN ON THE
The fact remains, however, that in the adult state dimensions
in Pantophthalmide as a whole are greatly in excess of the normal
in Diptera, and this small group of large flies consequently forms
an interesting case of the phenomenon known as giantism, con-
cerning which Williston (op. cit. p. 17) writes as follows :—
‘“‘Giantism in any group of animal life is a specialization, and is,
in general, an indication of approaching decadence; enduringly
small races ave never the descendants of giants, for decrease in
size means lessened vitality and incipient extinction. No strong
or dominant group of flies, like the Tachinide, Dolichopodide,
Syrphide, or Bombyliide, has ever had in the past a larger
average bodily size than is found among their living representa-
tives. On the other hand, those families composed to-day chiefly
of large forms are ones already past their prime. These con-
clusions seem established for the larger forms of life, and I believe
that they are in the main also applicable to insects.”
(f) Life-history and Bionomics.
Until the year 1906, knowledge under this heading was limited
toa brief, illustrated description by Brauer of a dried larva of
Pantophthalmus (Acanthomera) frauenfeldi Schin. from Bogota
(cf. “ Bibliography,” p. 552); and Osten ‘Sacken’s statement
(Biol. Centr.-Amer., Diptera, vol. i. p. 66), published three years
later on the authority of Mr. G. 0. Champion, that ‘‘ Acantho-
mere are found in forests, alighting on trunks of trees.” In the
year referred to, however, Karl TFiebrig, of San Bernardino,
Paraguay, working under the disadvantages contingent upon
isolation and lack of entomological literature, published a most
interesting account (cf. ‘ Bibliography,” p. 552), illustrated by a
series of figures, of the external and internal anatomy of the larva
of Pantophthalmus pictus Wied. (infra, p.575). From the larvee
found by him, Fiebrig succeeded in breeding out a pair of
imagines, which, owing to the disadvantageous conditions just
mentioned, he regarded as belonging to a new species, and
accordingly described under the name Acanthomera teretrumcum.
Thanks to Fiebrig’s observations, which were actually made in
1905, and were confirmed seven years afterwards by Mr. F. W.
Urich, in Trinidad, in the case of another species (Pantophthalmus
tabaninus Thunb.), we now know that larve of Pantophthalmide
have the extraordinary habit—unique among Diptera, so far as
the present writer is aware—of boring in the solid wood of living
trees, in which they make ‘nearly horizontal” tunnels, and feed
upon the exuding sap. Although Fiebrig does not mention the
name of the tree in which he found his larvew, he states that its
wood is “among the toughest of Paraguayan timbers, hard as
these in general are,” and that the tunnels made by the maggots
are “ sharply gouged out.”
In view of Fiebrig’s detailed account, which should be consulted
by the student, it is searcely necessary here to do more than draw
DIPLEROUS FAMILY PANTOPHTHALMID. 557
attention to the accompanying figure (text-fig. 1), which displays
the chief features in the external morphology of a larva—probably
nearly adult—of another species of the genus Pantophthalmus.
This larva, which is preserved in spirit, is 32°5 mm. (just over
12 inch) in length, while the average ventro-dorsal diameter ot
the body-segments is 9°75 mm. It was taken, with at least five
other larvee of the same kind, in the Island of Trinidad, B.W.L,
in borings in the trunk of an Hrythrina, a leguminous tree, which:
attains a height of from 15 to 20 feet.
Text-figure 1.
Cc THRAIX
Larva of Pantophthalmus tabaninus Thunb, (lateral view), from boring in trunk of
Erythrina tree (I. corallodendrum I..?), Trinidad, B.W.1., March 1912
(F. W. Urich). X 23.—From specimen kindly lent by Dr. Hugh Scott,
University Museum of Zoology, Cambridge.
a, labrum; 8, left mandible; ¢, left anterior (prothoracic) spiracle or stigma;
d, dorsum of 8th post-thoracic segment, which is very heavily chitinized; e, Fiebrig’s
“finger-shaped bodies”— in all probability an auxiliary respiratory apparatus,
employed when the posterior stigmata are unable to function, owing to the larval
tunnel being full of sap.
The stout labrum and the extremely powerful mandibles together form a “ drill,”
by means of which the larva is able to bore in solid wood; the equally heavily
chitinized dorsum of the 8th post-thoracic segment closes the tunnel from behind,
and protects the larva from an attack in the rear. The posterior stigmata open on
the posterior (ventral) surface of this segment, in a mouth-like chamber between it
and the 9th post-thoracic segment, which bears the “ finger-shaped bodies.” When
the latter sezment is in the position shown in the figure, the posterior stigmata are
completely closed.
In the adult Pantophthalmid larva, the number, size, and arrangement of the
backwardly directed spines on the 8th post-thoracic segment perhaps afford
characters of specific importance (cf. Brauer, loc. cit. Taf. ii. figs. 25 a, 25 b).
In the Trinidad larva, the longitudinal strips of chitin em-
bedded in the integument of the prothoracie segment, to which
the striped appearance of the latter is due, are reddish brown,
while the powerfully developed labrum and mandibles, and the
heavily chitinized dorsal surface of the 8th post-thoracic segment
are deep black; the prevailing tint of the remainder of the body
is that of old ivory. ‘There can be little doubt, however, that
prolonged immersion in alcohol has had a bleaching effect, since
Fiebrig describes the nine “soft” body-segments in the larva
discovered by him as “bluish green,” owing to the “ intense blue
tint” of the abundantly developed fat-body ; it is also worthy of
558 MAJOR E. E. AUSTEN ON THE
note that in life, according to the same author, the “ finger-shaped
bodies” (see text-fig. 1 and explanatory text) are bright green.
The dorsal surface of the prothoracic segment, though far less
heavily armoured than that of the 8th post-thoracic segment,
nevertheless, strengthened as it is in the manner already referred
to, is quite hard, and may well serve to protect the tunnelling
larva against possible injury from sharp splinters of wood.
An interesting point with reference to the living larva. of
/Pantophthalmus pictus is noted by Fiebrig, who writes that,
“when in the tree-trunk it produces rasping sounds, distinctly
audible up to a distance of some four paces.”
The pupal envelope, according to the same observer, is vertically
truncated at each end, “the anterior extremity being formed by ,
a stout, richly sculptured plate of chitin,” which serves to protect
the pupa from a frontal attack. Although this armoured shield
is not broken through by the imago in emerging, it may be that
the facial ‘‘ beak,” that is so conspicuous a feature in many Pant-
ophthalmide though not exhibited by all species, or always
developed in both sexes, when present assists the fly in making
its escape from the pupal envelope. The latter, we are told by
Fiebrig, “after the emergence of the imago, projects about half-
way out of the larval tunnel”; but the same authority is unable
to state whether the pupa makes its way to this point by its own
efforts, or whether the pupal envelope is dragged along with it
by the escaping fly.
“Metamorphosis,” writes Fiebrig, “appears to be very: pro-
tracted. A larva 35 mm. in length, and apparently nearly full-
grown, was kept under observation from July 31, 1905, onwards.
- The imago emerged in the afternoon of January 11, 1906, a few
weeks before which date it was found that the insect was still in
the larval state; this would point toa pupal stage of very short
duration.”
Whether larvee of Pantophthalmide are of economic importance,
as destroyers of timber of commercial value, is an interesting
point which awaits determination. So far as can be ascertained,
the wood of Hrythrina (E. corallodendrwm L.%), in which the
larvee of Pantophthalmus tabaninus Thunb. were found by Mr.
F. W. Urich, in Trinidad, is of little or no utility. But the
possibility of commercial value is at any rate not excluded in the
casé of the unknown Paraguayan tree referred to by Fiebrig, as
to which we are told that the wood is exceedingly hard and
extremely rich im sap, ‘yielding an abundance of evil-smelling
fluid on being injured.”
b
(g) Psewdo-Parasites.
Pseudo-parasites in the shape of Gamasid mites are of extra-
ordinarily common occurrence on Pantophthalmide, and many
instances of such infestation are recorded in the following pages.
The mites, on being submitted by the author to his colleague
Mr. A. 8. Hirst, F.Z.8., for identification, were found by him in
DIPTEROUS FAMILY PANTOPHTHALMIDA, 559
every instance except one to be conspecific, and to be nymphs of
a species of Z'rachytes, which Mr. Hirst will describe in a forth-
coming paper as new (cf. p. 565). In the single case forming the
exception, the pseudo-parasite was determined by Mr. Hirst as
anadult female of AMacrocheles, sp. incert. (vide p. 588). It would
seem that these pseudo-parasites must infest the borings of the
larve, and attach themselves to the flies as tho latter emerge
from the pupa-case.
(h) Systematic Portion.
Taking into account synonymy already established, the total
number of species of Pantophthalmide at present recognized
amounts to twenty-four. In the following pages ten species,
previously described by various authors, are admitted as valid
and commented upon, notes being given on the specimens of
these species in the British Museum (Natural History), or other-
wise available to the writer for examination ; the names of nine
other species, hitherto accepted, are sunk-as synonyms; and six
species and one genus are described as new. As the result of
this paper, therefore, the number of known species of Pant-
ophthalmide, including those that the author has been unable
to examine, will stand attwenty-one. It is explained below that,
in agreement with Kertész, and contrary to the course adopted
by Enderlein, the genus Pantophthalmus Thunb. is treated as
including Acanthomera Wied. In view of the limited amount
of material at the author’s command, and since as yet both sexes
are known in the case of only a minority of species, with much
regret no attempt has been made to construct a synoptic table.
Should the publication of the present paper fortunately result in
a considerable expansion of the National Collection of these
interesting Diptera, the construction of a reliable table of the
kind in question may eventually be possible.
Genus Pantophthalmus Thunberg.
Pantophthalmus Thunberg, Gétheborgs Kong]. Wettenskaps
och Witterhets Sambhiillets Nya Handlingar, iii. Delen, p. viii
(1819).
Acanthomera Wiedemann, Diptera Exotica, Parsi. p. 60(1821).
Megalomyia Bigot, Ann. Soc. Ent. France, 5° Sér., T. x., Bull.
p. v (1880).
Megalemyia Bigot, Ann. Soc. Ent. France, 6° Sér., T. i. p. 455
(1881).
After due consideration, it has seemed better to follow Kertész
(Cat. Dipt. ili. p. 294 (1908)), who gives the synonymy printed
above, rather than Enderlein (Zool. Anz., Bd. xli. p. 100 (1912)),
by whom Pantophthalmus Thunb. and Acanthomera Wied. are
treated as generically distinct. The course adopted by Kertész
appears the more reasonable one under the circumstances, in spite
560 MAJOR E. E. AUSTEN ON THE
of lack of complete agreement among the species mentioned or
described in the following pages, in respect of certain structural
details. Included among the latter are:—The degree of develop-
ment of the facial ‘‘ beak”; the shape of the terminal segment of
the maxillary palpus; the shape of the terminal division of the
third (compound) segment of the antenna in the female; and
the presence or absence of a distinct subfemoral spine on the hind
legs.
“Although, by concentrating attention on one or other of these
characters and disregarding the remainder, it would perhaps be
possible in the case of certain species to ‘select. distinctions of
apparently generic value, yet in other forms intergradations
occur which make the drawing of a hard and fast line of demar-
cation impossible. Thus, to take only one of the characters
referred to:—Enderlein (loc. cit. p. 99), writing of the terminal
division (called by him the tenth segment) of the female antenna,
says that this “in Pantophthalmus tapers gradually to a point,
but in Acanthomera and Khaphiorhynchus becomes but little
narrower towards the end, and then, just before the tip, i
suddenly pointed.” This statement is certainly true if we com-
pare the antenna of the female in species such as Pantophthalmus
tabaninus Thunb., or P. bellardii Bell. (both of which belong to
Pantophthalmus, sensu stricto) with that of the female of e.g.
Pantophthalmus (Acanthomera) chuni Enderl. But when we
examine the antenna in the female of Pantophthalmus pictus
(Acanthomera picta) Wied.—a species with a well-marked sub-
femoral spine, and the genotype of Acanthomera—we find that it
ig intermediate between the two conditions. Moreover, in the
female of Pantophthalmus vittatus (Acanthomera vittata) Wied.—
a species assigned by Enderlein himself (Joc. cit. p. 107) to
Pantophthalmus—the terminal division of the antenna is precisely
of the type regarded by Enderlein as characteristic of Acanthomera.
As regards the maxillary palpus, the terminal segment of this
structure is also subject to a certain amount of variation in shape
in different species. Although usually slender and cylindrical,
and, at least in the female sex, longer than the remainder of the
palpus, it is sometimes shorter and also stouter, albeit never
swollen in the-manner characteristic of the terminal segment of
the palpus in Lhaphiorhynchus.
That Enderlein is in error when he asserts (loc. cit. p. 160) that
in Pantophthalmus, sensi stricto, the facial beak is represented
merely by a “rounded tubercle-like swelling” is pointed out by
Hermann (Deutsche Ent. Zeitschr., Jahrg. 1916, p.48 (1916)), who
shows that in P. alienws Herm. the absence of a subfemoral spine
on the hind legs is correlated with the presence of a “long,
pointed, tooth-like facial process.” A further example of the
condition seen in P, alienus Herm, is afforded by the new species
described below as P. batesi (cf. p. 566), in the male of which, at
any rate, while the facial beak, though small, is conspicuous and
sharp-pointed, the posterior femora are either without a spine
DIPTEROUS FAMILY PANTOPHTHALMIDA. 561
on the under surface, or display the merest vestige of such a
structure.
The truth is that characters such as those mentioned above,
however valuable for the distinction of species, are of less than
generic importance, and, were it necessary, many arguments in
support of this contention might be furnished from the accepted
taxonomy in other families of Diptera.
Among characters of specific importance in the genus Pant-
ophthalmus, may be mentioned wing-markings and, although this
does not apply to the males of certain species which have the
thorax unicolorous or nearly so, the markings on the dorsum of the
thorax.
Wing-markings, in spite of their general similarity (with
certain exceptions) throughout the family Pantophthalmide, will
be found, if close attention be paid to details, to afford useful
assistance not only in the distinction of species, but in the correct
association of the sexes of the same species—a matter in some
ases probably of far greater difficulty. Osten Sacken, in the
course of a valuable disquisition on structural peculiarities and
characters of specific importance in Pantophthalmide (‘“ Acantho-
meride,” Biol. Centr.-Amer., Diptera, i. pp. 65-66 (1886)), does
not refer to wing-markings, but writes as follows with regard
to markings on the thorax :—“The arrangement of the stripes
and spots on the thorax, an apparent monotony notwith-
standing, offers excellent characters for the recognition of the
species, and especially for the assorting together of male and
female specimens of the same species.” As regards the value of
thoracic markings for the association of the sexes of a given
species, the statement just quoted is only partially true. It cer-
tainly hold goods in the case of Pantophthalmus tabaninus Thunb.
(the synonymy given on p. 562 below is based on and supported
by the characteristic thoracic markings exhibited by this species), .
and P. vittatus (Acanthomera vittata) Wied. (as also in that of
Rhaphiorhynchus planiventris Wied.); but it is entirely
inapplicable to a species such as Pantophthalmus pictus (Acantho-
mera picta) Wied., in which the dorsum of the thorax is con-
spicuously striped in the female, but almost uniformly silvery, or
at any rate without corresponding stripes, in the male. While
the presence of .sharply-defined, dark longitudinal stripes on the
dorsum of the thorax (scutum) is characteristic of the female sex
in Pantophthalmus (and also in Rhaphiorhynchus), there are, in
addition to DP. pictus Wied., at least three species, including one
described for the first time in the following pages, in which, while
the female is as yet unknown, the thorax in the male is without
such stripes. In these species the dorsum of the male abdomen,
be if noted, has a silvery sheen. It is to be hoped that males and
females of the species in question may ere long be caught in coitd,
or bred from larve or pupw taken in the same tree-trunk, other-
wise the correct association of the sexes in these cases may remain
a matter of doubt for an indefinite period.
562 MAJOR E. E. AUSTEN ON THE
In addition to sexual dimorphism of the kind just referred to,
difficulties of determination in this genus are increased by the
fact that in the case of certain species (at any rate P. tabaninus
Thunb. and P. pictus Wied.—see below) the female abdomen, at
least in dried specimens, sometimes shows colour-dimorphism as
well. Failure to realize the occurrence of this phenomenon led
Enderlein into error in constructing his synoptic tables (Zool.
Anz. xli. pp. 101, 107 (1912))—a fact which has already been
pointed out by Hermann (Deutsche Ent. Zeitschr., Jahrg. 1916,
p. 48 (1916)).
The number of species of Pantophthalmus recognized as valid
in Kertész’s ‘Catalogus’ (vol. iii. 1908) is fourteen, This total
does not include Acanthomera teretruncum Fiebrig (= Pantoph-
thalmus pictus (Acanthomera picta) Wied.), since the description
of the latter (1906) appeared between the ‘compilation and
publication of the volume of the late Dr. Kertész’s monumental
work in which the Pantophthalmide are included. Between
1908 and the present time, descriptions of nine species of Pant-
ophthalmus supposed to be new have been published, includ-
ing seven by Enderlein and one each by Hermann and Knab;
while Enderlein has rightly sunk Fiebrig’s designation Acantho-
mera teretruncum as a synonym, besides disposing of P. helleri
Ender]. in a similar manner. Thus for the moment the number
of species of Pantophthalmus assumed to be valid stands at
twenty-two. In the following pages, however, the names of no
fewer than nine of these are shown to be synonyms, while six
species are described as new; so that, as the result of the
present paper, the genus Pantophthalmus is to be regarded as
consisting of nineteen recognized species.
The genotype is Pantophthalmus tabaninuws Thunb.
Pantophthalmus tabaninus Thunb.
Pantophthalmus tabaninus Thunberg, loc. cit. p. vii (1819).
Acanthomera seticornis ‘Wiedemann, Auss. zweifl. Ins. i.
p. 108, Taf. ii. figs. 1-a—-1 ¢ (1828). [New synonym. ]
Acanthomera heydenii Wiedemann, op. cit. p. 555 (1828).
[New synonym. ]
Acanthomera immanis Wiedemann, op. cit. ii. p. 623 (1830).
| New synonym. |
Acanthomera rubriventris Bigot, Ann, Soc. Ent. France, 5° Sér.,
T. x., Bull. p. v (1880). [New synonym. ]
Pantophthalmus fastuosus Knab, Insec. Inscit. Mens. vol. ii.
no, 2, p. 27 (February 1914), [New synonym.]
According to Thunberg (loc. cit.), the type of P. tabaninus was
obtained in St. Barthélemy I., in the: West Indies.
Among the synonyms recorded above, the name Acanthomera
seticornis (like Pantophthalmus tabaninus itself) was originally
applied to the ¢ insect, while the designations Acanthomera
heydenit, A. immanis, and A. rubriventris were bestowed upon
DIPTERKOUS FAMILY PANTOPHTHALMID&. 563
the 9. A study of the thoracic markings, which are character-
istic of this species and present a striking agreement in both
sexes, leaves no doubt as to the correctness of the synonymy here
given. In spite of the fact that, Pantophthalmus gigas Wnderlein
(Zool. Anzeiger, Bd. xli. p. 110, fig. 7 (1912)), described from
the 2 sex, is stated by its author (doe. cit.) to be devoid of pale
spots on the lateral margins of the second and third abdominal
tergites, it would seem not unlikely that this supposed species is
also identical with P. tabaninus Thunb.
The representatives of Pantophthalmus tabaninus Thunb. at
present contained in the British Museum (Natural History) are
as follows:—-One 6, (?) Trinidad, B.W.I. (@. Robertson—ex coll.
W. W. Saunders); one 2, Venezuela, before 1847 (purchased
from — Dyson); one @, Brazil, before 1849 (Mrs. Noel); one 3,
locality unknown (ev coll. Alexander Fry); one. ¢, Trinidad,
B.W.I., before 1878 (Ars. Cutter); one. 3, Macapi, R. Amazons,
Brazil, 25.11.1896 (#. 4. Austen), “on board Messrs. Siemens
Bros.’ Cable 8.8. Haraday”; one 2, Nicaragua (H. A. Burns);
one 2, Colonia Hansa, Santa Catharina, Brazil (purchased from
H. Rolle). Presented by Mr. J. E. Collin, F.E.S., from the
Bigot collection, we also have the four specimens (including the
type) of Acanthomera rubriventris Big., which are all 9 9 from
Guatemala; and two ¢ ¢ of P. tabaninus, likewise from Guate-
mala, which, in Bigot’s series, were placed with a ¢ of P. versi-
color Austen (see below), above the label “Acanthomera seticornis,
Wied. 3,” in Macquart’s handwriting.
The Hope Department of the University Museum, Oxford,
possesses two gd ¢ and two 2 2 of P. tabaninus; of the former,
one specimen is labelled “ seticornis Wied.,” but is without any
indication as to provenance, while on the label of the other is
written ‘‘ Para. Higgins, 1868”; there is nothing to show the
origin of the two 2 2, which are labelled “ 22. planiventris.”
Lhe Liverpool School of Tropical Medicine has a 9 of. P,
tabaninus from Para, Dec. 1922 (Dr. Clarke).
In this species, in which the female usually, although not
invariably, much exceeds the male in size, the ground-colour of
the dorsum of the abdomen ranges from cinnamon-rufous * or
vinaceous-rufous to blackish-brown or black. The dark thoracic
stripes are the same in all cases, and, at least in the specimens
available to the writer for examination, the silvery-white
triangular spot near each posterior angle of the second and third
abdominal tergites is always visible, although in some individuals
less conspicuous than in others. In the case of the type of
P. tabaninus, which, from the details furnished by Thunberg, is
evidently a male, the dorsum of the abdomen, according to the
original description, is “ brown”; Wiedemann (loc. cit.) describes
the dorsum of the abdomen in Acanthomera seticornis as ‘ vivid
* Tor names and illustrations of colours used for deseeiDUg purposes in the
presont paper, seo Ridgway, ‘Color Standards and Color Nomenclature’ (Wash-
ington, D.C. Published by the Author, 1912),
Proc. Zoot. Soc.—] 923, No, XX XVII. 37
564 MAJOR E. E. AUSTEN ON THE
ferruginous-yellow” (sehr brennend rostgelb), and the same region
in A. immanis as “ ferruginous.” In the type of A. rubriventris
Big. the ground-colour of the first four abdominal tergites is
between ferruginous and vinaceous-rufous, while the hind borders
of tho second and third tergites, and the hind border of the
fourth tergite except its lateral extremities, are black. Of the
three paratypes of A. rubriventris, one agrees with the type in
the coloration of its abdomen, while in the case of the other two
the dorsum of the abdomen is reddish-brown. In the series of
specimens of P. tabaninus in the National Collection, other than
those just mentioned under Bigot’s designation Acanthomera
rubriventris, the six males have the dorsum of the abdomen
(except its basal angles and the lateral borders of the first three
tergites, which are black in this sex) cinnamon-rufous, vinaceous-
rufous, or ferruginous, while the venter is seal-brown. Of the
four females comprised in the same series, one has the dorsum of
the abdomen (except the lateral margins, which are black) dull
amber-brown, while in the case of the other three individuals the
corresponding region is for the most part blackish-brown ; the
venter in each specimen is dark seal-brown. Among the four
examples of P. tabaninus in the possession of the Hope Depart-
ment, University Museum, Oxford, the two males have the
dorsum of the abdomen cinnamon-rufous, apart from the black
lateral borders of the first three tergites, and the basal angles of
the fourth tergite, which are brown or blackish-brown. In one
of the two females in the Oxford collection, the dorsum of the
abdomen, with the exception of the small silvery-white spot near
each posterior angle of the second and third tergites, is uniformly
velvety-black. The corresponding region in the other specimen
is brownish black, with an ochraceous-tawny transverse area in
the centre of the hind border of the first segment, extending on
to the front margin of the second ; on the latter segment there is
also a deep, brownish-tawny, transverse band, which is not very
sharply defined, and does not reach the lateral extremities of the
tergite; on both the second and third tergites, in front of the
hind margin, there is a black transverse band.
It is interesting to note that, in the case of one of the
females of P. tabaninus in the National Collection (the specimen
referred to above as having been taken in Brazil prior to 1849,
by Mrs. Noel), the abdomen shows a very heavy infestation by
nymphs of a Gamasid mite. The pseudo-parasites, which, in the
dried condition at any rate, are light ochraceous-buff in colour,
and measure about 0:4 mm. in length, are present to the number
of many hundreds on the dorsal surface of the abdomen, where
they are packed so closely as to form large patches, while quantities
are to be seen crowded together beneath and partially concealed
by the hind margins of the tergites. Further specimens of the
mite, although in small numbers as compared with those on the
dorsum, are also visible on the venter. The male P. tabaninus
taken by the writer at Macapda, R. Amazons, 25.11.1896, on
DIPTEROUS FAMILY PANTOPHTHALMIDA. 565
board the Cable 8.8. Faraday, shows a considerable number of
what is evidently the same pseudo-parasite on the dorsal surface
of its abdomen, while many instances of similar infestation in
the case of specimens of other species of Pantophthalmidez are
noted below. Since therefore, even among the limited series of
these flies available to the author for examination, representatives
of various species are infested in this manner, it would seem that
for some reason members of this family are peculiarly subject to
to invasion by this Acarid. The author’s colleague, Mr. A. S.
Hirst, F.Z.8., to whom the mites have been submitted, has
kindly supplied the following note:—“ The pseudo-parasites are
nymphs of a speeies of Z’rachytes, a Gamasid mite, and presumably
are predatory forms, possibly feeding upon small mites or insects
inhabiting the tunnel of the fly-larva. This habit of attaching
itself to an insect host is no doubt of benefit to the mite, by
widening its area of distribution.”
Pantophthalmus frauenfeldi Schin.
Acanthomera frauenfeldi Schiner, Reise der dsterreichischen
Fregatte ‘Novara’ um die Erde, Zool. Theil, Bd. ii, Diptera,
p- 78 (1866).
Of this species, the National Collection possesses a 2 from
Bogota, Colombia (ex coll. — Stevens, 1856), in which country the
type was obtained ; and a second 9, from Peru (ea coll. J. M. F.
Bigot; presented by J. HE, Collin). The latter specimen, with
the type of Acanthomera crassipalpis Macq. (= Rhaphiorhynchus
planiventris Wicd.), and two other 2 9 of Rhaphiorhynchus
planiventris Wied., stood in the Bigot collection above the label
“ Acanthomera crassipalpis, Macq., n. sp. 2,” in Macquart’s
handwriting.
Including the facial beak and ovipositor, the original Museum
specimen measures just over 31 mm. in length, instead of 35 mm.,
which is the length of the type (also a 2) as given by Schiner.
The total length of Bigot’s example, measured in the same way,
is 82 mm.
In the Hope Department, University Museum, Oxford, there
is asolitary Q of P. frawenfeldi from Macas, Heuador (purchased
from Stevens, 1862). The dimensions of this specimen, which is
very much larger than the examples in the National Collection,
are as follows:—Total length, including facial beak and ovipositor,
42°25mm.; greatest width of abdomen (across hind margin of
second tergite) 16°25 mm.; wing-expanse 74°6 mm. The terminal
segment of the palpi in the Oxford specimen is slender, straight
and cylindrical, whereas the corresponding segment in the British
Museum @ from Bogota is curved outwards at the tip and
distinctly swollen towards the base; in the @ from the Bigot
collection the palpi are damaged.
Two nymphs of the usual species of Gamasid mite (cf. remarks
under LP. tabaninus above) are visible om the dorsum of the
abdomen of the 9 from Bogota.
37%
566 MAJOR E. E, AUSTEN ON THE
Pantophthalmus batesi, sp.n. (Text-fig. 2.)
3 .—Length (one specimen) 24 mm.; width of head 6:4 mm. ;
greatest width of abdomen (across hind border of second tergite)
8mm.; length of wing 18°5 mm.
Relatively small and (in 3 sex at least) somewhat narrow-bodied
species, with, in 3, a small but conspicuous, sharp-pointed facial
“‘ heak,” wnstriped thorux, and ochraceous-orange abdomen, which
along each lateral margin bears a series of sooty-black spots or
elongate blotches, and dorsal surface of which, except anterior
borders of second to fifth segments inclusive, has from certain angles
the appearance of burnished silver; wing-markings presenting a
general resemblance to those of P. tabaninus Uhunb., but with
conspicuous differences, notably a curved pale mark connecting fork
of third longitudinal vein with costa; posterior femora without,
or with merest vestiye of, a spine on under surface.
Head in § sex mere flattened in an antero-posterior direction
than that of P. tabaninus Thunb., and consequently less hemi-
spherical in ontline when viewed from above; face, except beak,
dusky drab; jowls and basi-ocvipital region clove-brown, clothed
with similarly coloured hair; occiput olive-buff; facial beak
blackish brown at base, distal attenuate portion chestnut-brown
palpi light sepia-coloured, darker towards distal extremities
second segment * clothed beneath, towards distal extremity, with
blackish-brown hair, terminal segment curved, slender, tapering
to the tip; first and second segments of antenne russet-coloured,
first segment thinly clothed with brownish hair, second segment
with some glistening orange-buff hair on distal margin (third
segment missing in case of type). Zhoraw: ground-colour of
dorsum cinnamon-drab, thinly suffused above with smoke-grey
pollen, while lateral borders of dorsum are olive-buff pollinose ;
main portion of dorsum (scutum) marked with five dull-black
spots, of which two, smaller than remainder and nearly circular,
are situated on line of transverse suture, one on each side of
middle line; remaining spots transversely elongate and situated
on hind border, one in middle line, the other two occupying the
postalar calli, except inner extremities of latter; secutellum dull-
black or blackish-brown, anterior angles and lateral borders
smoke-grey pollinose; pleure and pectus blackish-brown and
>
* Note on tho number of segments inthe maxillary palpus in Pantophthalmida.—
Fiebrig (Zeitschr. f. wiss. Insektenbiol., Bd. ii. p. 346, 1906), in describing the
species named by him Acanthomera teretruncum (=Pantophthalmus pictus
(Acanthomera picta) Wied.), speaks of “the relatively long, three-jointed palpi.”
Williston (North American Diptera, 3rd Ed., p. 173, 1908), in a short general
account of the ‘ Family Acanthomeride,” states that the palpi are “ two-jointed.”
Lastly Enderlein (Zool. Anz., Bd. xli. p. 99, 1912) says that in Pantophthalmida,
owing to the irregular transverse wrinkling of the palpus basad of the terminal
segment, it is impossible to determine whether this portion is further seemented or
not. ‘There can be no doubt, however, that the maxillary palpus in this family
consists of three segments, in which, while the second is often marked by con-
spicuous annuli or transverse wrinkles, the first owing to its shortness may easily
be overlooked, although in many species distinguishable by darker coloration and a
different arrangement of the hair.
DIPTEROUS FAMILY PANTOPHTIHALMIDE. 567
clothed with fine similarly coloured hair, dorsum, and lateral
extremities of pronotum, clothed wtih ochraceous-buff hair.
Abdomen: first tergite with each lateral extremity broadly black,
though just in front of hind margin of segment black area shows -
a deep indentation of ground-colour above and below ; second
and third tergites with a scutiform or elongate black spot
occupying each anterior angle, extreme hind margins of posterior
angles of second tergite also black; fourth tergite with anterior
two-thirds of lateral extremities sooty black; lateral extremities
of fifth tergite blackish-brown ; sixth tergite entirely ochraceous-
orange; a straight transverse band running right across fore
border of fifth tergite devoid of silvery sheen; on each of the
three preceding tergites corresponding non-silvered band tapers
towards lateral extremities of the tergite, which it fails to reach,
Text-figure 2.
Ay i
Pantophthalmus batesi Austen, 6. X 14.
while its hind margin, which exhibits a backwardly directed
projection in middle line, is more or less brace-shaped ; lateral
extremities of tergites clothed on dark areas with dark brown or
blackish-brown hair, and elsewhere clothed with yellowish,
cinnamon-coloured, or brownish hair, remainder of dorsum clothed
with minute, scattered, and very inconspicuous cinnamon-buft:
hairs, which on sixth tergite are somewhat longer; venter
orange-buff, partially suffused with a silvery sheen (especially
distinct on first three sternites), and sparsely ciothed with hair
similar in colour to that on dorsum, but longer. Wings light
sepia-coloured, marked as shown in fig. 2, the most conspicuous
markings being those on the costal border, consisting of two
irregularly shaped dark blotches and two cream-buff or pinkish-
buff marks; of the latter, the one nearer the base of the wing,
568 MAJOR E. E. AUSTEN ON THE
which extends from the costa to the upper distal angle of the
second basal cell and includes the anterior transverse vein,
resembles the corresponding mark in the wing of P. tabaninus
Thunb., but is more sharply defined; the veins separating the
second posterior marginal cell from the discal cell, and the
fifth posterior marginal cell from the second basal cell are each
enclosed in a sharply defined, cream-buff spot; a faint, elongate,
cream-buff spot is visible on the sixth longitudinal vein, in the
centre of the lower margin of the anal cell, and both basal cells
ave cream-buff at the base; veins sepia-coloured, except in the
lighter areas where they are cinnamon-buff or pinkish-buff, first
longitudinal vein where enclosed in more distal of the two dark
costal areas, and fifth longitudinal vein except at base and tip
of second basal cell, darker than most other veins or portions
of veins. Malteres cinnamon-buff. Legs: hind femora slightly
incrassate towards distal extremity; femora, front and middle
tibie, and last four segments of all tarsi russet, russet-brown,
or chocolate-brown, hind tibise blackish-brown, first segment of
all tarsi bright mustard-yellow, narrowly russet-brown at extreme
tip, last: segment of all tarsi russet, paler than three preceding
segments; claws russet-brown at base, then blackish-brown or
black; pulvilliand empodia ochraceous-buff or ochraceous-tawny ;
femora, front and middle tibie, and second and two following
segments of all tarsi clothed mainly with russet-brown, dark-
brown, or blackish-brown hair, first segment of all tarsi except
extreme tip clothed with glistening Naples-yellow hair, inner and
flexor surfaces of front tibie, anterior and posterior surfaces
of proximal portion of hind femora, and last segment and
extreme tip of first segment of all tarsi clothed with russet-brown
or chestnut-brown hair, hind tibie fairly thickly clothed with
blackish-brown or black hair, which, except hair on flexor surface
of hind femora, is longer than hair on legs elsewhere.
Brazil: Villa Nova, R. Amazons (4. IW. Bates).
The species described above, which is dedicated to the memory
of its discoverer, the well-known author of ‘'The Naturalist on
the River Amazons,’ is readily distinguishable from any of its
congeners included in the present paper. From P. tabaninus
Thunb., in addition to the points of difference to which special
attention has been shown in the foregoing description, P. bates:
is separable, inter alia, owing to its unstriped thorax, and, at any
rate in the ¢ sex, the silvery sheen on the dorsum of the
abdomen. From the two following species, in both of which, at
least in the ¢, the dorsum of the abdomen is likewise ‘‘silver-
washed,” P. batesi may be distinguished in the same sex, inter
alia, by its abdomen being narrower and spotted with black along
the lateral margins. From P. hellerianus Knderl., of which a
re-description is given below, P. batesi is further distinguishable,
owing to its much more clearly marked wings, to the presence of
dark spots on the thorax, and to the surface of the latter not
being silvery.
DIPTEROUS FAMILY PANLTOPHTHALMIDA. 569
Pantophthalmus hellerianus, Enderl. (‘Text-fig. 3.)
Acanthomera helleriana Enderlein, Zool. Anz., Bd. xliv. no. 13,
p. 581 (July 28, 1914).
Since the original description of this species, which is at
present only known in the male sex, in some respects requires
amplification, P. hellerianus Enderl. is redescribed below. Inthe
ease of the specimen in the National Collection, at any rate,
the brown or greyish-brown, W-shaped mark on the dorsum of
the thorax, on which Enderlein lays stress, is only visible when
the insect is viewed at a fairly low angle from behind, and even
then it is not very distinct. :
$ .—Length (one specimen) 29°5 mm.; width of head 8°4 mm. ;
greatest width of abdomen (across third tergite) 12°2mm.; length
of wing 23-2 mm.
Dorsal surface of both thorax and abdomen, at least in 3,
“washed” with silver, which on thorax is duller, but on abdomen
has a@ more pronounced silken sheen; facial ‘‘ beak” very small
and trconspicuous ; dorsum of thorax (scutum), when viewed from
certain angles, with more or less distinct traces of longitudinal
stripes ; dorsum of abdomen unicolorous, without definite lateral
spots, broad, fiat, and somewhat ovate pyriform in outline when
viewed from above, with sharp edges to third and following tergites ;
wings pale, with (except when viewed against a dark background)
faint, ill-defined markings; hind legs with a small subfemoral
spine.
Head in ¢ distinctly flattened in antero-posterior direction ;
upper portion of face, except immediately adjacent to bases of
antenna, isabella-coloured ; remainder of face, including base of
beak, as also jowls and basioccipital region, dark mummy-brown
or blackish-brown, tiptof beak, which scarcely protrudes beyond
enveloping hood formed by base, chestnut-brown, shining ;
occiput cream-buff pollinose; jowls clothed with brownish or
dark brown hair, with an ochraceous-tawny gleam at base, basi-
occipital region clothed with ochraceous-tawny hair; first and
second segments of palpt greyish cinnamon-brown, clothed with
dark brown hair, terminal segment tawny or cinnamon-rufous,
curved, slender, and tapering; first segment of antennce mummy-
brown, second segment tawny, fringed below with minute, ochra-
ceous-buff hairs (third segment missing in specimen described).
Thoraa: dorsum of main portion (scutum), when viewed at an angle
from behind, exhibiting a pair of admedian, darker (shimmering
greyish-olive), longitudinal stripes, commencing on fore border,
converging and diminishing somewhat in width as they proceed
backwards, and becoming narrower and indistinct after passing
transverse suture; posterior portion of scutum with a similar,
median, longitudinal stripe, commencing in prescutellar groove
and tapering off and disappearing about halfway between latter
and transverse suture; dorsum of scutum clothed with fine
yellowish bair; scutellum, which, except on anterior angles, is
570 MAJOR E, E. AUSTEN ON THE
mainly sepia-coloured, clothed with similar hair; outer extremities
of humeral calli cinnamon-rufous, shining, a ridge of fairly long,
shining, tawny-olive or cinnamon-buff hair on each side of dorsum,
connecting humeral with corresponding postalar callus, postalar
calli mainly mummy-brown, clothed with similarly coloured hair;
pleurz and pectus mummy-brown, clothed with hair of similar
colour, a tuft of bright ochraceous-tawny hair on each side, in
prothoracic region. Abdomen: ground-colour of dorsum russet,
lateral extremities of first tergite broadly, and those of fourth
tergite narrowly sepia-coloured, anterior angles of second and
third tergites similarly tinted; first tergite clothed with fine
cinnamon-buft hair, longer and more conspicuous on lateral
Text-figure 3.
\
hy) 2!
Ai Y
Pantophthalmus hellerianus Enderl., g. X 14.
extremities, where, except on anterior angles, it has an ochra-
ceous-tawny tinge; lateral extremities of second tergite clothed
with fine, short, appressed, cinnamon-buff hair (sepia-coloured
area occupying each anterior angle clothed with shining cin-
namon-brown hair); lateral extremities of third and two following
tergites fringed with cinnamon-brown hair; sixth tergite largely
clothed with similar hair, which is longer on hind border ; venter
uniformly tawny. JVings pale mouse-grey, first costal and sub-
jacent cell mummy-brown or sepia-coloured, two dusky (dusky-
drab), more or less quadrate blotches on costal border, separated
by cream-buff interspaces; proximal blotch including rather less
than distal half of first basal cell, but not extending to tip of
latter; distal blotch, which in specimen described at least: is
the darker of the two, including distal sixth or seventh of first
DIPTEROUS FAMILY PANTOPHTHALMID, 571
longitudinal vein, and area immediately below as far as third
longitudinal vein; tip of wing, including distal portion of first
submarginal and entire second submarginal cell, except its
extreme base, somewhat darker than hind border (when viewed
against a light background), and separated from distal costal
blotch by an ill-defined and irregular pale area; costa ochraceous-
tawny or light cinnamon-brown (mummy-brown at base and
above distal costal blotch), remaining veins cinnamon-brown,
ochraceous-tawny or ochraceous-buff (in darker areas corre-
sponding in tint to adjacent membrane). Haléeres: stalks cinna-
mon-buff, knobs blackish-brown. Zegs: hind femora of uniform
thickness (not incrassate towards distal extremity); femora and
tibiz russet-coloured, clothed with fine, dark brown hair, flexor
surfaces of front tibize clothed with short, glistening, ochraceous-
tawny hair; first segment of front and middle tarsi ochraceous-
buff (cinnamon-brown at extreme tips on flexor surface), first
segment of hind tarsi cream-buff (light cinnamon-brown at
extreme tips on flexor surface), last four segments of all tarsi
ochraceous-tawny, clothed with minute, appressed hairs of similar
colour; claws rufous-tawny ‘at base, then black; pulvilli and
empodia ochraceous-buff.
One g, Brazil: R. Amazons (precise locality unknown) (H. W.
Bates).
Pantophthalmus hellerianus, which, in the ¢ sex at any rate,
agrees with P. argyropastus Big., in having the dorsal surface of
the abdomen silvery, is readily distinguishable from that species,
inter alia, by the dorsum of the main portion of the thorax being
not only much paler but also devoid of dark spots (¢f. notes on
P. argyropastus below). For distinctions from P. batest Austen,
vide supra, p. 568.
Finally it may be added that the example of P. hellerianus in
the National Collection exhibits,.on the dorsal surface of its
abdomen, several specimens of the pseudo-parasitic Acarid
mentioned in the present paper in connection with P. tabaninus
Thunb., and other species; in this case there is also 2 number of
very much smaller mites on the dorsum of the thorax.
Pantophthalmus argyropastus Big.
Megalomyia argyropasta Bigot, Ann. Soc. Ent. Fr., 5° Sér., T. x.,
Bulletin, p. v (1880).
Megalemyia argyropasta Bigot, Ann. Soc. Ent. Fr., 6° Sér., T.1.
pp. 455, 458 (1881).
An examination of the holotype male of this species, formerly
in the Bigot collection and presented to the British Museum
(Natural History) by Mr. J. HE. Collin, F.E.S., shows that, in spite
of Bigot’s misleading statement to the effect that the thorax is
marked with ‘three, somewhat indistinct, blackish bands,” the
dorsum of the main portion of the thorax (seutum) is actually
unicolorous, with scarcely a trace of longitudinal stripes, but
572 MAJOR E, E, AUSTEN ON THE
spotted asin P. batest Austen, that is to say, with a pair of small,
blackish-brown, admedian spots on the transverse suture, a large
seal-brown blotch in front of the scutellum, and the postalar
calli similarly marked; the humeral calli are pale drab-grey
pollinose above, The dorsum of the main portion of the thorax
being dark brown, with a thin, smoke-grey, faintly silvery,
pollinose covering, clothed with fine, cinnamon-buff or drab-
coloured hair, the general appearance of this part of the body
resembles that of moleskin. The dorsum of the abdomen, which
is unicolorous and, when viewed at an angle from behind, appears
dull Mars-brown, has a bright silvery sheen when regarded from
other directions.
Pantophthalmus argyropastus is one of the smaller representa-
tives of its genus, the type, which is from Panama, measuring
only 24°5 mm. in length.
Pantophthalmus versicolor, sp. n. (Text-fig. 4.)
3 .—-Length (three specimens) 27 to 30 mm.; width of head
7:75 to just over 8mm.; greatest width of abdomen (across third
tergite, exclusive of fringe of hair) 12°2 to 14°25 mm.; length of
wing 22°25 to 24:6 mm.
Medium-sized species, with smallish head, which is distinctly
flattened antero-posteriorly, striped thorax, and (at any ratein o)
vid ferruginous or orange-rufous, sharp-edged abdomen ; facial
tubercle or ‘‘beak” in 3 either indistinguishable, its position
marked by a pit, or minute and vestigial; abdominal tergites without
lateral spots, though basal angles of first tergite evtensively, and
those of second tergite narrowly, mummy-brown ; wing-markings
in 3 resembling those in same sex of P. tabaninus Thunb. ; hind
femora of uniform thickness, without a spine on under surface, and
with terminal spine represented only by a blunt projection.
ITead: face and jowls mmmmy-brown, a transverse, horizontal,
smoke-grey, pollinose streak on front margin of jowls connecting
each eye with buccal cavity; occiput greyish-olive, posterior
orbits silvery-white pollinose; jowls clothed with mummy-brown
hair, which is shining ferruginous at base; first and second
segments of palpi clove-brown, clothed, like the terminal seg-
ment except distal extremity of latter, with brownish-ferruginous
hair, terminal segment russet (brownish at base), cylindrical,
straight, bluntly pointed at distal extremity; antenne in ¢
lighter or darker cinnamon-brown (terminal style ochraceous-
tawny), third (compound) segment viewed from side bluntly
lanceolate in outline, second segment midway between base and
distal extremity bearing a circlet of short, sparse, cinnamon-
brown or ochraceous-tawny hairs. Thoraw: ground-colour of
dorsum of scutum mummy-brown or olive-brown ; dorsum smoke-
grey or pale smoke-grey pollinose in front, on humeral calli,
lateral margins, lateral expansions of transverse suture, and on
postalar calli; central region of dorsum bearing three conspicuous,
dark, converging, longitudinal stripes (approximately equal in
DIPLEROUS FAMILY PANTOPHTHALMIDA. 573
breadth, except that outer stripes are usually somewhat broader
in front), extending from front to hind margin, but confluent
posteriorly after passing transverse suture, outer stripes black or
blackish-brown, median stripe paler, dark mummy-brown, some-
times less noticeable than outer stripes; behind transverse suture
a short blackish-brown or dark brown longitudinal stripe on each
side, extending from suture to postalar callus, and on inner side
of this stripe a somewhat oblique, shining, chestrut-brown stripe ;
dorsum, except main portion of upper surface of scutellum, which
is bare, thickly clothed with fine, silky, glistening, reddish-brown,
ochraceous-tawny or tawny-olive hair, which on lateral borders is
longer than elsewhere; scutellum mummy-brown, dark seal-brown,
or russet-brown, more or less distinctly shimmering silvery-white
Text-figure 4.
Pantophthalnus versicolor Austen, 6. X13.
or shimmering smoke-grey pollinose above, except on hind border;
pleurse and pectus chocolate-brown or warm sepia-coloured,
clothed with fine and fairly long, dark brown or reddish-brown
hair. Abdomen: second and two following tergites each with
a reddish-brown transverse band, just in front of hind margin
and widely separated from lateral extremities in each case, pos-
terior half of fifth tergite with an ill-defined blotch of similar
colour in centre, not reaching hind margin; first tergite thinly
clothed with fine hair, which is ochreous and recumbent except
on basal angles where it is brownish and erect; second and three
following tergites sparsely clothed with minute, appressed, glis-
tening, ochraceous-tawny hairs, which in the case of second and
574 MAJOR E. E, AUSTEN ON THE
third tergites at any rate are only noticeable’ on close scrutiny,
sixth tergite clothed with longer hair of similar colour; lateral
extremities of second tergite, except posterior angles, clothed
with dark brown or brownish hair, those of third and fourth
tergites fringed with longer, outstanding, closely set, shining,
cinnamon-rufous or russet-tawny hair, lateral extremities of fifth
tergite with a shorter, thinner, and paler fringe of similar char-
acter; first sternite mummy-brown or sepia-coloured (hind border
smoke-grey or drab-grey pollinose), clothed with fine and fairly
long, erect, dark brown hair; remaining sternites sepia-coloured,
lateral extremities of third and following sternites paler (ochra-
ceous-tawny); second and two following sternites (except lateral
extremities of third and fourth), andl central portion of fifth
sternite thinly clothed with fine, dark brown hair, which on
second sternite, at least anterior ly, i is longer and erect, but else-
where is shorter and appressed; sixth sternite, and lateral
extremities of the two preceding sternites, clothed with russet-
tawny hair; membrane connecting sternites with corresponding
tergites longitudinally wrinkled, dark brown in ease of first seg~
ment, and in case of succeeding segments dark brown in front
and ochraceous-tawny behind. Wings sepia-coloured, with pale
(cream-buff) markings as shown in: text-fig. 4, these markings
occupying corresponding positions to those in wing of P. tabaninus
Thunb., but somewhat more restricted in extent; darkest areas
in wing are those which are usual in this family, namely first
costal cell and cell immediately below, second costal cell except
its distal extremity, first basal cell except its base and tip, and a
smaller and somewhat ill-defined area including stigmatic region,
extending from costa to distal half or three-fourths of discal cell,
with an outward extension along third longitudinal vein; veins
for most part ferruginous or mummy-brown, paler in pale areas.
Squame blackish-brown, fringed with similarly coloured hair,
and also with a tuft of shining tawny or russet hair. J/alteres:
stalks ochraceous-buff or ochraceous-tawny, knobs mummy-
brown, Legs uniformly russet or chestnut-brown, tarsi some-
times paler (cinnamon-rufous); legs clothed with glistening
hair of similar colour, long and fine on front and hind femora
short elsewhere, hair on femora sometimes appearing dark brown,
sometimes tawny; claws orange-cinnamon or russet, their distal
extremities black; pulvilli and empodia ochraceous-buff or
orange-buff.
Guatemala and Mexico. Holotype, Pancina, Vera Paz,
Guatemala (G. C. Champion); two paratypes, Cuesta de Misantla,
Mexico (M. Zrujillo). The holotype is the specimen briefly
described, but not named, by Osten Sacken (Biol. Centr.-Amer.,
Dipt. i. p. 68).
Through the kindness of Mr. J. . Collin, the British Museum
(Natural History) has received a solitary ¢ of this species, which
unfortunately is in poor condition and is without a locality
label.- In the Bigot collection, whence it was taken, this fly was
DIPTEROUS FAMILY PANTOPHTHALMIDA, 575
associated with two ¢ o of P. tabaninus Thunb. and all three spe-
cimens stood above the label ‘“‘ Acanthomera seticornis Wied. 3 ,”
in Macquart’s handwriting (vide supra, p. 563).
In the male sex, at any rate, Pantophthalmus versicolor, by
means of the characters mentioned in the diagnosis printed in
italics above, is readily distinguishable from any of its congeners
at present known. From the male of P. tabaninus 'Thunb., for
which at the first glance it might possibly be mistaken, the male
of the present species can he distinguished at once owing to its
smaller and differently shaped head, duskier and differently
striped thorax, and broader and flatter, sharp-edged abdomen,
the sides of which are devoid of white spots, and are not black on
the basal half.
Pantophthalmus pictus Wied.
Acanthomera picta Wiedemann, Diptera Exotica, p. 61, Tab. ii.
fig. 2 (1821).
Acanthomera flavipes Macquart, Mém. Soc. roy. des Sc., de
YAgric. et des Arts de Lille, Année 1846, p. 42, Tab. i. fig. 2
(1847). [New synonym. ]
Acanthomera magnifica Walker, Ins. Saund., Diptera, Part i.
p. 74, pl. 1. fig. 1 (1850). [New synonym. |
Acanthomera teretruncum Fiebrig, Zeitschr. f. wiss. Insekten-
biol., Bd. ii. pp. 345-347, figs. 17-19 (1906).
Of this species, which is the genotype of Acanthomera Wied.,
the British Museum (Natural History) possesses a ¢ from an
unknown locality in Brazil (purchased from — Argent, 1847) ;
a1 2 (the holotype of Acanthomera magnifica Walk.), from
“South America ” (ea coll. W. W. Saunders); and a second @,
taken at Sapucay, Paraguay, 21.i1.1905 (W. Foster). ‘The type of
Acanthomera magnifica Walk. represents a form of the species in
which the dorsum of the abdomen is for the most part ferru-
ginous, tawny, or orange-cinnamon instead of black, the latter
colour being confined to the base (first visible tergite, except its
hind border), ovipositor, and a blotch at each lateral extremity
of the second to the fourth tergites inclusive. Jn the case of the
type of A. magnifica, moreover, the whitish transverse streak
near each posterior angle of the third ventral scute is wanting,
but this is doubtless due to individual variation.
From the Bigot collection, thanks to the generosity of Mr. J. KE.
Collin, the National series has been enriched by the type of
Acanthomera flavipes Macq. (stated by Macquart-—loc. cit.—to be
from Brazil), and two other Q @, all of which unfortunately are
without locality labels. In the case of Macquart’s type the
abdomen is russet-brown (“ brun vineux,” apud Macquart, loc.
cit.), a fact which misled Enderlein (Zool. Anz., Babel peel Ok
(1912)) into treating A. flavipes Macq. as specifically distinct from
A, pictw Wied. “Enderlein, however (¢. cit. p. 105), is perfectly
correct in regarding A. teretruncum Fiebrig as a synonym of
576 MAJOR E. E. AUSTEN ON THE
A. picta Wied., and indeed, so long ago as 1908, the present
writer placed a manuscript note in the Museum collection
-drawing attention to this identity.
The species under discussion is represented in the Hope Depart-
ment of the Oxford University Museum by two @ 9, one of ©
which is of ordinary size (total length, including ovipositor and
facial beak, 41:2 mm.; wing-expanse 73 mm.), while the other
is a diminutive example, with dimensions (total length, including
ovipositor and facial beak, 29 mm,: wing-expanse 50°5 mm.) much
below normal. Both specimens are unfortunately without locality
labels; the larger of the two is labelled ‘‘ Miers coll.,” while the
other, probably collected by Bates on the Amazons, bears a label
in Westwood’s handwriting, with the words “ i Mus. Saunders.
1867.”
In the wing-marking of Pantophthalmus pictus « characteristic
feature is a broaddark brown, E-shaped blotch, the shorter arm
of which meets the costa in such a way as to include the tip of
the mediastinal vein, while the longer arm embraces the third
longitudinal vein between the anterior transverse vein and the
bifurcation of the former, without being in contact with either.
Fiebrig’s interesting observations on the preliminary stages
and bionomics of ‘A, éeretruncwn” have already been referred
to (vide supra, p. 056). .
Pantophthalmus chuni Endezl.
Acanthomera chunt Enderlein, Zool. Anz., Bd. xli.’ no. 3,
pp. 102, 103, fig. 4 (December 20, 1912).
This fine species—described by its author from two 2 9 from
Peru, in the Zoological Museum, Stettin—is represented in the
National Collection by a single 2 from Brazil (R. Amazons—
precise locality unknown: 7. I. Bates), measuring 42 mm. in
length from the tip of the prominent facial beak to the end of
the ovipositor, and having a wing-expanse of 70°6 mm. On the
second and third abdominal tergites, conspicuous against the
velvety-black or sooty-black ground-colour, are half-a-dozen
specimens of the Acarid parasite already noticed in connection
with several other species.
By the courtesy of Dr. Hugh Scott, the writer has been per-
mitted to examine a 9 of this species in the University Museum
of Zoology, Cambridge. Except that its dimensions (length from
tip of facial beak to end of ovipositor 29 mm., wing-expanse
53°75 mm.) are much below normal, this example, which was
likewise taken on the River Amazons (Nevile Goodman, 1879),
agrees with that already mentioned.
In spite of certain apparent discrepancies, there can be no
doubt as to the correctness of the identification of the Brazilian
specimens referred to above. ‘lhe description of P. (Acanthomera)
chuni furnished by Enderlein is seriously misleading in more than
one respect. In the first place the German author speaks of the
DIPTEROUS FAMILY PANTOPHTHALMIDA. HTT
face as being “very broad and very greatly widened in front”;
a comparison, however, of the British Museum specimen with
examples of other species shows that the 9 of P. chuni is in no
way abnormal in this respect. Secondly, both in the diagnostic
table on p. 102 (loc. cié.) and in the detailed description on the
following page, Enderlein speaks of a single, relatively narrow
thoracic stripe. On the other hand, in actual fact, not only does
the dorsum of the thorax exhibit the three dark thoracic stripes
so constantly seen in representatives of the family Pantoph-
thalmide, but, even regarding the thoracic markings as con-
sisting of a single stripe, bearing on its surface a pair of paler,
admedian stripes, the measurements given by Enderlein himself,
which agree in detail with the corresponding measurements taken
from the British Museum specimen, show that as a characterization
“relatively narrow” is entirely unwarranted.
Pantophthalmus conspicabilis, sp. n. (Text-fig. 5.)
9 .—Length (one specimen), including ovipositor and facial
beak, 42 mm.; width of head 10°5 mm.; width of front at vertex
1:8 mm. (immediately above base of antenne just under 2 mm.);
length of facial beak just under 3 mm.; greatest width of
abdomen (across third tergite) just under 16 mm.; length of wing
31:5 mm.
In 2 sex, at least as represented by typical specimen, a grey and
black species, with prominent, sharp-pointed facial beak ; almost
parallel-sided front; conspicuously striped thorax; blunt-edged
abdomen, which on dorsal surface, on each lateral border, bears two
conspicuous silvery-white spots and a fringe of short hair; wings
marbled with sepia and light buff, somewhat resembling those of
Pantophthalmus (Acanthomera) chuni Hnderl., but differing in
details of pattern; and with hind femora incrassate towards tips,
and bearing a small spine on under surface and a large terminal
spine.
Head: face and jowls sepia-coloured or mummy-brown, inner
margin of each eye bounding face with a narrow, smoke-grey,
pollinose edging, which above sends off an extension to meet base
of facial beak, while below a similar but silvery-grey extension
connects pollinose edging of eye with buccal cavity; facial beak
long and sharp-pointed, its expanded basal portion dark brown
above, russet-brown on sides below, the shining tip blackish-
brown; front (frons) of moderate width, and only very slightly
broader at its anterior extremity than at vertex; front and
occiput, including posterior orbits, uniformly olive-buff or pale
olive-buff pollinose, except that the conspicuous ocellar tubercle,
which is distinctly separated from vertical margin of occiput,
is mummy-brown, and that in certain aspects, at least. in
typical speciinen, behind ocellar tubercle, on vertical margin
of occiput, and also on central anterior portion of front,
above base of antenns, ground-colour shows through pollinose
578 MAJOR E. E, AUSTEN ON THE
covering, and appears more or less tawny or ochraceous-salmon-
coloured; upper portion of occiput, except posterior orbits,
clothed with fine, ochraceous-buff hair, basioccipital region and
lower portion of occiput, except posterior orbits, clothed, like
jowls, with fine, brownish or dark brown hair; first and second
segments of palpi dull tawny-olive (second segment dark brown
shortly before distal extremity), clothed on sides and below with
blackish hair which is longer towards tip of second segment,
distal segment russet-coloured with a thin greyish-pollinose
covering, cylindrical and acuminate, considerably thicker than
base of proximal segment, but less swollen than is typically the
case in the genus Rhaphiorhynchus, distal segment at base and on
outer surface bearing a few short black hairs; antennw in &
5-6 mm, in length in case of type, agreeing generally in shape,
and in form and relative dimensions of the eight divisions of the
third (compound) segment, with antenna of Pantophthalmus
(Acanthomera) chuni Vinnderl., 2, as figured (Zool. Anz., Bd. xi.
no. 3, p. 104,-fig. 4, December 20, 1912) by its author; first and
second segments of antennz mummy-brown, distal extremity of
second segment sparsely fringed with short, ochreous hairs,
proximal division of third segment mummy-brown, next six
divisions tawny, terminal (elongate and attenuate) division paler
pinkish-cinnamon), entire antenne, when viewed from certain
angles, clothed with a thin, silvery-grey, pollinose covering.
Thorax: dorsum of main portion (scutum) silvery-grey (a deep
border on each side, extending from humeral to postalar calli
inclusive, silvery), median area in front of transverse suture when
viewed from certain angles tinged with pinkish-cinnamon (at
least in case of type), owing to ground-colour showing through ;
from inner end of each humeral callus a conspicuous, blackish-
brown longitudinal stripe extends backwards, each stripe being
fairly broad in front and diminishing in width as it passes back,
until at a point about one-third of the distance between trans-
verse suture and prescutellar groove it becomes suddenly attenuate,
and is thence continued as a narrow extension as far as the pre-
scutellar groove, in front of which it is connected with its fellow ;
attenuate portion of blackish-brown longitudinal stripes ‘scarcely
noticeable from certain angles, and completely visible only when
insect is viewed at a low angle from behind; in middle line,
between the two blackish-brown stripes, is an incomplete, paler
(brownish-drab) longitudinal stripe, which, commencing as a
deltoid expansion resting upon posterior commissure between the
attenuate ends of the blackish-brown stripes, extends forwards,
disappearing (in case of type, at least) before reaching trans-
verse suture, but reappearing on level of latter, and tapering to
a point and terminating between humeral calli; on upper
margin of Jateral silvery border on each side, between inner
extremities of humeral and postalar calli, is a narrow, longi-
tudinal brownish streak, which, following configuration of
thorax, curves outwards in front of transverse suture, and makes
DIPTEROUS FAMILY PANTOPHTHALMIDA. 579
a similar but rather flatter curve behind it; scutellum mummy-
brown (posterior border, which is thickly covered with small,
polished, rounded tubercles, mainly blackish-brown), lateral
extremities of dorsal surface silvery-grey; dorsum of scutum
clothed with fine, short, silky, yellowish or brownish hair, which
becomes longer on hind border; scutellum, except anterior half
of dorsal surface, which is bare, clothed with short, dark brown
or blackish-brown hair; pleure and pectus mummy-brown
(appearing darker when viewed from certain angles), clothed
with fine, dark brown hair. Abdomen (at least in case of type,
and except as regards the lateral silvery-white spots) uniformly
blackish-brown or brownish-black above and below (ventral
surface perhaps rather more brownish than dorsal); second and
Text-figure 5.
FC
Pantophthalmus conspicabilis Austen, 9. x 1}.
third tergites each with a conspicuous, silvery-white spot on each
side, close to posterior angle; lateral borders of dorsum clothed
with blackish-brown hair, which on second and two following
tergites forms a not very conspicuous fringe on each side (on
lateral margins of fourth tergite the fringe is longer and sparser
than on the two preceding tergites); first sternite, and central
area of second, clothed with fine, dark brownish hair. Wings
marked as shown in text-fig. 5, the dark blotches on costal border
sharply defined and conspicuous; veins sepia-coloured or
cinnamon-brown, in pale areas usually paler ( pinkish-cinnamon).
Halteres: stalks cinnamon-brown, knobs blackish-brown. Legs :
front femora with a small but distinct terminal Spine, which on
Proc. Zoou. Soc.—1923, No. XX XVIII. 38
580 MAJOR E. E. AUSTEN ON THE
middle femora is represented by a blunt triangular projection ;
hind legs with a very small, somewhat blunt and triangular, sub-
femoral spine; terminal spine on hind femora large, long (1 mm.
in length) and sharp; length of hind femur (exclusive of terminal
spine) 15:6 mm.; hind femora distinctly thickened towards distal
extremity, clothed with fine, short, blackish hair, which is longer
on flexor surface and, owing to greater density, especially notice-
able between subfemoral spine and tip, distal extremities of
extensor and posterior surfaces clothed mainly with bright,
appressed, russet-coloured hair; hind tibie thickly clothed with
fine, black hair, longer than elsewhere on extensor surface, where
it forms a conspicuous fringe; femora reddish-brown, darker
towards distal extremities, tips of hind pair reddish-brown ; front
tibie russet-coloured, clothed on flexor surface with short,
glistening, ochraceous-tawny hair; middle tibize russet-brown,
clothed for most part with dark brown hairs; hind tibie black ;
first segment of tarsi buff-yellow, extreme tip brown; remaining
segments of front tarsi sepia-coloured, second segment, except
distal third, and extreme bases of following segments cinnamon-
buff, last three segments of front tarsi clothed above with dark
brown hairs, tip of last segment and distal angles of preceding
segments clothed with bright ochraceous-tawny hairs; second
segment of middle tarsi cinnamon-brown, darker at distal
extremity, last three segments of middle tarsi dark brown, clothed
above with similarly coloured hair, tip of last segment clothed
with tawny hair; last four segments of hind tarsi blackish-brown,
second segment paler at base; claws orange-cinnamon or russet,
their tips black; pulvilli and empodia ochraceous-tawny.
Colombia (— Chevrolat): ex coll. W. W. Saunders.
In the @ sex, at any rate, the handsome species described
above, although at the first glance presenting a certain superficial
resemblance to Pantophthalmus (Acanthomera) chuni Ender.
(vide supra), is distinguishable inter alia owing to the pallor of
the front (frons) and of the dorsum of the thorax; the reduction
in the striping of the latter as regards the median longitudinal
stripe and the distal extremities of the paired dark stripes; and
the shorter, much more densely hairy and consequently apparently
stouter hind tibie. In the case of the type, at least, of P. con-
spicabilis, as compared with the specimen of P. chuni referred to
on p. 576, the lateral silvery-white spots on the dorsum of the
abdomen are blunter and rounder.
This species is in any case closely allied to, and may ultimately
prove to be identical with, Pantophthalmus kertészianus (Acan-
thomera kertésziana) Enderlein, described (Zool. Anz., Bd. xliv.
no. 13, p. 578, July 28, 1914) from two 2 2 from Peru, in the
Museums of Budapest and Stettin. Since, however, Hnderlein
(é. cit. p. 579), in describing the wing in P. kertészianus, says
that the ‘“‘ basal fourth and a large spot on the anterior margin”
are “pale ochreous,” it seems wiser, provisionally at any rate, to
regard P. conspicabilis as distinct.
DIPTEROUS FAMILY PANTOPHTHALMID&. 581
Pantophthalmus conspicuus, sp.n. (Text-figs. 6, 7.)
9 .—Length (one specimen), including ovipositor and facial
beak, 36°2 mm.; width of head 9°4mm.; width of front at vertex
1°38 mm.; greatest width of abdomen (across third tergite)
14°5 mm.; length of wing 28 mm.
In & sex, at least in case. of typical specimen, a grey, honey-
yellow or isabella-colowred (probably in life green) and black
species, with facial beak represented by a blunt, rounded prominence,
conspicuously black-striped thorax, blunt-edged abdomen, and no
subfemoral spine on hind legs; lateral borders of first four
abdominal tergites black, with two large, silvery-white spots on each
side; wings marked as shown in teat-fig. 6, broadly cream-buff or
buff-yellow at base; hind femora somewhat incrassate towards tips,
hind tibiee thickly clothed with black hair, appearing stout.
Head: front (frons) light greyish-olive pollinose, of moderate
and uniform width, ocellar tubercle black; face and jowls
mumimy-brown, tip of bluntly rounded facial beak reddish, inner
and lower orbits bordering face:and jowls narrowly olive-buft
»ollinose, jowls and basioccipital region «clothed with dark brown
or blackish-brown hair; occiput dark olive-buff pollinose, appear-
ing butfy-brown in certain areas and mummy-brown below
vertex, clothed with fine, brownish hair, which has an ochraceous-
tawny or cinnamon-brown sheen, posterior orbits light buff
pollinose, bare; first and second segments of palpi sepia-coloured,
together shorter than distal seginent and clothed with dark
brown or blackish-brown hair, distal segment russet-coloured,
cylindrical, tapering somewhat towards ‘tip and ending bluntly,
with a few blackish hairs on inner side towards base; antennce
thinly olive-buff or smoke-grey pollinose, first and second segments
tawny, distal extremity of second segment fringed with bright
ochraceous-tawny hairs, third (compound) segment russet-brown,
terminal division paler (cinnamon- or pinkish-cinnamon-coloured).
Thorax: dorsum of main portion (scutum) pale smoke-grey pol-
linose (ground-colour largely chestnut-brown or russet, ground-
colour of humeral calli pinkish-buff), with two-straight and
complete, converging, longitudinal black stripes of uniform width,
which in front are in contact with inner ends of humeral calli,
and posteriorly meet together in prescutellar groove; in middle
line a brownish, dusky stripe (visible at least when insect is
regarded at an angle of 45° from behind), commencing on front
margin as an elongate black spot, extends backwards and becomes
indistinct shortly after passing level of transverse suture ;
scutellum, except lateral extremities, which are whitish-grey
pollinose and clothed with pale hair, blackish-brown, clothed
above with short blackish hair, and below with brownish hair:
dorsum of scutum clothed in front with short yellowish hair, and
posteriorly with short blackish or black hair, on each side above
pleure and in front of transverse suture an area of longer, dark
brown or brownish hair, posteriorly ochraceous-tawny at base;
38*
582, MAJOR E. &. AUSTEN ON THE
pleure and upper border of pectus black, clothed with black or
blackish hair, central area of pectus light greyish-olive pollinose
(ground-colour chestnut-brown), clothed with fine, cinnamon-buff-
coloured hair. Abdomen; first four tergites, at least in case of
type, honey-yellow or isabella-coloured, here and there irregularly
mottled with dark green or greyish-olive, their lateral extremities
black, fifth tergite dark olive, rather less than posterior half. of
preceding tergite similarly coloured in centre, second and third
tergites each with a large, bluntly triangular, silvery-white spot
in each posterior angle, not quite reaching hind margin in either
case, first tergite with a pair of very small whitish spots in
corresponding position but further from hind margin, ovipositor
black or blackish-brown (distal extremity cinnamon-coloured) ;
Text-figure 6.
rei
Pica
THRZI-X
Pantophthalimus conspicuus Austen, 9. x Il,
lateral borders of first four tergites, except silvery-white spots,
clothed with fine, short, black hair, which, though longer on outer
edges, does not form a conspicuous fringe; first four tergites,
except lateral extremities, sparsely clothed with short, or minute
and inconspicuous, ochraceous-buff or ochraceous-orange hairs;
first four sternites clove-brown (their lateral extremities blackish-
brown), with a shimmering, smoke-grey, pollinose covering and
sparsely clothed with short, ochraceous-orange hairs, ventral
surface of projecting lateral extremities of first four tergites
black, fifth sternite blackish-brown, sparsely clothed with
similatly coloured hair, coloration of ventral surface of ovi positor
same as that of dorsal. Wings: infuscated (blackish-brown) area
chiefly confined to distal third and posterior two-thirds, but first
DIPIEROUS FAMILY PANYTOPHTHALMID”. 583
basal cell with an elongate dusky blotch occupying part of its
distal two-thirds, but not extending either to tip or to hind
border of the cell, an elongate brownish streak in second costal
cell above foregoing blotch, a large pale spot on fork of third
longitudinal vein, and a more or less distinct pale streak along
costa at tip of marginal cell; infuscated area in discal cell not in
contact with boundaries of cell at any point except on anterior
basal transverse vein, across centre of which it is connected
with large infuscated area in second basal cell, which again
is continuous with infuscated axillary cell and alula, through
infuscation occupying greater part of anal cell; veins mummy-
_ brown or cinnamon-brown. J/alteres: stalks tawny- -olive, knobs
dusky dull green or dusky olive-green. Legs: coxe and tro-
chanters tawny or russet-coloured, clothed mainly with tawny or
ochreous hairs, front coxze on outer side blackish and clothed with
similarly coloured hair; front and middle femora with a bluntly
triangular projection at distal extremity, terminal spine on hind
femora not unusually large; front and middle femora chestnut-
brown, with an ill-defined dark brown band before distal ex-
tremity, hind femora similarly coloured but darker, femora
clothed mainly with dark brown or blackish hair, front and
middle femora sparsely clothed at base on under side with short
tawny hair, tips of all femora, at least above, and base of hind
femora, clothed with glistening cinnamon-rufous hair, par-
ticularly noticeable at tips of hind femora on extensor surface,
where it is short and closely appressed ; front tibiee russet, clothed
largely with short, glistening tawny or cinnamon-rufous hairs,
with longer and sparser dark brown hairs on extensor surface,
middle tibize chestnut-brown (darker on anterior surface), extensor
surface sparsely clothed with dark brown hair, flexor surface and
tips, which are paler, clothed with short, glistening, appressed,
cinnamon-rufous hair; hind tibie black (extreme tips paler and
clothed with ochraceous- -tawny hair); first three segments of front
and middle tarsi, except tips, ochraceous-buff, last two segments
and tips of preceding segments cinnamon-brown; front and middle
tarsi entirely clothed with glistening, appressed, ochraceous-
tawny hairs, longer at distal extremity of upper surface of ter-
minal segment, minute or short elsewhere; first segment of hind
tarsi warm buff or cinnamon-buff (extreme tip faintly cinnamon-
brown), clothed with short, glistening , appressed, cream-buff hair;
last four segments of hind tarsi cinnamon-brown, clothed (as
also extreme tip of first segment) with glistening, appressed,
ochraceous-tawny hairs; claws tawny or russet-coloured, their
tips black; pulvilli and empodia ochraceous-tawny.
British Honduras: Corozal, 26. ix. 1922, “came to light, at
night” (Dr. #. L. Davis). The collector of the type, who has
been kind enough to present the specimen to the National
Collection, writes that, although resident in British Honduras
-for very many years, he cannot. remember having met with ‘las
species previously. "
584 : MAJOR E. E. AUSTEN ON THE
Apart from the coloration of the: abdomen, as exhibited in the
case of the typical specimen at least, Pantophihalmus conspicuws
in the 2 sex exhibits a certain superficial resemblance both to
P. chuni Enderl. and P. conspicabilis Austen. From both of
these species it is, however, distinguishable inter alia owing to
the wings being broadly pale (cream-buff or buff- yellow) at the
base, and to the facial beak being bluntly rounded at the tip
instead of acuminate. From P. chuni, apart from any other
characters, it is further distinguished by its stout, furry hind
tibiz, and pale front (frons). Cees
The type of P. conspicwus was originally preserved and sent to
the Museum in spirit, and there is reason to believe that, owing |
either to this, or to natural post-mortem changes, the coloration
of the disc of the abdomen (exclusive of the lateral borders) has
suffered alteration, and that it was originally green. ‘This at any
rate seems a natural deduction from the fact that traces of green
are visible here and there in the first four abdominal tergites,
and that the halteres are still more or less green (vide descrip-
tion above). In any case, this is not the only instance in which
a representative of this family is more or less green in life, since,
according to the late Prof. Hermann (Deutsch. Ent, Z., Jahrg.
1916, Heft i. pp. 43,.45, April 1, 1916), the Brazilian Pant-
ophthalmus alienus Herm., in the female sex at any rate, in
which the ground-colour of the abdomen is dull velvety-black,
has the hind margins of the first four tergites apple-green, while
there are indications that in life the scutelluin and halteres are
of the same colour. It is of course well known that in those
instances among Diptera—highly exceptional if the Chironomid
be excluded—in which the predominant colour in life is green,
the green pigment after death is often or usually, largely or
entirely, replaced by yellow. As cases in point, it will suflice
to mention Oxycera trilineata Fabr., and other Stratiomyide,
such as the South American Odontomyia pulchra Wied.; Tabanus
fasciatus Fabr., of West and Central Africa; and the Kuropean
Didea alneti Elin.
What is almost certainly the male sex of P. conspicwus is
represented bya solitary ¢ (cf. text-fig. 7) from the R. Amazons,
Brazil (HZ. W. Bates), in the Hope Department, University
Museum, Oxford (ea coll. W. W. Saunders)., The dimensions of
this specimen are as follows :—Length 29°6 mm.; width of head
just over 8 mm.; greatest width of abdomen (across hind margin
of second tergite) 9°-4 mm.; length of wing 24:'4mm. Viewed
from above and slightly from in front, the dorsum of the thorax,
except the hind border of the scutellum, is unstriped and
uniformly silvery, forming a striking colour-contrast with the
dorsum of the abdomen, which is ochraceous-tawny ; the lateral
extremities of the first four abdominal tergites are black and
deflexed, while the posterior angles of the second and third
tergites are each occupied by a large, bluntly triangular, silvery-
white spot, lateral borders and spots being scarcely noticeable
DIPTEROUS FAMILY PANTOPHTHALMIDS. 585
from above. ‘The other principal external characters are as fol-
lows:—MHead: frontal and vertical triangles smoke-grey pollinose,
ocellar tubercle dark brown ; face and jowls sepia-coloured, jowls
clothed with hair of similar hue, inner orbits bordering face
narrowly pinkish-buff pollinose ; facial beak represented by a low,
elongate oval, tawny-brown area, which is scarcely noticeable ;
occiput shimmering golden-ochraceous-buff pollinose, with darker
reflections, and clothed with fine yellowish hair; palpi similar
in colour to those of 9, but distal segment considerably (about
one-third) shorter than second; antenne sepia-coloured, style
paler, third (compound) segment cuneate, flattened from side to
side, and deepest (in a dorso-ventral direction) at about one-
fourth of its length from the base. Thorax: rather less than
Text-figure 7.
ff
Presumptive ¢ of Pantophthalmus conspicuus Austen. X13. (From a specimen
in the Hope Department, University Museum, Oxford).
posterior half of seutellum blackish-brown, clothed with brownish
hair, dividing line between infuscated and silvery areas straight ;
dorsum of scutum, except lateral borders immediately above
pleure, clothed with fine, short, reflexed, whitish hair, lateral
borders clothed with longer hair, light ochraceous buff in front
of, and yellowish behind transverse suture, under side of
postalar calli blackish-brown; viewed at a slight angle from
behind, dorsum of scutum shows unpigmented longitudinal
stripes, corresponding to the paired black stripes and commence-
ment of a median stripe exhibited by the 2 ; pleure and lateral
borders of pectus dark clove-brown or blackish-brown, clothed
with similarly coloured hair, remainder of pectus light ochraceous-
buff, clothed with fine, silky hair of similar colour. Abdomen:
586 MAJOR E. E. AUSTEN ON THE
venter ochraceous-buff, pinkish-buff pollinose, sparsely clothed
with fine, glistening ochreous hairs, lateral extremities of sternites
brownish ; dorsum, except lateral borders of first four segments,
clothed with ochreous or ochraceous-buff hairs, for most part
minute and inconspicuous except on first and sixth segments,
lateral borders of first four tergites, except silvery-white spots,
clothed with black or blackish hair. Wings (cp. text-figs. 6 & 7)
with markings corresponding extremely closely to those exhibited
by °, but tips paler. Halteres cinnamon-buff. Legs agreeing
in all essential respects with those of @ ; coloration of hind coxe
and their hairy covering as in case of adjacent area of pectus.
Pantophthalmus vittatus Wied.
Acanthomera vittata Wiedemann, Auss. zweifl. Ins. i. p. 109
(1828).
Acanthomera fulvida Bigot, Ann. Soc. Ent. Fr., 5¢ Sér., T. x.,
Bulletin, p. v (1880). [New synonym. ]
A note in the National Collection in the handwriting of the
late Baron C. R. Osten Sacken, and bearing the initials “O. 8.,”
confirms the correctness of the identification of P. vittatus
(Acanthomera vittata) Wied, as given above.
The following are the specimens of this species in the British
Museum: a 6d, from the R. Amazons, Brazil (HZ. W. Bates) ;
a 9, also from Brazil, but exact provenance unknown, taken
prior to 1849 (Mfrs, Noel); and two 2 Q (the type and paratype
of Acanthomera fulvida Big.), from Guiana (ex coll. J. Bigot:
presented by J. HK. Collin, F.E.S.). _
The two latter examples represent a form of P. vittatus in
which the dark lateral spots on the dorsum of the abdomen are
equally well developed in both sexes. Bigot himself (loc. cit.)
mentions only a single specimen, but, since the alternative
measurements given by this author agree with the respective
lengths of the 9 9 referred to, his remark “‘ Specimen wnicum”
is probably a lapsus calamt.
There are six specimens of P. vittatus—all 9 9 —in the Hope
Department of the University Museum, Oxford. Of these, three
are simply labelled “ Miers coll.”, and bear no further indication
of provenance ; of the remainder, one has a label with the words
“S$. S. Saunders, Bahia, Bz.”; another is labelled “‘H Mus.
Saunders, 1867”; and a label attached to the sixth example
shows that this specimen was taken in Brazil, and purchased
from Higgins in 1871. Considerable variation in size is notice-
able in the Oxford series; while the dimensions of the largest
@ (total length, including ovipositor and short facial beak,
32:25 mm.; wing-expanse 57 mm.) are approximately the same
as those of Mrs. Noel’s specimen in the National Collection, the
corresponding measurements of the smallest example in the
possession of the Hope Department are 24-6 mm. and 40 mm.
respectively—practically the same as those of the smaller of the
DIPTEROUS FAMILY PANTOPHTHALMIDA. 587
two 2° 2 from the Bigot collection, presented to the British
Museum by Mr. Collin.
The g and @ in the British Museum, collected respectively
by H. W. Bates and Mrs. Noel, are each pseudo-parasitized by
nymphs of the usual species of Gamasid mite. In the case of
the ¢, there is only a single Acarid, which is on the upper
surface of the abdomen; the pseudo-parasites on the 2 are
present on both sides of the abdomen, and, although only in
small numbers, are more plentiful above than below. One of the
specimens in the Oxford collection is similarly infested.
Pantophthalmus bellardii Bell.
Acanthomera bellardii Bellardi, Saggio di Ditterol. Messicana,
Appendice, p. 16, Tav. ili. fig. 11 (1862).
Acanthomera championi Osten Sacken, Biol. Centr.-Amer.,
Diptera, vol. i. p. 67, Tab. iii. fig. 16 (1886). [New synonym. ]
Pantophthalmus helleri Kuderlein, Zool. Anz., Bd. xli. no. 3,
pp. 108, 110, tigs. 8, 9 (Dec. 20, 1912).
Although nothing can be said with reference to the male of
P. bellardii, since it is at present unknown, in the female sex,
as shown by the extensive series of specimens available to the
writer for comparison, the species is normally one of the finest
of its family, aud is conspicuous, even among the huge Diptera
with which this paper is concerned, as much on account of the
dark coloration of the body and wings, as by reason of the shape
of itsshining abdomen. While females of P. bellardi of ordinary
size are certainly not larger—perhaps if anything actually less
bulky—than corresponding examples of P. tabaninus Thunb.,
the great breadth often exhibited by the flat abdomen, combined
with the shining black surface of the first four abdominal tergites
(except the base, hind border and lateral extremities of the first,
and the hind borders and lateral extremities of the remainder),
serves to make the present species noticeable among its congeners,
among which, in actual size of the largest examples, females of
P. tabaninus are its only rivals. It is therefore not a little
curious, as well as unfortunate, that the holotype of P. bellardi
-—formerly in the Bigot collection, and generously presented to
the British Museum (Natural History) by Mr. J. E. Collin,
T.E.S.—is one of the smallest Pantophthalmids yet seen. ‘The
dimensions of a large female of P. bellardii in the National
Collection are as follows :—Total length (ovipositor and short,
blunt, facial beak included) 53-5 mm.; greatest breadth (measured
across the third abdominal segment) 20°56 mm.; wing-expanse
86 mm. (nearly 3} inches). ‘The corresponding measurements
in the case of the type are respectively 24, 9°5, and 43 mm.
Moreover, apart from its diminutive proportions, the type of
P. bellardii is anything but typical, since it is faded, rubbed, and
somewhat shrunken, having in all probability originally been
preserved in spirit before being pinned. Nevertheless, the
588 MAJOR £. E. AUSTEN ON THE
specific identity of Bellardi’s type with even the largest member
of the series of specimens under discussion is beyond question,
and the examination of a small female of P. bellardii, which has
been in the National Collection for upwards of fifty-five years,
entirely supports this conclusion, In the latter specimen, as in
the type, the shining transverse bands on the dorsum of the
abdomen are russet-brown instead of black, and a similar colora-
tion is exhibited by one or more of the larger females of
P. bellardit in the British Museum.
The particulars with reference to the fifteen specimens of
this species (including the type and paratypes of Acanthomera
championt O. Sack.) in the National Collection are as follows :—
One 2 (holotype of the species), Mexico (— Sallé), ea coll. Bigot,
presented by J. E. Collin, F.E.8.; one 2 (type of Acanthomera
championt O. Sack.), Chontales, Nicaragua (#. M. Janson); one
2, Orizaba, Mexico (— Sallé); one 2, Bugaba, Panama (G. C.
Champion); one 9, R. Dagua, Colombia (IV. F. H. Rosenberg) ;
one 9, provenance unknown (ev coll, W. W. Saunders). The
remaining nine specimens, presented to the British Museum
(Natural History) by Mr. J. E. Collin, F.E.S., previously formed
part of the collection of the late J. M. I’. Bigot, where they were
placed above the label ‘‘ A. heydeni, Wied.” * Of these examples,
one is from Mexico, two are from Nicaragua, and the remainder
(if the general Jabel can be trusted) from Brazil.
Of the two females of P. bellardii belonging to the Hope
Department, University Museum, Oxford, one is from Chontales,
Nicaragua, 1870 (#. Al. Janson), the other from Peru (“EK Mus.
Saunders, 1867 ”y.
_ From what has been stated above, it will have been seen that
P. bellardii, at any rate in the female sex, is, like other species
of Pantophthalmids, subject to great variation in size.
Infestation by Gamasid nymphs of the usual species is
exhibited by at least two of the specimens in the National
Collection ; in the cases referred to, the pseudo-parasites in small
numbers are elustered together beneath the hind margins of one
or more of the distal abdominal tergites, commencing with the
fourth. An Acarid pseudo-parasite of a different kind, deter-
mined by Mr, A. 8. Hirst as an adult female of a species of
Macrocheles, was found on the front (frons) of one of the
Brazilian females from the Bigot collection.
According to Enderlein (Zool. Anz., Bd. xliv. no. 13, p. 585,
July 28, 1914), P. helleri Ender). is a synonym of P. (Acantho-
mera) championi O, Sack. -It would be impossible to arrive at
such a conclusion from a perusal of Enderlein’s original descrip-
tion of P. helleri (vide supra), in which the front (frons) in the
female is explicitly stated to be narrowed posteriorly, and to be
less than twice as long as broad.
* Acanthomera heydenii Wied., though treated as a valid species by Enderlein
(Zool. Anz., Bd. xli. p. 102 (1912)), is in reality synonymous with Pantophthalmus
tabaninus Thunb. (cf. p. 562).
DIPTEROUS FAMILY PANTOPHTHALMIDA. 589
Pantophthalmus splendidus, sp.n. (Text-fig. 8.)
@.—Length (one specimen), including ovipositor and facial
beak, 37°0 mm.; width of head 8:4 mm.; width of front at
vertex 1:25 mm.; greatest width of abdomen (across base of third
tergite) 12°6 mm.; length of wing 28 mm.; wing-expanse
64°65 mm.
In & sea, at least as represented by type, handsome, blackish-
brown, black and orange-rufous species, with small, inconspicuous
and only moderately sharply pointed facial beak ; dorswm of thoraa
bearing a pair of admedian, sharply defined and conspicuous, pale
olive-buff pollinose, longitudinal stripes, apparent colour of which
is, in certain aspects, somewhat altered by hairy covering ; blunt-
edged abdomen, in which anterior portion of both second and third
tergites is shining, and dorsum of which is black at base, then
largely orange-rufous, with a black border on each side, im which
are a small white spot, and two short, transverse, silvery-white
streaks; dusky wings, with sharply defined, cinnamon-buff mark-
ings, as shown in text-fig. 8; hind femora and tibice not so thickly
clothed with hair as to appear furry, hind femora fairly stout,
somewhat, though not very noticeably, incrassute towards tips, and
with small subfemoral and terminal spines.
Head: front (frons), jowls and basioccipital region black or
blackish-brown, face mummy-brown, inner orbits bordering face
and at least upper portion of front more or less buffy-brown or
sepia-coloured; front of moderate width, somewhat broader at
anterior extremity than at vertex; ocelli, at least in type, orange-
cinnamon ; facial beak shining black, its base enveloped above
with a narrow, matt, mummy-brown covering ; jowls and basi-
occipital region clothed with fine, blackish-brown or dark
mummy-brown hair; occiput smoke-grey pollinose, posterior
orbits narrowly pale smoke-grey ; palpi black or blackish-brown,
distal segment somewhat swollen, pointed at tip, distinctly shorter
than remaining portion of palpi, which is clothed with blackish
hair, distal segment clothed with similar hair at base, and bear-
ing two or three hairs of same kind at tip ; antenna slender (1%. é.
depth of third (compound) segment not greatly increased at base),
blackish-brown or black, paler towards distal extremity, terminal
division of third segment greyish cinnamon-coloured at tip, first
and second segments somewhat sparsely clothed with black hairs.
Thorax: dorsum, so far as can be seen, with no trace of small or
minute, shining, rounded, wart-like tubercles, with which as a
rule in Pantophthalmidae this region is characteristically and
plentifully besprinkled ; scutellum, sides of main portion of dorsum
(scutum) behind humeral calli, and central area of dorsum in
front of preseutellar groove and behind ends of olive-buff pollinose
stripes chestnut-brown, this hue being darker on sides of scutum
and brighter in case of scutellum ; admedian, olive-buff pollinose
stripes separated in middle line by a narrow, tapering. chocolate-
brown stripe, and bordered on each side by a much broader,
590 MAJOR E. E. AUSTEN ON THE
blackish or blackish-brown longitudinal stripe, which, commencing
on front border and occupying entire space between humeral callus
and corresponding admedian stripe, is continued backwards to
prescutellar groove, becoming constantly narrower as it passes
backwards, and in particular diminishing markedly in width after
passing beyond end of olive-buff stripe, which itself disappears
shortly after crossing transverse suture ; behind latter, bordering
each blackish or blackish-brown stripe on outer side, is a fairly
broad, curved, shining stripe; humeral calli light ochraceous-bulf
in front, blackish-brown behind, outer extremity in each case
shining, pointed, and reddish-brown ; olive-buff pollinose stripes
clothed with fine, ochraceous-tawny hair, remainder of dorsum of
Text-figure 8.
oi i
THER2zIX
ie y
Pantophthalmus splendidus Austen, 9. X 1}.
scutum covered with fine russet-brown or russet-coloured hair,
lateral borders of scutum in front of transverse suture clothed
with dark brown hair, scutellum and oblong median area
immediately in front of it clothed with bright russet-coloured
hair; pleure and pectus mummy-brown, clothed with similarly
coloured hair, rounded knob below base of wing on each side
shining blackish-brown, Abdomen: first tergite black, clothed
with fine black or blackish hair, and having on each lateral edge,
a short distance in front of posterior angle, a small white spot;
second and two following tergites orange-rufous (shining anterior
border of second tergite mainly reddish-brown, at least in case of
type), lateral borders black, shading off somewhat into brown on
DIPTEROUS FAMILY PANTOPHTHALMIDS. 591
inner margin, a short, transverse, silvery-white streak just in
front of each posterior angle of second and third tergites; apart
from silvery-white streaks just mentioned, on both second and
third tergites non-shining area is confined to posterior half of
segment, is of less depth than shining area in front of it, and has
its extremities, which are far from reaching lateral margins,
tapered and rounded off; fifth tergite black, moderately shining
and narrowly margined (incompletely so in front) with orange-
rufous; remaining tergites blackish-brown or dark brown
(terminal lobes or valves ochraceous-buff, sparsely clothed with
ochraceous-tawny hairs); orange-rufous area of dorsum, as also
fifth tergite, thinly clothed with glistening, decumbent, ochraceous-
tawny hair, shining anterior border of second tergite clothed with
similar hair, second and following tergites otherwise for most
part thinly clothed with black hair ; venter black (second and two
or three following sternites shining), thinly clothed with black
hairs. Wings (apart from cinnamon-buff markings) mummy-
brown over rather less than proximal three-fourths of costal
border, then lighter or darker sepia-coloured, typical specimen at
any rate with a distinctly paler area in centre of certain cells,
such as discal and second posterior; cinnamon-buff markings
more sharply defined and more restricted than in case of several
other species of Pantophthalmus already described, chief markings
being at base of basal cells (whence extensions are emitted, to
costa on humeral transverse vein, and along fifth longitudinal
vein), between costa and proximal extremity of discal cell (includ-
ing anterior transverse vein and an extension thence for some
distance along third longitudinal vein), and streaks embracing
respectively distal extremity of second, and anterior branch of
third longitudinal veins, and distal boundaries of second basal and
discal cells; costa clove:-brown at base, otherwise mainly cinna-
mon-brown, veins elsewhere mummy-brown in darker, ochra-
ceous-tawny in lighter areas. Halteres: (in dried condition, at
any rate) stalks mummy-brown, knobs buff-yellow. Legs, except
first segment of hind tarsi, uniformly black and clothed with
black hair; first segment of hind tarsi light orange-yellow,
clothed with short, ochraceous-orange hairs, tip of segment
blackish-brown or black, clothed with similarly coloured hairs,
extreme base of segment also blackish-brown; claws black ;
pulvilli and empodia ochraceous-buff.
Panama: Chiriqui Volcano, between 5000 and 9000 ft., 1915
(1. J. Watson).
Owing to its distinctive coloration and markings, Pantoph-
thalmus splendidus—in the 9 sexatany rate—cannot be confused
with any of its congeners already known.
The specimen from which the above description has been drawn
up bears, on the fifth and following abdominal segments, a small
number of nymphs of the species of Gamasid mite (genus /'rachytes)
that has been mentioned so often in the course of this paper as
occurring on various species of Pantophthalmide. In addition to
592, MAJOR E. E. AUSTEN ON THE
these pseudo-parasites, a few other nymphal mites, apparently
conspecific with the foregoing, were found on the left side of
the thorax, among the hair on the anterior border of the meso-
pleura,
Pantophthalmus variegatus, sp. n. (Text-fig. 9.)
9 .—Length (four specimens), including ovipositor and facial
beak, 29°5 to 36°25mm.; width of head 7°5 to 9°2mm.; width
of front at vertex 1:4 to 1:5mm.; greatest width of abdomen
(across hind border of second tergite) 9:8 to 13°0 mm.; length of
wing 25:0 to 28°-5mm..
Strongly marked species, with, in 2 sex at any rate, dorsum
of thorax vinaceous-buff, conspicuously striped and spolted with
black or blackish-brown, and with dorsum of abdomen ochraceous-
orange or orange-rufous, with sharply defined black lateral borders,
in each of which are four transversely elongate, silvery-white spots,
the fourth pair of spots less conspicuous than the others and some-
times indistinct ; facial beak prominent but not sharp pointed ;
wings strongly infuscated, mummy-brown at base, then sepia-
colowred or fuscous, with ochraceous-buff or cream-buff markings as
shown in text-fig. 9; hind femora incrassate towards tips, with very
small terminal spine, subfemoral spine Likewise small, difficult to
distinguish among the hair.
Head: front (frons) warm. buff, of moderate and uniform
width; in middle line, just above bases of antenne, a narrow,
short, somewhat sagittate, brownish streak; ocellar tubercle black,
bluntly ovate, sharply defined and conspicuous ; face dark seal-
brown (facial orbits narrowly olive-buff), below facial beak with
an ochraceous-orange transverse band, extremities of which turn
upwards on each side of base of beak, a similar but paler (light
ochyraceous-buff) band, about half, the depth of the former,
immediately.above bases of palpi; jowls and basioccipital region:
mummy-brown or cinnamon-brown, clothed with similarly
coloured hair; facial beak of moderate length, blunt (not sharp-
pointed) at tip, which is shining black or blackish-brown,
proximal three-fourths of beak enveloped, except below, with a
matt, dark seal-brown, smooth (not transversely rugose) covering ;
occiput light buff (immediately adjacent to posterior orbits
ochraceous-buff), clothed with glistening, ochraceous-buff hairs,
which become brownish towards .tips; first and second seg-
ments of palpi sepia-coloured or dark brown, clothed with
reddish-brown or dark brown hair, and together longer than or
nt least as long as distal segment, latter blackish-brown or black
(sepia-coloured at base), moderately swollen beyond proximal
fourth, and sparsely clothed, at least on outer side and at base,
with short, reddish-brown hairs; antenne warm buff, second
seement with a circlet of russet. coloured hairs at distal extremity,
third (compound) segment slender, showing but little increase of
depth at base. Zhorax: dorsum of main portion (scutum)
DIPTEROUS FAMILY PANTOPHTHALMID&. 593
vinaceous-buff pollinose, with, as shown in text-fig. 9, on anterior
two-thirds a pair of fairly broad, black or blackish-brown longi-
tudinal stripes, and on each side, behind transverse suture, a pair
of similarly coloured spots, the larger of which occupies postalar
callus, while the other, much smaller spot is situate above base of
wing, just behind outer extremity of transverse suture ; a short
distance in rear of latter, each longitudinal stripe becomes: partially
obliterated or is tapered off obliquely from in front outwards,
and is then continued to prescutellar groove by a narrower and
paler extension, each extension ending on anterior wall of groove
in a somewhat ill- defined, darker, spot-like termination, which
is in contact with its filly g ; seutalar black or blackish- Eprown)
its outer extremities silver-grey pollinose on a pale ochraceous-
Text-figure 9.
Pantophthalmus variegatus Austen, 9. 1}.
buff ground; dorsum of scutum clothed with ochraceous-buft
hair, stri ipes, ‘spots, and scutellum clothed with brownish or dark
brown hair; humeral calli warm buff or cream-buff, outer
extremities and a spot on upper surface at base dark brown;
pleure and pectus mummy-brown, clothed with similarly colour ed
hair. Abdomen: first four tergites, at least in typical series,
ochraceous-orange or orange-rufous, their lateral extremities,
which are somewhat deflexed, black and sharply defined, with a
transversely elongate, silvery-white spot in posterior angle of
each tergite, spots on fourth tergite smaller than the others and
sometimes indistinct; remaining tergites, including ovipositor,
mummy-brown, a larger or smaller area in centre of base of fifth
594 MAJOR E. E. AUSTEN ON THE
tergite ochraceous-orange or orange-rufous; sixth and seventh
tergites, fifth tergite except area just mentioned, and lateral
borders of preceding tergites, except silvery-white spots, clothed
with dark brown, blackish or black hair; first tergite, except
lateral extremities, thinly clothed with fine, decumbent, glistening
ochraceous-buff hairs, remainder of ochraceous-orange or orange-
rufous area of dorsum very sparsely clothed with short or minute
hairs of similar colour; venter mummy-brown, thinly clothed
with fine, dark brown hair. Wings: markings very similar to
those exhibited by Pantophthalmus batesi Austen (cf. figs. 9 and 2);
costa, from a short distance beyond base, mainly cinnamon-
coloured; veins elsewhere partly mummy-brown, partly sepia-
coloured, except in pale areas where they are cinnamon-buff ;
teguliform swelling at base of costa ochraceous-buff, large and
conspicuous. JJalteres (in dried condition at any rate) sepia-
coloured. Legs: first segment of all tarsi (except extreme tips),
also extreme bases of following segments in case of front pair,
honey-yellow, ochraceous-buff or cream-buff, clothed with short,
glistening, cream-coloured or cream-buff hair; legs otherwise
clove-brown or chocolate-brown (last four segments of hind tarsi
sometimes blackish-brown or black), clothed with fine, similarly
coloured or blackish hair, which is closely set on hind femora and
tibise, so that these segments, or at least hind tibie, have a furry
appearance; claws black, sometimes russet-coloured at base ;
pulvilli and empodia ochraceous-buff.
Brazil. Holotype, precise locality unknown, before 1849
(Mrs. Noel); one paratype, Parana, about 1906 (H. Dukinfield
Jones); a second paratype, precise locality unknown, ex coll.
J. Bigot (presented by Mr. J. HE. Collin, F.E.S.). A third para-
type, without any indication of locality, but bearing two labels,
each with the words “ seticornis, Wied.”, and in one case with tke
addition of “EK Mus, Saund.: 1867. 10d” in J. O. Westwood’s
handwriting, is in the possession of the Hope Department,
University Museum, Oxford. ‘The holotype is the specimen
called by F. Walker (List Dipt. Ins. in coll. Brit. Mus., Pt. 1.
p. 210, 1848) Acanthomera seticornis Wied. The paratype from
the Bigot collection was placed by its original owner above the
label “A. frawenfeldi, Schiner.” It may be of interest to note
that, in the wings of the paratype from Parana, the second
posterior cell is closed and petiolate, although in all other respects,
apart from post-mortem shrinkage of the abdomen, the specimen
ig normal,
While the thoracic spots in the case of Pantophthalmus
variegatus, 2, are similar to those exhibited by both sexes of
P. vittatus Wied., though the posterior lateral spot is much
larger in the species characterized above, the distinctive thoracic
stripes in P. variegatus will at once serve to separate it, not
only from P. vittatus, but also from any other Pantophthalmid
in which at any rate the female sex is known.
It only remains to add that nymphs of the usual species
DIPTEROUS ‘FAMILY PANTOPHTHALMIDZ. 595
of Gamasid. mite, ‘albéit in’ small numbers, are present on the
abdominal tergites of nt least two of the paratypes, as: well as on
those of the holotype of P. variegatus.
Genus Rhaphiorhynchus Wiedemann.” . ae ;
Rhaphiorh es Wiedemann, Diptera Exotica, Pars i. p. 59
(1821). ,
- In‘spite of the lapse of more than a century since this genus
was first described, the genotype, Ithaphiorhynchus’ planiventris
Wied., is still the only species that can be assigned to it; as
explained below (cf. p. 597), it now seems more “reasonable ta
found a distinct. genus for Lthaphiorhynchus rothschildi Austen,
Text-figure 10.
=
' « Left maxillary palpus of Rhaphiorhynchus planiventris Wied.; a, g 5h Qs.
Inner (ventral) aspect in each case.
ws
which was described in 1909. At the same time, the distinctive
shape of the terminal segment of the palpus in Rhaphior hynchus
planiventris, especially i in the female, precludes any thought of
including this species in the genus Pantophthalmus. As shown
in fig. 10, the distal segment of the palpus in Ehaphiorh ynchus,
while distinctly swollen and fusiform in the male, is enormously
dilated in the opposite sex and, viewed from below, is decidedly
reniform. A prominent and sharp- pointed facial beak, and o
Proc. Zoou. Soc.— 1923, No. XXXIX. | 39
696 MAJOR E. E. AUSTEN ON THE
well-developed subfemoral spine on the hind legs are present in
both sexes,
Rhaphiorhynchus planiventris Wied.
Rhaphiorhynchus planiventris Wiedemann, Dipt. Exot. 1.
p. 60, Tab. ii. fig. 1 (1821). . i
Acanthomera crassipalpis Macquart, Mém. Soc. roy. des Sc.,
dé Agric. et des Arts de Lille, Année 1846, p. 43, Tab. i.
fig. 3 (1847).
Acanthomera bigoti Bellardi, Saggio di Ditterol. Messicana,
Appendice, p. 16, ‘lav. iii. fig. 10 (1862). .
The specific identity of Acanthomera crassipalpis Macq. and
A. bigoti Bell, with Ith. planiventris Wied. was recognized long
ago by Baron O. R. Osten Sacken (Biol. Centr.-Amer., Diptera,
yol. i. p. 66 (1886)).
The undermentioned specimens of 2th. planiventris are in the
British Museum (Natural History) :—One ¢, David, Chiriqui,
Panama, and oné 9, Bugaba, Panama, 800-1500 ft.—both col- —
lected by Mr. G. C. Champion, and determined by Osten Sacken
(loc. cit.); one 9, Provincia Sara, Santa Cruz de Ja Sierra, Bolivia,
February—April, 1904 (J. Steinbach). From the Bigot collection,
Mr. J. E. Collin has kindly presented the type of Acanthomera
crassipalpis Macq. (a 9, stated by Macquart, loc. cit., to be from
Guatemala), and twoadditional 2 9, from Chontales, Nicaragua,
all of which, with a 2 of Pantophthalmus (Acanthomera) frauen-
feldi Schin. (vide supra, p. 565), were placed by Bigot aboye a
label inscribed * Acanthomera crassipalpis Macq.,n. sp. 2 .”
In the Hope Department of the University Museum, Oxford,
this species is represented by three 9 9, of which two are from
Chontales, Nicaragua, 1870 (#. MM. Janson), while the third is
simply labelled ‘ America, aeq.”
he Liverpool School of ‘I'ropical Medicine possesses a 2 of
Rh. planiventris from the R. Amazons, Brazil, “on board
S.S. ‘ Hildebrand,’ 25. ii. 1920, 6.30 p.m.” (Dr. Clarke).
All three Oxford specimens show the nymphal stage of the
usual species of Gamasid mite on the dorsal surface of the
abdomen ; in two instances, however, only a single parasite is
visible, while in the third case the nymphs number less than
a dozen, as
Genus Atopomyia*, nov.
Distinguished, at least in g Sex, from both Pantophthalmus
Thunb. and Rhaphiorhynchus Wied. by the narrow, elongate
shape of the body, and the forin of the third (compound) segment
of the antenna. The latter, instead of its broader (proximal)
portion being more or less abruptly truncate (or tapering quickly
to 4 point), and bearing terminally a fine, setiform arista (as in
* droztos, strange, extraordinary, pia, a fly.
DIPLEROUS FAMILY PANTOPUTHALMIDE, — ' 597
Pantophthalmus §), or being lanceolate (as in Rhaphiorhynchis
planiventris Wied., 3, apud Macquart, Mém. Soe. Roy. des Sc.,
de VAgric. et des Arts de Lille, Année 1838, 2itme Partie,
pl. 20. figs. 3,3 a, 1838), is narrow and subulate, with its
terminal-annulus elongate, rod-like, abruptly attenuate and
pointed at the tip, and bearing a few minute hairs towards the
end, in addition to a longer recumbent hair above and below,
just before the middle. Distinguished further, from both genera
mentioned, by the palpi (at least in ¢) being densely clothed
with stiff hair, instead of the distal segment. being practically
bare; distinguished also from Rhaphiorhynchus by distal segment
Text-figure 11.
Left antenna of Atopomyia rothschildé Austen, §, from the inner side.
of palpi not being characteristically swollen as in that genus, and
being decidedly shorter than remaining portion of palpus, instead
of vice versd, Face with a large, thick beak, pointed at tip,
thickened portion more or less distinctly wrinkled transversely.
Dorsum of thorax, at least in g of genotype, devoid of the small
or minute, rounded, wart-like tubercles, which, though varying
greatly in different species in size and number, are characteristic
of the majority of Pantophthalmide, though apparently not
present in all, at any rate in both sexes. Hind legs with a large
subfemoral spine; terminal spine on hind femora very small, at
least in genotype.
Genoty pe—Lhaphiorhynchus rothschildi Austen,
Atopomyia rothschildi Austen.
Rhaphiorhynchus rothschildi Austen, Novitates Zoologice,
vol. xvi. p. 129 (May 1909), and vol. xvii. pl. xv. fig. 1
(December 1910).
Although at the time of describing this striking species the
author considered that there were no real grounds for separating
it from Lthaphiorhynchus (cf. Austen, é. cit. vol. xvi. p. 130), it
seems advisable on further consideration to reverse this view,
especially when regard is paid to the important palpal characters
to which attention is drawn in the generic diagnosis above.
The type of A. rothschildi—a § from Buenavista, Bolivia
(J. Steiubach)—is in the Tring Museum, and the British Museum
(Natural History) possesses a paratype of the same sex from
Ecuador (Buckley). During the fourteen years that have elapsed
sincé the species was described, no further specimens of it have
3u*
598 ON THE DIPTEROUS FAMILY PANTOPHTUALMIDA,
been added to the National Collection or seen by the author ; but
it may be noted that two ¢ ¢ of A. rothschildi, from Peru and
Chiriqui, Panama, respectively, were in the possession of the late
Professor Hermann (cf. Hermann, Deutsch, Ent. Z. yelalanise 1916,
Heft i. p. 47, 1916).
According to Lord Rothschild (Nov. Zool. vol. xvii. p. 461,
December 1910) ,Atopomyia rothschildi mimics the Hymenopteron.
Pepsis elevuta Fabr., which is doubtless the explanation of the
curious contrast in its general fucies presented by this species, as
compared with any other Pantophthalmid at present known.
ON NEW CRYPTOSTOME BEETLES, : 599
31. New Cryptostome Beetles. By S. Mauntm, F.Z.S.
[Received May 7, 1923: Read May 29, 1923.]
(Text-figures 1-8.)
In the present paper are described one new genus and six new
species of Hispinze and Cassidinaz, Two facts are interesting :
the association of two Hispids of the same genus, structurally
different but having strong resemblance in superticial colouring,
in the same food-plant ; and secondly, the new Cassid beetle from
Palestine possesses a claw-structure which is not found usually
in this group of beetles. These facts are described in detail
below. :
Text-figure 1.
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Dorsal view of Callispa almora.
CALLISPA ALMORA, sp. n.
Body elongate. Colour shining yellow; mouthparts, antenne,
trochanter, apices of femora, and bases of tibiw pitch-black;
underside. brown. oe : Seer
600 MR, 8S. MAULIK ON
A
Head smooth, impunctate, and slightly produced between the
bases of the antennee, The antenne are hardly thickened towards
the apex, the apical joint pointed, subnitid; the third joint is
longer than the second, but slightly shorter than the first and
second combined. Prothoraa almost as long as broad, slightly
narrowed in front; the basal margin bisinuate, tho sides straight
and margined, the anterior angles rounded, the posterior acute.
The upper surface with a longitudinal depression on each side,
and also with one in the: middle at base ; in the lateral depressions
are some coarse punctures; the middle longitudinal area slightly
convex, but having similar punctures except along a narrow
middle line ; some of the punctate portions are slightly depressed.
Scutellum obtong, sides straight, anterior edge broadly rounded ;
surface smooth, shining, impunctate, Alytra very sli ehtly broader
at base than the prothorax, parallel-sided, punctate-striate; besides
the short scutellar row on each elytron there are eight rows at
the base and ten rows in the middle; the interstices are smooth
and flat. The punctures are much stronger in the middle area
than elsewhere. Underside smooth, shining, impunctate except
for a few coarse punctures on the sides of the thoracic sterna.
Length 5 mm. .
India: the Himalayas, Almora, 4500 ft., Khaula (//. G.
Champion).
Type in the British Museum,
Described from six examples.
This species is relited to C. nigritarsata Maulik, which is from
Ceylon, but differs in the coloration, the relative lengths of the
second and third joints of the antennz, and the structure of the
prothorax with its punctation.
DowNESIA SASTHT, sp. n,
Body elongate, narrow, ‘ Colour shining brown, with the eyes,
antenna, and the apical area of the elytra black.
Head with the eyes large; the interocular space smooth, with a
few punctures. ‘The antennw are short, scarcely reaching the
middle of the pronotum, very slightly thickened towards the apex,
and covered with hair, more so towards the apex ; the first joint
rounded, the second longer than the third, the third, fourth, fifth,
and sixth almost equal to each other in length, from the seventh
to the eleventh slightly thickened, the last the longest and bluntly
pointed. . Prothorax quadrate, longer than broad, very slightly
narrowed behind, the sides parallel and margined ; the anterior
angles rounded, the posterior ones emarginate. Basal margin at
the middle deeply channelled. ‘he upper surface sparsely and
confusedly punctate, there being less punctures in the middle than
elsewhere; the front rounded margin impunctate. Scutellum
small; impunctate. “lytra broader at the base than the prothorax,
parallel-sided, yery slightly constricted in the middle; at the
rounded apical angles the margin is explanate. Hach elytron has
NEW CRYPTOSTOME BEETLES, 601
six rows of punctures on the apical surface: the first runs along
the suture right to the apical surface, where it does not become
double, the next row of punctures divides itself into two on the
black apical surface of the elytra, then another row commencing
from the humerus soon becomes double, finally there is the
marginal single row; on the apical area the third interstice is
raised into « short, sharp costa; all the interstices have the
appearance of being slightly raised ; on the brown portion, of the
elytral surface the punctures are surrounded: by a black ring,
and hence appear much larger than those on the black area; a
scutellar row is'‘absent. Underside smooth, shining, impunctate.
" Text-figure 2,
Dorsal view of Downesia sasthi.
The tarsi of the front legs almost equal in length to the tibise
and larger than those of the other legs. The claw-joint of the
tarsus projects beyond the third joint. —
Length 5-6 mm.
W. Himalayas: Kumaon (Z/. @. Champion). Found on Thy-
sanolana agrostis.
Type in the British Museum. -
Described from nine examples.
These nine examples were sent with four examples of
Downesia gestroi Baly, all found in sheaths of Neyraudia, to
602 ©.» MR, 8. MAULIK ON ~
Mr. G. C..Champion, who was. good enough to afford me. the
opportunity of examining them. aK RE AG :
. ‘Lhe superficial resemblance between D. gestroi and D. sasthi is
‘so remarkable that they were considered as the same species. ‘he
structural differences between them are considerable, and warrant
the erection of a new species. The close association of these two
structurally different forms, but possessing a strong superticial
resemblance, in the same food-plant is interesting, and is sugges-
tive of some form of mimicry. D. ceylonica Maulik, D. ratana
Maulik, and D. kanarensis Weise have also similar coloration.of the
elytra, viz. the apical portion is black while the basal is brown. .
Kani, gen. nov.
This genus is erected for the insect of which a detailed
description is given below, according to the following characters :
(1) the shape of the body, parallel-sided, rounded posteriorly, the
pronotum transversely elliptical ; (2) the structure of the antenne,
which partly lie in the channel on each side of the cavity in which
the head is imbedded ; (3) the peculiar structure of the tarsus,
in which there is no felt covering on the underside, and the
deeply bilobed character, especially of the third joint, is absent.
The structure of the tarsi suggests that they have been adapted
to a desert life. The drab colour of the insect seems to support
this view. At present there are no data available with regard to
the larve and the bionomics of the insect. By the shape of the
body and the antennal channels it is related to the genus Glypho-
casis, but the structure of the tarsus places this genus in a unique
position. :
KARI BRUNNEA, sp. n.
Body oblong, parallel-sided, rounded posteriorly ; the front of
the pronotum slightly drawn forwards ; in profile the dorsal side
of the insect has a slight elevation posteriorly. Colour dull
brown; the posterior undulated edge of the pronotum opposing the
anterior similarly undulated edge of the elytra, black; the eyes,
mouthparts, and the five apical joints of the antennz black or
piceous.
Head imbedded in a hollow, each side of which forms a channel
in which lie the six basal joints of the antenns. The clypeus flat,
indented with large coarse pits, some of which are confluent with
each other and bear a few erect hairs. The antennz pass just
beyond the base of the prothorax on the underside, the first joint
thickened, club-shaped; the second small, rounded; the third
longer than the fourth, which is equal in length to the fifth ; the
sixth slightly shorter than the fifth ; seventh to eleventh joints
form an elongated thickened club. ‘The whole antenna is sparsely
covered with short, erect, whitish hairs. Prothorax transversely
oval, broader than long, as broad at the base as the elytra; the
middle of the slightly drawn out front edge with a very shallow
NEW CRYPTOSTOME BEETLES. 603
‘emargination ; each half of the basal margin bisinuate, the lateral
margins rounded. ‘The edge all round, except the basal black
‘portion, bears short, erect, whitish hairs, is uneven, more so at the
lateral rounded portions, and is generally darker than the ground-
colour of the pronotum. The upper surface is very uneven,
having the following structure: the middle of the basal area is
enormously raised into two high ‘peaks (they being the highest
points of the surface) with a depression between them ; on each
side of these there is a broadly convex, uneven, and oblique ares
sloping down to the edge; anterior and posterior to these swollen
areas are deep depressions ; in front the surface also slopes down,
and is broadly convex and tuberculate, narrowing down in the
middle into a longitudinal ridge which reaches the front edge; on
either side of this ridge the surface is deeply depressed; the frontal
area, including the swollen and the depressed portions, contain
Text-figure 3.
Dorsal view of Kari brunnea.
round black spots; the posteriorly situated depressed areas also
contain similar spots; some of these may be impressed pits or
punctures. Scutellum triangular, with the surface fairly plane.
Elytra: the front black edge is bisinuate on each side and
roughly serrate. The edge all round is roughly serrate, darker
than the body-colour, and bears short, erect, whitish hairs.
Each elytron is punctate, and with coste and tubercles. The
whole surface is very sparsely scattered over with short, erect,
whitish hairs which are visible under a high power. The suture
is strongly raised. On each elytron there are four coste: the first,
that nearest to the suture, is the strongest, running the whole
length parallel to the suture, and in its posterior portion at several
places swollen into tubercles and joined to the suture and to
the second costa by many short transverse cost; the second and
604 » ~ MR. S, MAULIK ON,
third cogte are feebler, and arising from the humerus they break
up jnto tubercles posteriorly without reaching the apical area of
the elytron; the fourth costs is hardly developed at the base, but
more strongly so posteriorly, bending round to join the first costa
on the apical area; between the fourth costa and the margin,
Text-figure 4.
Ventral view of Kari brunnea.’
fr i “i Se ct eadiGy
which is also raised, the surface is much depressed, containing
deeper punctures and much broken up at various places by
swellings. Underside more shiny than the upperside. The
epipleurz of the elytra and the under surface of the pronotum
bear a few scattered dark punctures. The legs are very sparsely
Text-figure 5.
Under surface of tarsus of Kari brunnea.
scattered over with short, erect, whitish hairs. The first joint
of the tarsus is constricted at the base and dilated at the apex ; the
second is broader than long, slightly emarginate at the apex ; the
third is similar to the second in structure, not deeply bilobed as
is usual in the Chrysomelide; the claw-joint arises from the base
NEW CRYPTOSTOME BEETLES. 605
of the third, long, more or less eylindrieal, projecting much
beyond, as broad at the base as at the apex; the claws are simple.
There is no felt’ covering underneath, as is usual in the Chryso-
melide, Viewed from above, the tarsus appears to have uniform
breadth throughout its length, and is covered with stiff, erect
hairs. Each side of the first, second, and third joints has a little
strongly chitinized projection whichis reminiscent of the lobes
of the joints; these projections are clearly visible when the tarsus
is viewed sideways, and on their underside they bear tufts of
bristles, so that there are altogether three pairs o them.
Length 5 mm.
Locality. Palestine, 10 miles east of J erusalem, 1000 ft.,
17. iv. 1922 (P. A. Buston).
Type in the British Museum.
Described from one example.
CASSIDA RATI, Sp. n.
Body subtriangular. Colour yellow-brown ; elytra with ill-
defined darker patches, the explanate margin at the anterior and
posterior angles with deep red-brown patches, the sutural angles
deep red-brown; underside (except the legs) shining black.
'
Text-figure 6.
Dorsal view of Cassida rati.
Head with the clypeus triangularly elevated towards the base
of the antenn, otherwise the surface is plane, having stiff, long
hairs at the sides. ‘lhe antenne pass beyond the anterior angles
of the explanate margins of the elytra; the first joint elongate,
thickened ; the second short; the third, fourth, and fifth almost
606 . MR. S. MAULIK ON
equal to each other i in n' length ; : the sixth shorter, slightly dilated
pt the apex; the seventh still shorter, more thickened ; the eighth
to eleventh slightly thickened, black, and more hairy. Prothorax
much narrower than the base of the elytra, broader than long.
The sides rounded and front margin Widely arched; each half of
the basal margin bisinuate and edged with black. ‘The surface
is convex, smooth, shining, impunctate, and slightly sloping from
the base to the front; on each side of the disc there are two
indistinct, roundish, darker patches. The explanate margins are
transparent, with a honeycomb structure. Scutellum triangular,
smooth, impunctate. Elytra much broader at the base than the
prothorax.: A little posterior to the sentellum the surface is raised
into a hump (not visible in the illustration). On each elytron
there are nine rows of rather coarse punctures: the two nearest
to the suture run right up to the’ end, the others converge and
terminate on the apical area of the elytron; besides these rows
a few punctures may be said to represent a short scutellar row,
and some very coarse pits, more regular posteriorly, along the
extreme margin may represent a tenth row of punctures. The
interstices are more or less costate, particularly the first two.
The surface has some indistinct patches of dark-brown colour ;
the red-brown fascia on the explanate margin at the anterior
angles (which are rounded) continues as a lateral band, joining
the similar fascia at the posterior angle. The explanate margins
transparent and with a honeycomb structure. Underside: “the
abdominal sternites sparsely covered with brownish hairs ; those
on the side longer. The claws simple, projecting slightly beyond
the bilobed joint. bag
Length 7 mm.; breadth 6 mm.
India: Manipur (Doherty).
Z'ype in the British Museum,
Described from one example.
CAssIDA MANIPURIA, Sp. n.
Body subtriangular, ‘Colour PEllgw brown, with darker brown
patches on the elytra. The anterior lateral angles of the ex-
planate margin of the elytra are very sharp.
Head with the clypeus slightly convex, smooth, impunctate.
The antenne extend beyond the anterior angles of the explanate
margin of the elytra ; first joint elongate, club-shaped ; the second
almost as long as the third, constricted at the base a little dilated
towards the apex; the third, fourth, and fifth almost equal to each
other in length; the sixth shorter; seventh to eleventh a little
thicker, more hairy; the last two piceous. Prothoraz elliptical,
much broader than long, the front margin widely arched. The
dise convex, slightly sloping from the base to the anterior
margin, smooth, impunctate, The explanate margin transparent,
with a honeycomb structure, Scwtellwm triangular, smooth,
shining, impunctate. Zlytra much broader at the base than
» NEW CRYPTOSTOME BEETLES, -; 607;
the prothorax. That the’anterior lateral angles of the expla-
nate margins of the elytra are acute and sharp is an important,
character. A little posterior to the scutellum there is a hump
(not visible in the ‘illustration). On each’ elytron there ig :a
short scutellar row consisting of a few punctures, and ten rows
of coarse punctures, including the extreme marginal one which
consists of a few coarse pits, ‘I'he rows are not very regular, in
some places the punctures being confused. ‘I'he interstices are
more or less costate. The dark brown patches. are obsolescent,.
there being two, one below the humerus and the other at the,
posterior angle, dark pitch-black ; these may form with the obso-
lescent patches a lateral band. The explanate margins are
transparent, with a honeycomb structure. Underside uniformly
Text-Sigure 7.
Dorsal view of. Cassida manipuria.
brown, with the middle area of the abdomen.a shade darker.
The claw-joint hardly projects beyond the bilobed joint ; claws
simple. s
Length 7 mm.; breadth6 mm. . me eee i
India : Manipur (Doherty).
Type in the British Museum.
Described from two examples, one of which is defective.
CASSIDA DIMBAKAR, 8p. Nn.
Body elongate-ovate. Colour uniformly brown above; under-
side black, except the legs and the sides of the abdomen.
Tead black, with the clypeus flat. The antennae hardly
reaching the base of the elytra; the first joint thickened, club-
608 ON NEW CRYPTOSTOME BEETLES.
shaped s the second smaller; the third, fourth, and fifth almost
equal to each other in length; the sixth and seventh slightly
shorter; the eighth to eleventh formed into a thickened club,
which is piceous. Prothorad broader than long, as broad as the
elytra at the base, the lateral angles being near the base; the front
niargin uniformly arched. The upper surface slopes from the base
to the anterior margin, convex and impunctate; the explanate
margin with a honeycomb structure. Scutellum small, tri-
angular, smooth, and impunctate. Llytra ovate, with the sides
and the explanate margins sloping down. On each elytron,
Text-figure 8.
Sleece= OL,
Ccecoes (A
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Dorsal view of Cassida dimbakar.
besides a very short scutellar one, there are eleven other rows
of punctures; the fourth row is irregular and confused, the rows
meeting in pairs on the apical area, ‘The interstices more or less
costate. Underside: abdominal sternites slightly covered with
hairs.
Length 5°5 mm.
India: ex coll. Linn, Soc:
Type in the British Museum.
Described from one example.
ON TUE DISPLAY OF THE MAGNIFICENT BiND-OF-PARADISE. 609
32. On the Display of the Magnificent DBird-of-Paradise,
Diphyllodes magnifica hunstein. By D. Suru-Smiru,
IZ. i
[Received June 9, 1923 : Read Juno 12, 1923.]
(Text-figures 1—4.)
At a meeting of this Society, held on February 20th, 1923,
I made some remarks on the nuptial display of the various
species of the Paradiseide, and referred especially to that of
Diphyllodes, of which there was a male example of the form
Text-figure 1,
First attitude of display.
f
from south-east New, Guinea (D. m. hunsteini) in the Society’s
collection. The remarks then made and the drawings exhibited
were the result,.of brief observations, the bird having only
recently commenced to display; butsince then-it has piven many:
610 MR, D, SETH-SMITH ON THE DISPLAY i
exhibitions, having been displaying on and off for the past five
or six months,
Further observations have revealed certain details in the
attitude adopted that were not at first apparent.
Dr. Philip Manson-Bahr has spent many hours in sketching
this very difficult subject, and his series of beautiful coloured
drawings now exhibited will be fully explained by him *, For
my own part I-decided to try and photograph the bird during
display, knowing that in the dull light in which the cage
is situated and the presence of intervening bars it was in pos-
sible to obtain presentable pictures, but hoping to secure records
4
Text-figure 2.
Second attitude of display.
of the different positions adopted, from which it would be possible
to make accurate drawings. .
Before describing the display of this species it will be as well
to give a brief description of the bird itself. About the size of
a Starling, it has the top of the head covered with velvet-like
feathers of a brownish colour; the back is metallic reddish
chestnut, becoming orange on the lower back, the wing-coverts
yellow with o distinct shade of orange, the abdomen and under
tail-coverts blackish..purple. ‘The tail, which is very short
» * Dr, Manson-Bahr’s serios of coloured drawings of this bird have been presented
by him to the Zoological Society and placed in the Library, _ :
OF THE MAGNIFICENT BIRD-OF-PARADISE. 611
except for its two middle feathers, is dark brown, the central
pair of feathers being produced into very narrow plumes some
nine inches in length and of a brilliant metallic green on the
upper surface—they cross one another and spread outwards in
opposite curves. Covering the under surface of the body from
the throat, and occupying the whole of the breast, is a shield of
rich velvet-like green, the lower feathers tipped with metallic
blue, and down. the middle of this, commencing below the black
throat, is a narrow band of square-ended feathers-of a rich
metallic emerald-ereen. From the hind neck grows a broad
Text-figure 3.
Final attitude with mouth open.
fan-shaped hood of pale yellow plumes, with the texture of spun
glass, flanked on each side by tufts of reddish-brown feathers.
The bill and feet are blue, and the inside lining of the mouth
delicate green.
Birds-of-Paradise comprise one of the few groups of Passerine
birds that are polygamous, and where polygamy prevails in birds
it is generally accompanied by great adornment of the male eex.
Various attitudes of display are adopted, calculated to exhibit
these adornments to the best advantage.
The methods of adornment of the males in the Birds-of-
Paradise are very varied and the attitudes adopted during
display differ in every genus, that of Diphyllodes being entirely
Proc. Zoou. Soc.—1923, No. XL. 40
612 MR. D. SETH-SMITH ON THE DISPLAY
unlike that of any other Bird-of-Paradise so far as our know-
ledge of these birds goes; and, before the specimen now in the
Society’s Gardens commenced to display, this remarkable per-
formance had apparently never been witnessed. .
There is more than one attitude of display assumed in the
ease of most birds, and in D. m. hunsteini there appear to be
three or four which, as a rule, follow one another in regular
sequence, though frequently the final attitude is assumed without
the others.
When inclined to display the bird sits on the perch with the
plumage rather relaxed and utters two or three notes sounding
Text-figure 4.
Final attitude with mouth closed.
like gua-qua-qua, and then suddenly adopts the attitude depicted
in text-fig. 1, the pectoral shield being brought into prominence
and its sides thrown up considerably above the level of the head—
at the same time the tail being thrown sideways. In the second
stage the body is brought into a horizontal position, the sides of
the pectoral shield erected, and the central tail-feathers held
above the level of the body with their brilliant metallic surfaces
visible from the front (text-fig. 2).
Ina third stage, which is not often observed, the head is thrown
further back and the erected tail more forward, the two long,
\
OF THE MAGNIFICENT BIRD-OF-PARADISE. 613
plumes reaching to the head and being swayed from side to side.
In both of these and in the final stage the bill is from time to
time opened to exhibit its delicate green lining.
In the final stage, which is depicted in text-figs. 3 and 4a
complete transformation takes place, the body being suddenly
brought into an erect position, the nuchal hood of pale yellow
feathers flashed into prominence above the head, the pectoral
shield lowered to form a broad ribbon-like strip of bright green,
down the centre of which now appears a narrow band of iridescent
feathers of emerald-green or brilliant blue according to the angle
of the light in which it is viewed, while at the lower extremity
of the shield appears a narrow edging of the same hue.
40**
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THE BRAIN OF THE ZEUGLODONTIDA. 615
33. The Brain of the Zetiglodontidse (Cetacea). By Ray-
monn A. Dart, M.Se., M.B., Ch.M., Professor of
Anatomy, University of the Witwatersrand, Johannes-
burg; with a Note on the Skulls from which
the Endocranial. Casts were taken. By C. W.
Anprews, D.S8e., F.R.S., ¥.2Z.8., British Museum
of Natural History.
[Received May 18, 1923: Read June 12, 1929.]
' (Text-figures 1-24.)
TasLE oF ContTENTS.
Page
MSATNGradwetionucscssva-ccssssssese ste creche tence fences tetaveneeeeeeeemOLO
SMa teninl eae ee eee ks eo nea Ca ney, ate ane LG
3. The two Kudocranial Casts unaccompanied by Osseous
Remains ............ .. 618
a; Zeuglodon sensitivus, sp.nov. (M. 12123.)
b. Zeuglodon elliotsmithii, sp. nov. (M. 12066.)
4, The three Endocranial Casts forming a Phyletic Series ... 627
a. Prozeuglodon atrox Andrews. (M. 9265.)
b. Zeuglodon intermedius, sp. nov. (M. 10173.)
c. Zeuglodon osiris Dames. (M. 10228.)
6. Table of Measurements of the Endocranial Casts ......... 634
6. The Zoological Position of the Zeuglodontide ............... 635
7. Comparison of Zeuglodont and Prosqualodont Brains...... 638
8. The Trigeminus and the Law of Infiltration in Cerebellar
Evolution ......... 643
9. Note on the Skulls from whicle the Endocranial Casts
were taken, by C. W. ANDREws, D\Sc., F.R.S. ..+...... 648
FIO, LENCE D NF cds noemandandsocandedcondsoedononenacadsoteaciinos baboon
J. Inrropucrion.
“The Cetacea are connected with the marine Catnivorés
through the genus Zeuglodon, as Huxley has shown, and the points
of resemblance are so marked that the affinity cannot: be doubted”
(O. C. Marsh, 1877). |
It has been suggested, in the past, that these water-living
forms, which preserve in their skeletal parts characteristics
linking them with the ancestors of modern Carnivora, were the
actual ancestors of the whales and the dolphins, 7. e., the
Cetacea. The Zeuglodonts were therefore grouped together as
the Archxoceti. This suggestion of a Carnivore (or Creodont)
ancestry seems to have been universally admitted until Matthew
and Gregory (1910) pointed out various characteristics which
‘6.16 a PROF. Ri A. DANT ON TIE *
seem to link the Zeuglodontide more closely with Insectivore
antecedents. In the meantime, the intimacy of the relationship
between Zeuglodontide and Cetacea has been reaffirmed by
numerous obser vers, including amongst many.others Abel (1905-
1913) and Winge (1919, 1921).
Previous investigation of the group has naturally centred
around the osteological remains, and has shown (Andrews, 1907)
that they probably arose ‘on the - nor thern shores of the
Ethiopian land in the early part of the Eocene period,” and that
“by the end of the Middle EKocene the true Zeuglodonts had
come into existence, and had,spread_ rapidly over the earth, their
remains being found in the upper part of the Hocene of America,
England, and New Zealand.”
Since the appearance: of Professor G. Elliot Smith’s important
contribution to the study of the form of the endocranial cast.
(1903) no new feature of the cerebral anatomy of these forms has
been recognised, so far as I have discovered, although Stromer
(1908). has fieured what is apparently a unique natural endo-
cranial cast in a form which he styles Zeuglodon osiris.
Shortly after the publication of Professor Elliot Smith’s article,
Mr. H. J. L. Beadnell gave him an alinost perfect natural endo-
cranial, cast of a Zeuglodont collected in the Egyptian Fayum at
the locality known as the Gar-el-Gehannem. Professor Elliot
Smith kindly placed it at my disposal some timeago. I was assisted
in the “‘ development” of this natural cast by Professor D. M.S.
Watson, who carefully removed most of the adherent bone and
matrix, giving the result which has been accurately reproduced
by Miss Davison in text-figs. 1, 2, and 3. ‘This specimen has
since been.given ‘to the British Museum by Professor G. Elliot
Smith and is distinguished by the collection number M. 12123.
The majority of the remaining figures have been drawn by
Mr. Poulton, artist to the Anatomical Institute of University
Collegé, London,
Owing to the courtesy of the British Museum officials, of
Dr. A. Smith Woodward, and especially of Dr. C. W. Andrews,
who has assisted me at every turn in this work, I have been able
to draw upon the paleontological resources of that institution.
The Zeuglodont material there has been examined and extremely
satisfactory casts made of the brain-cases, through the skilful
-work of Mr. L. E. Parsons,
_ The conclusion of this research would have been. impossible
apart from the courtesies that have been extended to me not only
-by the above-mentioned persons, but also by Sir Arthur Keith
and Mr. Burne of the Royal College of Surgeons. ‘To one and all
I tender my grateful thanks.
2. MATERIAL.
One of the most important results that has emerged from the
research is the definiteness with which we can now determine
BRAIN OF TILE ZEUGLODONTID™. 617
certain different Zeuglodont species. The species Zeuglodon osiris
Dames, 1894, was originally known from its lower jaw and certain
other. skeletal. remains. Stromer (1903) correctly identified the
skull of this creature, having in his possession an almost complete
head. Ina later paper, however, Stromer (1908) has caused con-
fusion by referring an entirely different Zeuglodunt to the same
species. It became necessary, therefore, to discover which form
actually was Zeuglodon osiris and, working with this information,
to arrange our series.
In this matter two lines of evidence have substantiated our
findings. illiot Smith (1903) described an’ imperfect endo-
cranial cast (M. 8150) which was known to have come from
a skull (M. 8150) determined as Zeuglodon osiris Dames by
Dr. Andrews in Egypt at that time. Fortunately, casts both of
the cranial cavity and of the skull itself were amongst the British
Museum material. In addition there were two skulls in the
Museum, both of which were termed Zeuglodon osiris. A. more
intimate survey of these two skulls (M. 10228 and M. 10173) has
rendered necessary their separation into two different species.
Further, by means of the matrix in which they are embedded,
Dr. Andrews is able to recognise them as probably coming from
two entirely different beds of the Middle Eocene epoch. The
first (M. 10228) possesses the same characters as M. 8150, and is
actually Zeuglodon osiris Dames (see text-fig. 14) from the Qasr-
el-Sagha Series. These two skulls (M. 8150 and M. 10228) con-
form entirely to the description of that type given by Stromer
1903).
This primary orientation having been achieved, it became
apparent that the second skull (M. 10173), hitherto classed
indiscriminately as Zeuglodon osiris, was not Zeuglodon osiris but
another form altogether. The matrix in which it is embedded
shows that it probably comes from the Birket-el-Qurun Series
(Operculina-Nummulite beds). Its characters are intermediate
between those of Zeuglodon osiris of the Carolia beds and Prozeu-
glodon atroc Andrews (M. 9266) from the Ravine beds. The
endocranial casts confirm these facts; indeed, it was the serial
conformity of the casts which led us to a re-examination of the
osteological features.
Therefore, three of the endocranial casts here described come
from skulls whose osteological features are known. Further, as
a group, these three skulls and their casts may be conveniently
regarded as a phyletic series. In view of this fact, Dr. C. W.
Andrews has kindly written an account of the osteological
features of the new form (M. 10173), which we will call provi-
sionally Zeuglodon intermedius, sp. nov., and has stated its geo-
logical horizon to have probably been the Birket-el-Qurun scries
(vide p. 35, ‘The Topography and Geology of the Fayum Province
of Egypt,’ IH. J. L. Beadnell, Survey Department, Cairo, Egypt,
SOD) am
618 PROF. f. A. DART ON TIE
vs
3. THe Two EnDOcRANIAL CAsts UNACCOMPANIED BY
OssEous REMAINS.
Unfortunately there are no bony remains associated with the
natural cast (M. 12123) spoken of at the outset. This cast,
although belonging to some Zeuglodont, is so entirely different
from any one of the phyletic series that it certainly comes from a
species of Zeuglodont not known in the British Museum collection.
Lack of information prevents our associating it with any other
Zeuglodont hitherto described. This cast presents characters
which indicate its affinity with the natural cast (M. 12066)
described by Elliot Smith (1903)—which also was unaccompanied
by any osseous remains—rather than with any member of this
so-called phyletic series. This natural cast does not come from
another member of the same species as that described by Elliot
Smith, nor is it an endocranial cast of Zeuglodon osiris. Com-
parison with a duplicate (M. 12066) of the natural cast deseribeil
by Elliot Smith and with Zeuglodon osiris (M. 10228) demon-
strates a wider degree of separation from both of these than could
be accounted for on a specific differentiation alone; we may be
dealing here with different genera.
Tn the case of these casts (M. 12123 and M. 12066) the absence
of knowledge concerning the skeleton and the exact horizon from
which they have come makes it an invidious matter to establish
new species. At the same time it is necessary, in order to avoid
confusion in description, to associate some name with each of
these casts since they represent at least different species, if not
genera. Since we owe to Professor Elliot Smith our first detailed
account of the brain in the Archzoceti, and especially the recog-
nition that “the differences” (7.e., between the endocranial cast
of Zeuglodon osivis and the natur al cast (M. 12066) in his hands)
“are sufficiently pronounced to indicate a generie distinction
between the two specimens”; I propose to term the form from
which this type of cast is derived Zeuglodon elliatsmithii, sp. nov.
The second natural cast (M. 12123)—the one which Professor
Elliot’ Smith has given to me for description—I will call
Zeuglodon sensilivus, sp. nov., since it was in this cast that a
marked hypertrophy of the trigeminal apparatus was first
recognised.
3. a. ZEUGLODON SENSITIVUS, sp. nov. (M. 12123.)
The most complete endocranial cast of a Zeuglodont yet
discovered is that represented by Stromer (1908) % sitw in the
skull of what he has there called Zeuglodon osiris. It is not
the Zeuglodon osiris of his earlier (1903) paper, but more nearly
resembles the .rozeuylodon «atrox Andrews or is intermediate
between Prozeuglodon atrex and the Zeuglodon intermedius of
this paper..
The: peculiar resemblance of the general configuration of the
fore-brain and olfactory peduncles in the Zeuglodont brain to
BRAIN OF THE ZEUGLODONTIDZ. 619
the corresponding regions of the reptilian brain was emphasized
by Elliot Smith (1903), and is shown to distinct advantage in
the lateral aspect of the natural endocranial cast (text-fig. 1) here
under description.
This appearance is accentuated by the apparently reptilian
‘“‘lobus olfactorius”’ formed by the anterior part of the cerebrum
in Stromer’s specimen. This particular resemblance is probably
superficial. Instead of passing with a gradual expansion into the
front end of the hemisphere as the stalk of a pear is attached to
the smaller end of the fruit, the olfactory peduncle in the
Zeuglodont is attached to the hemisphere on its ventral surface
in the characteristically mammalian fashion, in front of the area
of which the still prominent tubercula olfactoria form a part
(see text-fig. 3). Behind the position of the attachment of the
olfactory peduncle there is an obvious depression in the lower
and anterior part of the lateral aspect of the cerebral hemispheres,
which I consider to be a definite Sylvian depression (in the sense
Text-figure 1.
Medan tentorial
Cepression
Lobvs meds
Sagittal sinus
cerebell
Cerebral hemisphere. are *
ZG»
Vee
Olfactory. aN se R aa A i
eduricle ONE Res AS Re, yeah Wa Paratlocculus
eC ¢ tty Ny ~ c6erebel
Optic nerve SAU Qa een RF ‘
sored espe; SIN ALS Ae Jugulor leash
OQohthatruc / Ft) PY;
mesa "Maxillary eee Mezdeuley ‘eq
nerve. P : eA Acoustic nerve
Gasserian Migale cerebral
garghon. vesse/
Lateral view of natural endocranial cast of Zewglodon sensitivus, sp. nov. M. 12123.
About 4 nat. size.
that that term is used in the lower Mammalia). This view is
corroborated by the fact that the groove is occupied by the large
middle cerebral vessels. Stromer’s specimen shows not merely a
large portion of the endocranial cast, but also a mould of the.
whole interior of the brain-case, including the olfactory peduncle
. from its origin in the olfactory bulb in its insertion into the
brain. The length of the olfactory peduncle there is more than
double the antero-posterior length of the cerebral hemispheres
themselves, while the relative size and shape of the bulbar
terminal dilatation is clearly distinguishable.
I am able to recognise in this cast (M. 12123) that the optic
nerves. formed the basal angles of Elliot Smith’s “ prismatic
olfactory peduncle” (vide “anterior view” inset to text-fig. 2).
Although this involves a considerable reduction of the actual
dimensions of the olfactory peduncles in cross-section, as stated
by Elliot Smith, it is still obvious that the sense of smell was of
significant importance to the Archeoceti.
620 - PROF. R. A. DART ON THE ”
“ Zeuglodon ist mindestens zu den hemianosmatischen Siiuge-
thieren zu rechnen im Gegensatze zu den anosmatischen Zahn- und
Burtenwalen und ist: auch durch normal verlaufenden Nasen-
rachengiinge von letzteren unterschieden,” says Stromer. In his
insistence on the retention of a sense of smell by these creatures,
he is justified even though his term ‘ hemianosmatic” seems
philologically meaningless.
Along the dorsal border of the lateral aspect of! the cast the
olfactory peduncle appears to merge into the sagittal sinus, which
is continued over the cerebrum towards the anterior border of
the cerebellum, where it dips into the ‘median tentorial depres-
sion’ and receives veins from the cerebellar surface.
The cerebrum on either side is seen to be bounded by the
sagittal sinus above and by the cerebellum posteriorly (which
excludes all view of the mid-brain region from the dorsal or
lateral aspect). Ventrally there is to be seen anterior to the
region of the cerebellum designated “ paraflocculus,” a rounded
roll-like structure continuous anteriorly with the ophthalmic and
maxillary divisions of the trigeminal nerve. This bulging mass
is a huge Gasserian ganglion which rivals the cerebral hemisphere
in ‘size, helps to mould the lateral contour of the brain-case, and
gives rise to its three proportionately large trunks. Coursing
over the lateral aspect of the Gasserian ganglion (well illustrated
in text-fig, 1) are the middle cerebral vessels. Between these
confines (sagittal sinus above, cerebellum behind, and Gasserian
ganglion below) in Zeuglodon sensitivus the cerebrum bulges out,
displaying a perfectly smooth hemispherical surface with the
exception of the previously noted Sylvian depression.
Behind the cerebrum lies the apparently irregular mass of the
cerebellum elevated to a height of approximately 15 mm. above
the cerebrum. [From this it is separated by a tentorial sulcus,
which is of especial depth in the mid-line, forming a “ median
tentorial depression.” ‘lhe most obtrusive feature of the cere-
bellum from the lateral aspect is unquestionably the lobus floecu-
laris (consisting of the flocculus and the paraflocculus).
Elliot Smith has called this region the ‘“ paraflocculus” in his
description of the Zeuglodont brain, and this name is retained in
these figures. It seems that in most aquatic mammals the para-
flocculus is the portion of the lobus floccularis which undergoes
greatest expansion. The term paraflocculus may therefore be
regarded, for the purposes of this paper, as synonymous with
the lobus floccularis,
Despite Bolk’s association of the paraflocculus with tail-move-
ments, bears which have no tail (as Elliot Smith has pointed
out to me) have, nevertheless, a well-marked paraflocculus. But,
whereas the paraflocculus is present in most land-mammals, it
becomes especially hypertrophied in all marine Mammalia irre-
spective of the stock from which they have sprung. Thus
creatures so divergent as Otaria and Monachus (Carnivore),
Trichechus manatus (of Ungulate origin), and Phocena (probably
BRAIN OF TIE ZEUGLODONTIDA. ‘621
of Tnsectivore origin) all possess hypertrophied paraflocculi. This
convergent hypertrophy of the same cerebellar region must arise
from some cause common to all. Ferrier was one of the first to
emphasise the functions of the cerebellum as primarily concerned
‘with equilibration. Ingvar (1918) has supported this conception,
und he finds a certain pattern of equilibratory localisation in the
‘organ, ‘This pattern is that of a ‘compass,’ a lesion in any
‘given part of the cerebellum entailing a defect in co-ordinating
‘muscle-movement in such a way as to resist falling in that
direction ; 7. e., the animal falls in the direction indicated by the
site of the lesion. This theory seems adequate to account for
the parafloccular expansion exhibited by marine Mammalia, for
Text-figure 2.
Oey, Anterior view.
P' Olfactory peduncle.
Optic nerve.
egittal sinus.
Cerebral hemisphere.
V2
Vy,
a vA . .
ZY f mm ganglion.
i LY; / : Gasserran ganglio
p tuft
fi a, Uy
Meovsr tentorial |, fi
depression. rtf
em 4 Wy vesse/,
Piece of
matrix,
Acou-tic
nerve.
Psratlocculus, (ash, Lobus medius cerebellt
Dorsal and anterior views of natural endocranial cast of Zeaglodon sensitivus, sp. nov.
“M. 12123. About } nat. size.
the paraflocculus is situated entirely laterally in the cerebellum,
and these creatures are.under the necessity of resisting con-
tinuously the tendency to “rolling,” which the fluid medium
postulates. In brief, it is a mechanism evolved to preserve
an even keel. is
The ‘“ paraflocculus” is bounded above by the region which in
the subsequent account is referred to as the lobus medius.
‘Anteriorly, the paraflocculus abuts on the cerebrum and the
Gasserlan ganglion; inferiorly, upon the eighth nerve and the
petrous temporal and the casts of the foramen lacerum medium
and the foramen Jacerum posterius (‘jugular leash” of the
figures). Posteriorly, it is in contact with, the exoccipital bone.
622 PROF, R. A. DART ON THE
Medially from this region can be seen an elevation of. the lobus
medius of the cerebellum, which is the most posterior structure
from the lateral aspect. It probably corresponds to the region
which gives rise to the cerebellar tonsil of human anatomy.
Text-fig. 2 presents all those features emphasized by Elliot
Smith (1903) in his specimen (Zeuglodon elliotsmithit of this
paper). This natural cast reproduces very faithfully the convo-
lutional pattern of the cerebellum and the position of the various
meningeal vessels, particularly the sagittal sinus. ‘This latter
Text-figure 3.
OlFactory pedurcle.
Al Optic nerve.
! | x Oohthalmie. division
Pt Nat of trigeninus.
mine:
Maxillary Wviston
AN SF Crigemnus.
\ 1% /ntertubercular sulcus.
Tubercu/um olfactorium.
WN, Gasserian ganglion.
x f
INe-7Tuber cincereu/m.
\
, Vandi bular
WW ovssvo77 of
Medulla oblongata.
‘Ventral view of natural endocranial cast of Zeuglodon sensitivus, sp.nov. .M. 12123.
About } nat. size.
structure was described by Elliot. Smith (1903) as the “ dorsal
rostrum.” ‘The knob-like elevations along its course are appa-
rently veins entering it from the diploe of the skull. hey bear
an extraordinary resemblance to human Pacchionian bodies.
Posterior to the prominence of the sagittal sinus and separating
it from the cerebellum is again seen the deep “ tentorial median
depression ” already referred to, from which the tentorial sulcus
runs laterally on each side separating cerebrum and cerebellum.
From this aspect, too, the cerebellum is seen in complete detail—
BRAIN OF THE ZEUGLODONTID. 623
the enlarged lobus floccularis on either side and in the mid-region
the so-called lobus medius. The lobus medius of the cerebellum
is shown to be markedly asymmetrical. A bilaterally symmetrical
projection into the posterior part of the tentorial median depres-
sion possibly represents the lobulus simplex of Bolk. If so, it is
very small, and the lobus anticus is completely hidden from sight
underneath it.
The hemming in laterally and anteriorly of the hemispheres at
their ventral margins by the Gasserian ganglia is obvious from
this aspect—forming a unique arrangement for a mammal. In
no other mammal have these ganglia been described as visible
from the dorsal aspect of the brain. The ganglia therefore are
peculiar as constituting a factor in the modelling of the roof of
the cranial cavity.
The ventral surface of the natural cast (text-fig. 3) gives im-
portant confirmation of the inferences drawn already. Through
this cast we are able to infer the extent and size of the huge
Gasserian ganglia. With these must be related a correspondingly
extensive area of grey matter in the medulla oblongata and cord
(substantia gelatinosa Rolandi), . Lhe resemblance of this aspect
of the casts to the ventral surface of the brain of Ornithorhynchus
(which is the only mammal that provides a suitable comparison,
by virtue of its similar functional specialisation) is undoubtedly
the clue in this arrangement. Anteriorly, the ophthalmic trunk of
the trigeminus overlaps the optic nerves and olfactory peduncles.
The course of all three structures is parallel for some distance
until the region of the tuberculum olfactorium is reached. The
maxillary division of the trigeminus is there attached to the
ganglion? Between the ganglion and the tuberculum the optic
nerve (not visible as a distinct elevation on the surface of the
cast) must have skirted the lateral aspect of the tuberculum till
it reached its posterior margin, where it bent medially to meet
its fellow of the opposite side in the optic chiasma, the position of
which can be recognised at the posterior end of an ‘“ inter-tuber-
cular sulcus” lying in the mid-line between the two tubercula.
Directly posterior to this point there is a single median elevation
present in the three casts. It is obviously the site of the
strongly-marked tuber cinereum and hypophysis. Behind this
region the Gasserian ganglia diverge, giving place first of all to a
slight central depression, then to a broad flat surface. This wide
divergence and a bulging of the cast on either side of the mid-line
may be due in large measure to the forward jutting of a tuberculum
quinti upon either side correlated in size with the enormous
Gasserian ganglia.. In the cast we have no indication that an
elevation due to a pons Varolii, as such, was present, and probably
it possessed no larger pons than the Prototherian. The “ bulging
of the cast” postero-medial to the Gasserian ganglion may there-
fore be interpreted as the upper portion of the tuberculum quinti.
Behind it lies a transversely running depression in the region
where the pons would normally lie. It is possible that this
624 . ~PROF. R. A. DART ON THE
depression is due to the constricting influence of a small pons, such
as was probably present... In other words, the substantia gelatinosa
expanded anterior and posterior to this site to form the promi-
nences evident in the cast. More posteriorly the base of the
brain passes gradually into the medulla oblongata. In many
mammals (e.g., Ungulata) the transverse width of the medulla
oblongata is very appreciable, but in these creatures the trans-
verse width is never so great (relatively) as in animals (e. g., Orni-
thorhynchus) which rely to a greater extent upon the “ fifth nerve»
sense.” The great increase in transverse width found in the
Zeuglodontide is to be regarded as a corroboration of the obser-
vation already made concerning the size of the tuberculum quinti.
In the cast there is no trace of the origin of either the third,
fourth, or sixth cranial nerves. The origin of the seventh. and
eighth is also obscure, though the latter is exposed for a portion
of its course through the petrous portion of the temporal bone.
‘The cast ends abruptly before the emergence of the 1X, X, and
XI complex of cranial nerves. The study of the skeletal parts
‘Text-fignre ‘4,
Lobus meaius cerebell.
Farattoccu/us.
Broken mergin
“A OF cast A
- Posterior view of natural endocranial cast of Zeuglodon sensitivus, sp. nov, ;
.M. 12123. About } nat size.
confirms what has been stated concerning the splaying apart
of the Gasserian ganglia (and incidentally of the posterior parts of.
the cerebral hemispheres and the lobi flocculares of the cerebelluin)
by the. expansion of the tubercula quinti. ss
No new anatomical feature from this aspect (text-fig. 4) is
revealed. The natural cast has been severed transversely at the
hinder end of the cerebellum. The important fact to be remem-
bered is that the lobus anticus and lobus posticus are probably,
totally hidden from external view ; and the central mass of the
cerebellum, as exhibited in dorsal or posterior view, is composed.
entirely of the lobus medius (of Elliot Smith), while laterally on
either side lies the lobus. floccularis. _ Posteriorly to the acoustic
nerve and ventrally from the lobus floccularis (* paraflocculus ” of
the text-figures) is to be seen an ill-defined mass, which represents.
the cast of the jugular vein and the associated nerve-structures,
which finally emerge at the foramen lacerum posterius. I have
called this region the “jugular leash.”
BRAIN OF THE ZEUGLODONYID.E. 625
3.6, ZEUGLODON ELLIOTsMITHI, sp. nov. (M. 12066.)
Turning now to the lateral view of Zeuglodon elliotsmithii,
M. 12066 (text-fig. 5), and comparing it with Zeuglodon sensi-
tivus (text-fig. 1) we recognise in both the general Zeuglodont
characters, which may be summarised as including a hyper-
trophied cerebellum tending to grow forwards over the cere-
abrum, hypertrophied trigeminal apparatus, and a diminutive
reptilian-like cerebrum bounded by the two former behind and
below, and an unusually well-marked sagittal sinus above and in
the mid-line.
It will be noted that in Zeuglodon elliotsmithii the fore brain
is very flattened and slopes upwards and backwards evenly
towards the cerebellum, that the cerebellum is only slightly
elevated (1-10 mm.) above the flattened cerebrum, and that
appears almost vertical from the Jateral aspect. In all these
features this brain contrasts strongly with that of Zeuglodon sensi-
twus. Further, the trigeminal apparatus, although hypertrophic,
*'Text-figure 5.
Sagittal sinus
Lodus mediuvs:
cerebelli.
Ofactory peduncle.
i Paraflocculus.
- Trigenrus.
Gasserian
broken margin of cast... ganglion. Middle cerebral vessels. t
Lateral view of endocranial cast of Zeuglodon elliotsmithi, sp. nov. M. 12066.
About 4 nat. size.
does not seem quite so pronounced as in Zeuglodon sensitivus,
although the fragmented nature of the cast, in this region, renders
a final statement upon this particular point impossible.
The features to which reference has been made are to be recog-
nised from the dorsal aspect also. Note further from this aspect
that the lobus medius cerebelli of Zeuglodon elliotsmithit is sym-
metrical, whereas the other is markedly asymmetrical (cf. text-
fig. 2), a point which is also clearly evidenced in the posterior
views (text-figs. 4 and 7).
As these figures have been drawn to scale it is clear that many
features, such as the general flattening of Zeuglodon elliotsmithit,
shown by the increased bi-parafloccular width, its general triangular
outline as seen from the dorsum, and its slender transversely-
elongated cerebellum mark the brain as characteristic and distinct
from that of Zeuglodon sensitivus. These facts are brought out
by a comparison of the table of brain-measurements (vide infra):
Thus both the greater width and length of the. cerebral hemi-
spheres emphasize the flatness already referred to, while both the
626 PROF. R. A. DART ON THE
total length and height of the brain-mass were considerably
smaller in this species.
The bulk of the endocranial cast (average of five separate
measurements by water displacement) of Zeuglodon elliotsnuthi
Text-figure 6,
Olfactory peguncte.
Trigeminus.
Middle cerebral
vesse/s.
Mik ee
a < cy j
oo it wr ese
Dorsal view of endocranial cast of Zeuglodon elliotsmithii. M. 12066.
About 4 nat. size.
was 300c.c. Elliot Smith gives (1903) 410c.c. as the bulk of
this ‘natural cast (including that of a considerable quantity of
matrix attached to the base of the brain and some small frag-
ments of bone),” but states that the actual weight of the brain
Text-figure 7.
Lobus Menus
cerebellt.
Broken margin of cast.
Posterior view of auleoranial’e cast of Zeuglodon elliotsmithii. M. 12066.
_ About $ nat. size.
wns neobally “nearer 300 grammes.” In this latter estimate I
aim inclined to agree. But whereas the bulk of Zeugledon elliot-
smithii could not have been very much greater than 300 c.c., that
of the natural cast of Zeuglodon sensitivus (in which the medulla
-
BRAIN OF THE ZEUGLODONTIDA. 627
oblongata is absent) was 490c.c., or nearly 200 c.c. larger. . This
is of some interest not only in demonstrating the specific (if not
generic) difference between these two Zeuglodonts but hlso in
showing that the brain-capacity of certain Zeuglodonts was con-
siderably greater than that of some existing Cetacea—because
the brain-weight of Kogia (Haswell) is only 455 grammes.
All these distinctions between these two forms are certainly
not to be accounted for by a difference in the sex or age of the
individuals. They should therefore be separated as distinct
species, as has been done here, until fresh data shall be forth-
coming when it may be possible to associate one or the other of
these forms, Zeuglodon sensitivus or Zeuglodon elliotsmithi, with
the species now recognised from skeletal parts alone, such as
Zeuglodon zittelr. :
4. THE THREE ENDOCRANIAL CASTS FORMING A PHYLETIC SERIES.
4, a. Prozeu@Lopon ATROox Andrews. (M. 9265.)
In the ‘Descriptive Catalogue of the Tertiary Vertebrata of
the Fayum of Egypt,’ Dr. Andrews described briefly the out-
standing features of this cast, pointing out its correspondence
with Elliot Smith’s general account of the Archzoceti (1903).
It is unfortunate that in this important cast from the lowest bed
Text-figure 8.
Lobus medius cerebelli:
fi,
WU
y yy’
vessels. Paratlocculus.
Case Jugulsr least
i?
ganglion. Acoustic nerve.
Lateral view of endocranial cast of Prozeuglodon atrox Andrews. M. 9265.
About } nat. size.
of the Middle Eocene there is no reproduction of the anterior
portion of the cranial cavity, because this region would supply
important data concerning the degree of trigeminal specialisation
in this early representative of the group—data which now must
be inferred from the evidence afforded by other regions.
It is astonishing to find in this earliest-known Zeuglodont an
extravagantly expanded cerebellum. It is the dominant portion
of the brain from lateral, dorsal, or posterior aspects. It rises at
Proc. Zoon. Soc.—1923, No. XLI. 4]
628 PROF. R. A, DART ON THE
certain’ points taken along the tentorial suleus to a height of
35mm. above the cerebral surface and nowhere is it less than
Text-figure 9.
Sagittal sinus.
Median tentorial
Flattened cerebrum.
Middle cerebral epression.
vessels Covering
Gasserian
ganglion. Tentorial
2 OU
(eo be Ne "fe iN :
Zig Pe
EE
—%
Nedullz oblongata. — Wt
Dorsal view of endocranial cast of 4.a. Prozeuglodon atrox Andrews. M, 9265.
About } nat. size.
20 mm. higher than the cerebral surface. This one characteristic
immediately brands this brain (however Zeuglodont in type) as
something entirely distinct from what has hitherto been described
Text-figure 10.
Lobus medius cerebell:.
Posterior view of endocranial cast of Prozeuglodon atrvox Andrews. M. 9265.
About 3 nat size.
in the literature. The size of the cerebellum is immense and the
size is not confined to one part of the organ only; the parafloc-
BRAIN OF THE ZEUGLODONTID®. 629
‘culus is hypertrophied, but so, too, is the lobus medius and, in
addition, the latter is markedly asymmetrical.
These facts are readily appreciated from the figures. The
lateral aspect is instructive as revealing the relative size of the
‘cerebrum and the cerebellum. Coursing over the latter from the
top to lateral angle is a ridge, probably indicating the site of a
venous channel, which converges upon the jugular leash posterior
to the acoustic nerve.
From the posterior aspect we are impressed with the vertical
“lie” of the paraflocculi and their size relative to the lobus
medius. The massive “jugular leash” undoubtedly accommo-
dates the venous channels and the posterior cerebral nerves.
The dimensions of the medulla oblongata should be noted for
comparison with the later forms to be described.
Text-figure 11,
Intertubercular sulcus.
Gasserian
- ganglion. Tuber cinereus.
uiv 4 >
PE Psraflocculus.
Acoustic
nerve.
Yugular le2sh.
Nedulla obfongata.
Ventral view of endocranial cast of Prozeuglodon atrox Andrews. M. 9266.
About 4 nat. size.
The ventral view of Prozeuglodon atrox should be compared
with text-fig. 3. It will then be recognised that the structures
-on the basal surface of Prozeuglodon atrox are entirely comparable
with those found on the basal surface of Zeuglodon sensitivus.
It is typically Zeuglodont in character—a fact which we will find
to be of great significance in comparing these endocranial casts
with that of Prosqualodon. No features call for special attention
at this stage of the discussion other than those noted in the
figures themselves.
4, 6. ZEUGLODON INTERMEDIUS, sp. nov. (M. 10173.)
‘Zeuglodon intermedius from the middle beds of the Middle
Eocene shows from the lateral aspect features comparable. in
41*
‘630 : PROF, R. A. DART ON THE
most points with those of Prozeuglodon atiox, save for the more
marked sagittal sinus, larger Gasserian ganglia, and a greater
backward thrust of the paraflocculus.
From the dorsal aspect it is evident that there is a greater
Text-figure 12.
Cerebral henus27reé.
Lobys medus.
cerebelli.
Olfactory -
peguncle.
Medullz
(E29 guissons
HEUER) Jugular leasis.
Gasserian ganglion Widdle cerebral vessels
Lateral view of endocranial cast of Zeuglodon intermedius, sp, nov. M. 10173.
About 4 nat. size.
lateral thrust of the paraflocculi also, a fact still more emphasized
in the posterior view, The prominence of the sagittal sinus 1s.
Text-figure 13.
Olfactory peauncle.
Maxillary division
of trigeminus.
Sagitta! sinus
Trigermaus & mowsie
cerebra/ vessels.
Ze Zi, GG / )
his fe il hy hte me
MANNIE cel
WAS Gav yall : Parafloccul
SSS S araflocculus
= ule uy if ' :
WAY
A\l
‘, , » ti
Dorsal view of endocranial cast of Zeugludon intermedius, sp. nov. M, 10173..
About } nat. size.
well shown from the dorsal aspect also, but the median tentorial
depression is not quite so pronounced.
An arresting feature of this cast is the transversely running
furrow subdividing the central mass of the cerebellum and
BRAIN OF THE ZEUGLODONTIDA, 631
disappearing laterally below the paraflocculi. Here it separates
the paraflocculus above from the structures of the jugular leash
below. It is due to a kind of secondary tentorial bony projection
on the interior aspect of the occipital bone. The presence of this
bony strut is perhaps to be associated with the increasing width
of the occipital region, This furrow is wider and more dorso-
ventrally situated in the cast of Prozeuglodon atrox; whereas in
Zeuglodon osiris (vide infra) it is still transverse but very wide
and scalloped out, showing that with the extreme width of the
skull of Zeuglodon osiris the bone has become greatly reinforced
in thickness internally. It is well to note such cases as these
Text-figure 14,
Lobus medius cerebells.
Juguler
ap
oblongata.
Posterior view of éndocranial cast of Zeuglodon intermedius, sp.nov. M. 10173.
About 4 nat. size.
where the modelling of the skull undoubtedly influences the
shape of the endocranial contents, because it shows that the inter-
play of factors (¢.¢., brain upon skull and skull upon brain) in
cavity-modelling is ceaseless and reciprocal.
Coincident with the lateral expansion of the brain-substance
there has been no widening of the medulla oblongata or the
aperture of the foramen magnum (vide Tables), but the thickness
of the medulla oblongata has decreased somewhat and so it
presents a somewhat more ellipsoidal transverse section.
The cubical capacities of these two casts was 790 c.c. for
Prozeuglodon atrox and 785c.c. for Zeuglodon intermedius. As
the cast of Prozeuglodon atrox was very incomplete anteriorly it
is evident that its cubic bulk must have been considerably over
800 c.c.
The degree of likeness between these two forms both cranially
and endocranially is so great that they may be regarded as very
closely related phyletically. Because of this the decreased volume
of the brain of Zeuglodon intermedius, in accordance with the
‘law of increasing brain weight” expressed by Marsh, indicates
632 / PROF, R. A, DART ON THE
a, degenerating condition in this later Zeuglodont. type. When
we realise that part passu with this decrease in volume there
is an actual increase in the lateral thrusting of the brain-mass.
and with it of the Gasserian ganglia, and a relative flattening
of the mass of medulla oblongata, it seems clear that this
degeneration is primarily trigeminal in nature,
4, c. ZEUGLODON ostris Dames. (M. 10228.) |
A comparison of the lateral aspect of the cast of Zeuglodon
osiris from the upper beds of the Middle Eocene with the pre-
ceding forms is very striking. One gathers the impression that
the brain-mass is shrivelled and shrunken, a conception that is
strengthened by examination of the dorsal and posterior views.
If we take the features seriatim we find first of all that the
Gasserian ganglion is relatively small as compared with the other
forms, but its stranded position (laterally to the cerebrum—see
dorsal view) shows that this creature is descended from a type in
Text-figure 15.
Cerebral hemisphere.
Lobus medius cerebeli, -
Olfactory
peduncle,
Trigeminus.
é ; ob/ongata.
| asserian Spon Jugular leash.
Lateral view of endocranial cast of Zewglodon osiris Dames. M. 10228.
About } nat. size,
which the trigeminal apparatus was of a much more imposing
nature, The “gouged out” cerebrum reveals a degenerate con-
dition of this organ. Obviously the olfactory region has become
even more atrophied than in the other known Zeuglodonts.
Despite the opportunity offered by this atrophy anteriorly and
the diminished size of the cerebellum posteriorly the cerebrum is
diminutive.
The cerebellum is still large but it is crumpled. The para-
flocculi no longer bulge over the acoustic region, but are
contracted towards the lobus medius.. The ‘latter is very
asymmetrical. The space. between the middle cerebral vessels
and the jugular leash, and lateral to the paraflocculus, seems to
have been devoid of brain-substance in the recent condition and
to have been occupied by venous sinuses.
BRAIN OF THE ZEUGLODONTID. 633
The lobus medius, or area crescens, is not only very asym-
metrical but is poorly developed,and the paraflocculi form by far the
| greatest. part of the cerebellar mass. ‘The posterior view ‘affords
' perhaps the completest picture of this atrophic brain, for here we
recognise the general flattening out or “‘ cake-like” appearance of
Text-figure 16.
Olfactory peduncle.
ew Sagittal sinus.
Wi, IK "
“A SS
Trigeminus. Trigeminus ~
Cerebral hen sphere
Middle cerebrs/
vesse/, , Gasserian ganglion
& middle
‘f"\ cerebral vesse/s
wh
an LS Fa
< we IS NL Pareflocculus.
PS.
INAS ON SN
Jugular leash.
Me
Wedulla oblongata.
Dorsal view of endocranial cast of Zeuglodon osinis Dames. M. 10228.
About + nat. size.
this brain (which came from a perfect skull) and also the
extremely long but narrow cross-section of the medulla oblongata.
‘This view of the medulla oblongata coupled with the lateral view
Text-figure 17.
Lobus medius cerebelli.
Parstlocculus. , Ze
AK RR
: WW Ree a4 , Yh ) N ah Opals
4!’ er WW)! armor nti,
gr \ Yb, wr i ff,/s oo R ROK AA
cs < Ky | = =< : =a LLL: = hy, \\ 1)
PENN ; nN We Zz
TE a nil ul Z
, _ Medulla oblongata.
" Posterior view of endocranial cast of Zeuglodon osiris Dames. M. 10228,
About % nat. size,
of alg region corroborate the “aflmanes we have already drawn
that in Zenglodon osiris we have the degenerate ciispr ing of @
specialised race.
The degree of this degeneration is perhaps most graphically
shown if we compare the cubic capacity of this brain (48(c.c.)
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BRAIN OF THE ZEUGLODONTIDA. 635
with those of the foregoing which are almost twice the size.
An examination of the tables and the diagrams of the skulls will
show that this discrepancy in brain-capacity is not due to any
diminished bodily size in Zeuglodon osiris, for although the skull
may be somewhat shorter it is absolutely wider than those of the
other two forms.
One can only conclude that we have here an obvious dwindling
in brain-substance which has affected not only the trigeminal
and olfactory regions but with them the cerebrum, cerebellum,
and the medulla oblongata. The whole evidence goes to show
that the devolutional potentialities exhibited in the specialisations
(chiefly trigeminal) of Prozeuylodon and the group it typifies have
found in Zeuglodon osiris (M. 10228) their logical finale.
6. Tur ZooLocicAL Posirion oF THE ZEUGLODONTIDE.
Certain facts arising from our study of these endocranial casts
claim our immediate attention. The first of these is the tri-
geminal specialisation.‘ In all of the Zeuglodonts (although
coming from different horizons) the essential features noted in
Zeuglodon sensitivus apply. In all, the Gasserian ganglion assists
in modelling the roof of the cranial cavity. The inference is
justified that the group, as a whole, rested from the outset upon
this specialisation for its subsequent achievements.
It seems to be a law of general evolutionary application that
specialisation of one “sense” entails as its corollary the atrophy
of one or more of the other avenues of sense-perception. Thus,
birds become specialised as to sight but lose their appreciation of
smell; the same is true, though in different ways, for Teleostean
fishes and for Primates. In Ornithorhynchus, which specialises
in its “‘ fifth nerve sense,” we find a relative atrophy of both smell
and sight. In Zeuglodontide the sense of smell has certainly
been largely lost as a result of their adoption of a water habitat,
and it seems likely that sight also was of diminished value.
To the loss of smell and the abortion of the basal (olfactory) parts
of the cerebrum with their effect in limiting the longitudinal exten-
sion of the hemispheres and consequently of the cranial cavity,
reference was made by Elliot Smith (1902 and 1903). A relative
loss of sight, with a resulting diminution of the mid-brain and
thalamic regions, may assist in accounting for the smallness of the
cerebrum in Prozeuglodon and also for its lack of growth in the
two later forms from the upper horizons. It assists in the under-
standing of the propinquity of the cerebellum to—or, rather, its
overgrowth over—the cerebrum, and affords further reason for
the diminished longitudinal extension of the cerebral axis (because
of a wasting mesencephalon and optic thalamus and the con-
sequences entailed thereby) throughout the series.
But it seems to me that the lateral expansion not merely of
the brain-stem itself but also of the cerebellum has been provoked
mainly by the trigeminus, and that a specialisation of this nerve
has been the most significant factor in determining the queer
636 . PROF, R, A, DART ON THE
contour of the Zeuglodont brain. Of course, the enlarged cere-
bellum has played a role in the lateral expansion of the cranial
cavity, but the extraordinary dimensions, even for an [ocene
animal, of this cerebellum can only be explained by the tribute
coming from trigeminal nerves whose Gasserian ganglia rival the
cerebral cortex in size. Hence the influence of the cerebellum is
fundamentally trigeminal in origin.
It is true then that, although the trigeminus has played the
chief part in this expansion, the internal factors include also an
olfactory and a visual factor and may well have been assisted in
Zeuglodonts, as Andrews (1908) suggested, by the “pressure
on the anterior end, more or less in the direction of the long
axis,” during motion through the water, “and during very rapid
movement this pressure must be considerable.”
Now a reduction in value of the sense of smell and possibly of
sight and a compensatory dependence upon the sense of touch in
the muzzle—i, e., a certain degree of trigeminal specialisation—
are equally to be anticipated in the ancestors of modern Cetacea.
Might it not be that the Zeuglodontida, in this respect as in their
osteological conformity, are to be regarded as ancestral to Cetacea ?
This might be admitted if it were not for the demonstration of
the already-marked specialisation of Prozeuglodon atrow and the
gradual deterioration by specialisation within the phyletic
series as already discussed. In this light the passage of Cetacean
ancestors through a “trigeminal” stage in evolution can only be
cited as convergence, or may be explained by the hypothesis that:
the Zeuglodontide and the true ancestors of Cetacea may have
had a common ancestry in the earliest; Kocene.
Despite the positive views concerning the ancestral relation-
ship of Zeuglodontide to Cetacea stated at the outset, it must
not be assumed that paleontologists are agreed concerning this
interpretation.
As early as 1877 Marsh said: “That the connection (between
Zeuglodon and Cetacea) was a direct one, however, is hardly
probable, since the diminutive brain, large number of simple
teeth, and reduced limbs in the whales all indicate them to be an
old type which doubtless branched off from the more primitive
stock leading to the Carnivores.”
Weber (1886, p. 243) also referred to Zeuglodon as ‘einen verun-
gliickten Versuch Cetaceen herauszubilden,” and although there
have been wavering opinions by many since, which even Weber
himself has shared, many, such as Miiller, Fraas, and Stromer,
have enunciated the same view.
Fraas, in 1905, summed up the evidence to that date available
and showed that the Zeuglodontide, arising from some possible
Creodont stock, have passed through some stage corresponding
to his /ocetus (Mesocetus) and, by differentiation, have produced
various forms, including the gigantic ones which generally have
marked the acme of evolutionary progress in many groups of
animals, after which the race has disappeared. He has felt it
BRAIN OF THE ZEUGLODONTID&. 637
impossible to regard such later gigantic forms as ancestral to:
the later Squalodon series of animals, which appear to be more
definitely related to the progenitors of the modern Cetacea.
According to Winge (1919, 1921) Fraas “‘considered both Proto-
cetus and with it other ZGeuglodonts as a side branch from the
Carnivores which did not lead in the direction of true whales.”
Stromer (1908, p. 174) also states: ‘Ich halte also einstweilen
fiir geboten die Zeuglodontidze fiir eine ihnliche Parallelreihe der
Denticeten anzusehen, wie sie neuerdings innerhalb vieler engeren
Saiugethier-abtheilungen nochgewiesen wurden. Sie hatten schon
im Qbereézan ihre Bliitezeit unter Entwickelung von Riesen-
formen wahrend die anderen fast gleichartig aber viel weiter sich
differenzierend langsamer sich entwickelten und wieder in mehrere
Zweige auseinandergingen, die auch in vieler Beziehung einander
parallel fortliefen. In diesem Sinne also nehme ich wie Weber
(1886, S. 243) Zeuglodon als ‘einen verungliickten Versuch
Cetaceen herauszubilden’ halte aber fiir noch nicht beweisen dass
alle Archeoceti so auf zufassen waren.” ° .
* Abel, in his later works (1913), does not appear to have any
doubt that the Zeuglodontide lie off the direct line, with the
possible exception of the so-called Microzeuglodontide. Because:
of their great specialisation, the degree of which would entirely
unfit them for such an evolution, I am entirely in agreement
with the opinion that no Zeuglodont here examined can be
regarded as ancestral to Cetacea. ;
It is a curious fact that these highly specialised animals have
so wide a geological distribution, and one in favour of their
marine life. This wide distribution may have occurred before
the trigeminal specialisation, so characteristic of all these Fayum
forms, was attained. This makes the examination of the endo-
cranial casts in Zeuglodontide found elsewhere in Europe, in
America, and in Australia of the greatest importance because the
facts put forward demonstrate our lack of knowledge concerning
the evolutionary history of the whales and also because “the
distribution of the Zeuglodontide and other shallow-water fauna”
has been used (Stromer, 1906, and Andrews, 1907) to indicate
the location of the shore-lines of previous land-connections
between the Old and New Worlds. I consider that in the
demonstration of this tactile specialisation in Zeuglodonts their
restriction to a shore-line distribution is shown to be highly
probable. .
The parallel origin of the Zeuglodontide and the Sirenia— ,
probably in the Lower EHocene—is very striking. The grade of
cerebral organisation in both is approximately similar, but the
cerebellum of Zeuglodontide seems to indicate that its life was
somewhat less retired than that of Sirenia. Andrews (1907) has
shown that “freedom from competition and, to some extent, from
powerful enemies, would offer: exceedingly favourable conditions.
for the rapid spread and multiplication” of these groups in the
Eocene seas. ‘Thus both groups became widely disseminated, but.
638 ‘PROF. R. A. DART ON TUE
whereas the Sirenia have persisted as “living fossils,” despite
their humble grade of intelligence, the Zeuglodontide have long
since disappeared.
This disappearance is to be correlated with the fact that,
whereas the Sirenia chose an eminently retired and sluggish
mode of existence, the Zeuglodontide were more active. They
came into more direct conflict with other marine forms and,
specialised as they were, did so.at a disadvantage and were over-
whelmed in the struggle for existence.
The present study therefore indicates that the scepticism of
Marsh and Fraas was entirely justified and that Winge (1921)
is incorrect in regarding the Zeuglodontide as ancestral to
Cetacea. The origin, dispersal (into so many strikingly different
forms and all over the globe), and disappearance of this group
geologically is a demonstration of the conception put forward by
Marsh (1877) that ‘In every primitive type which was destined
to survive many geological changes there seems to have been a
tendency to throw off lateral branches which become highly
specialised, and soon die out, because they are unable to adapt
themselves to new conditions,” .
7. CoMPARISON OF ZEUGLODONT AND ProsquaLopont BRAINS.
The above paper was practically ready for the press when
Prof. Elliot Smith received from ‘Tasmania, through the great
courtesy of Prof. Flynn of the Zoological Department in the
University at Hobart, a splendid cast of the cranial cavity of
Prosqualodon davidi which he has recently described. Plaster
replicas of the skull from which this cast was made were pre-
sented by him to the British Museum of Natural History and
to the Zoological Department of University College, London. I
have had the privilege of studying these casts and the endocranial
cast.
The lateral view of this cast (text-fig. 18) reveals an astound-
ing likeness to the Zeuglodont endocranial cast. We meet with
the same cerebellar enlargement, expanding forwards over the
cerebrum, and quite an enlarged trigeminus.
It is obvious that such striking similarities would not exist in
the absence of some close relationship between these forms, but
the data will reveal that this relationship is not a filial one as it
has frequently been eonceived.
It is to be noted that the paraflocculus is recognizable laterally,
but is submerged by a widely and generally expanded cerebellum.
The cerebrum is emancipating itself from the cerebellar growth
as its great height posteriorly shows. Anteriorly the rapidly
dwindling olfactory apparatus is attached in a “ nipple-like”
fashion to the cerebrum and considerably ventral to it is a
well-marked optic chiasma. The middle cerebral vessels reveal
a course comparable with that seen in the Zeuglodonts and
posteriorly there is a well-marked jugular leash.
BRAIN OF THE ZEUGLODONTIDS. 639
The dorsal view corroborates what we have already detected
from the lateral aspect. The wide expanse of the “area crescens”
cerebelli is’ the most obtrusive feature and after that the
expanding cerebrum. ‘The double olfactory peduncle is clearly
seen but no Gasserian ganglion is visible (text-fig. 19).
The posterior view is instructive in revealing an enlarged oval
transverse section of the medulla oblongata and still further
illustrates the nature of the cerebellar expansion (text-fig. 20).
Perhaps the most informative of all is the ventral view, because
we are able to recognise the true size of the trigeminus (which
is quite large), the relatively enormous width of the base of the
brain, the marked development of the optic chiasma, and the
relative atrophy of the olfactory apparatus. There is an entire
absence in this brain of anything corresponding to the tuber-
culum olfactorium or “ intertubercular sulcus” of the Zeuglo-
donts, but we find a ridge on either side of the middle line medial
Text-figure 18,
Lobus medius
cerebel/t,
© (generally expended)
Medulla
ob/ongatd.
Olfactor.
ve
pecuncle.~-
Optic
wa Juguler
leash.
Trigeminus. ae
Middle cerebral vesse/s.
Lateral view of endocranial cast of Prosqualodon davidi Flynn.
About 4 nat. size.
to the trigeminal region which may be due to the carotid artery
and accompanying venous sinuses. ‘The tuber cinereum is not
fay in the cast, but must lie between these ridges (text-
fig. 21). :
” Prosqualodon davidi Flynn comes from the Miocene deposits of
‘Tasmania, and the characters which it presents linking it to the
Zeuglodonts might well be interpreted as illustrating its origin
from the Zeuglodont family if our information were not so
complete as it is now.
The actual bulk of this endocranial cast is approximately
750 c.c., 2.é., not quite so great as that of Zeuglodon intermedius.
This fact itself is sufficient to show that the Eocene form which
gave rise to Prosqualodon must have possessed a brain-capacity
very much less than that of Prozeuglodon atrox or of Zeuglodon
intermedius. This follows from the well-known “ law of increasing
brain weight” put forward by Marsh and supported by all
‘640 - PROF, R. A. DART ON THE
paleontological endocranial investigation up to the present
time. ;
Quite apart from this there is ample evidence against the view
that a Zeuglodont, even one so primitive as Prozeuglodon, could
have given rise to Prosqualodon. This is very evident if we call
to mind those features pointed out in detail for Zewglodon sensti-
tivus (vide text-fig. 3) and repeated by all the Zeuglodonts. It
was shown in that case that the site of the insertion of the
olfactory peduncle in the Zeuglodonts had been drawn, as it were,
during atrophy under the fore brain on to the basal aspect. This
contraction of the area between the olfactory peduncles and the
tuber cinereum demonstrated for us the relative atrophy of the
e Text-figure 19. ,
hip eee! an ees ‘ee i Olfactory peduncle.
Cerebral hemisphere.
Trigemmnus.
Niddle :
cerebral! Lobus medius
vessels. cerebell,
(genera My
exparided)
Paratlocculus.
Medulla oblongata.
Dorsal view of endocranial cast of Prosqualodon davidi Flynn.
About $ nat, size.
-optic chiasmatie region in Zeuglodonts, and, fortunately, the cast
of Prozeuglodon atroz in this crucial region is sufliciently perfect
to illustrate the fact that these changes were already well marked
in the Zeuglodonts of the-earliest Middle Hocene deposits of the
Fayum (vide text-fig. 11).
In the case of Prosqualodon davidi, on the other hand, we find
that the course of its evolution has been entirely different,
There is evidence here of an initial expanding of this basal area
between the insertion of the olfactory peduncle (vide text-fig. 21)
and the tuber cineyeum (which lies somewhere between the
“ carotid ridges”). Crossing this wide interspace in Prosgualodon
-davidi we find in the well-defined optic chiasma the evidence of
BRAIN OF THE ZEUGLODONTIDA, 64]
ithe retention of elaborate visual capacities in the Miocene—an
utter impossibility in the offspring of the Zeuglodonts here
-described at any epoch.
It is of some value to appreciate how great a degree of
‘trigeminal specialisation is compatible with future evolution.
For it is evident that Prosqualodon davidi has an enlarged
trigeminus, even though we do not find such gross enlargement
of the Gasserian ganglia as in the Zeuglodonts. It has already
been indicated that “a certain degree of trigeminal specialisation”
is to be expected in the ancestors of Cetacea. It seems unques-
tionable that the initial widening of the Prosqualodont, as well
as that of the Zeuglodont, brain and medulla oblongata is due
not merely to the passive recession of the sense of smell but
rather to the active hypertrophy of the trigeminal apparatus,
which in an aquatic mammal provides so much more information
Text-figure 20,
Lobus medius cerebel/y,
(generally expanded)
|
\
Ai, \ ‘ hy
Paratlocculus.® : E ; i
Yee
Megullz oblongata. Irigemunus.
Posterior view of endocranial cast of Prosqualodon davidi Flynn.
About + nat, size,
‘concerning food, friends, and foes ‘than do the senses of smell
or sight,
Prosqualodon davidi teaches us therefore that the evolution of the
‘Cetacean stock, while it depended to some degree upon an initial
trigeminal specialisation, was not effected by any sudden reliance
‘upon this sense to the neglect of other important senses, but
depended upon an orderly and “ balanced” correlation. of this
hypertrophy with a concurrent aggrandisement of the visual and
auditory senses. In this connection it is significant that the most
‘expanded portions of the fore brain in Prosqualodon davidi appear
to be the “occipital” and “temporal”: regions; 7. e., posteriorly
and laterally, where one may reasonably conclude that these
senses were finding cerebral representation. Whether Fraas ix
‘correct in ‘believing that even the more primitive Protocetus
642 PROF, R. A, DART ON THE
is also off the main line of Cetacean evolution would probably be
indicated by an examination of the endocranial casts with regard
to these details,
The conformation of .the district between the olfactory
peduncles and the chiasma—in fact, the whole ‘ mamma-like”’
appearance of the antero-ventral portion of the fore brain in
Prosqualodon davidi, contrasting as it does with the same region
in the Zeuglodonts—is consonant with the homologous region in
true Cetacea, A comparison of this region with the ventral
aspect of the brain in the foetal Monodon or adult Kogia (vide
figures in Elliot Smith’s account, 1903) demonstrates the truth
of the conception that the ancestors of true Cetacea went through
Text-figure 21.
Carotid ridge.. » ye = OM Optic chiasma.
Sa SS
bi \N Trigeminus,
, yyy (Anterior
Middle cerebral NAG, 7 ‘seiseeuslf, ovisions.)
vessels. i jp LEX
fi: ‘ | VY ae
J \ Trigeminus,
OW WR, Mandibular
a\hy avision)
ie a
VMN! x
(eg
Vy, \S iN =)
aS
cS ———————
Jugular
leash,
Ventral view of endocranial cast of Prosqualodon davidi Flynn.
About 4 nat, size.
a stage of “ expansion of the basal regions of the brain” (witness.
the “desert region” of Broca in Cetacea) not displayed by
Zeuglodonts but evident enough in Prosqualodon.
What this expanded “desert region ” or tuberculum olfactorium
in whales exactly signifies, we are not as yet able to definitely
state. Since this area is the site of the palostriatal cortex
overlying the palostriatum (Dart, 1920) it follows that the
persistence of a large “desert region” is dependent on factors.
other than olfactory, 7. e., factors which have determined the size
of the paleostriatum itself. The consideration of what these
factors actually may be lies outside the scope of this article, but.
the essential point affecting our argument here is that these-
BRAIN OF THE ZEUGLODONTID. 643
factors, whatever their nature, played an enhanced réle in the
ancestors of Cetacea, whereas they were recessive and negligible
in the Zeuglodontide.
The evidence afforded by this cast therefore supports the
conclusions which we drew in the earlier part of this paper
concerning the deflection of the Zeuglodont line from the course
of true Cetacenn evolution, but supports the conception that
Squalodon has at least close allinities with the true Cetacean
stock.
8. Tur TRIGEMINUS AND THE LAw or INFILTRATION IN
CEREBELLAR Eyonurron.
In conclusion, it is necessary to call to mind certain general
facts about the cerebellum which appear to throw light upon the
strange and generalised hypertrophy of that organ in this group.
We know that animals with very sensitive whiskers and
bristles (such as certain Rodentia) depend largely for their
power of equilibration upon stimuli which arise in nerve-endings
situated in relation with the proximal end of these whisker-like
structures and affected by their faintest movement. Such stimuli
are communicated to the cerebellum by way of the trigeminus.
In Ornithorhynchus, instead of bristles, there is a remarkable
development of special receptive end organs in the delicate snout
served by the trigeminus. The bristles round the mouth-parts
of the Sirenia may be active in a similar way in addition to their
function as tactile end organs.
The more recent researches of Ingvar (1918) have placed the
morphological survey of the cerebellum by Elliot Smith upon an
even firmer basis: for he has extended his research into the
Reptilian and Avian series and has shown that here, too,
the same sulci and three lobes are distinguishable. Such a
division into three lobes does not appear to obtain in Fishes or
Amphibia. It seems therefore that the advent of the middle
lobe synchronises with the origin of a neopallium in the cerebral
cortex. ‘The researches of Haller (1900), of Unger (1906), and
of Crosby (1917) have shown that the first clearly defined
neopallium primordium makes its appearance in the Reptilia, and
in these creatures the tripartite structure of the cerebellum is
also clearly to be recognised for the first time.
Recognising the relationship which the development of the
neopallium has to the expansion of the cerebellum in Mammalia
it is, at first, most disconcerting to find in a group of Eocene
mammals with such ill-developed neopallium a cerebellum of
extravagant proportions which has not merely obliterated all
traces of the mid-brain from the surface but threatens to cover
the cerebrum also by its forward expansion. :
The cerebellum is usually relatively large in Eocene mammals
and yet such a bizarre arrangement as is present in Zeuglodonts
can have only one explanation. We may eliminate the neopallinm
Proc. Zoou. Soc.—1923, No. XLII. 42
644 PROF. R. A. DART ON THE
as an explanation because of its slight development and the
doubtful presence of a pons Varolii, and it is equally unlikely
that the spino-cerebellar contribution could have been so great
as to account for the whole of the cerebellar expansion. But
we do know that so diminutive and primitive a mammal as
Ornithorhynchus, with a well-developed trigeminal apparatus can
and does possess not only an unexpectedly large neopallium, but
also a relatively large convoluted cerebellum. The absolute and the
relative size of the Gasserian ganglia in Zeuglodon sensitivus, and
indeed in all Zeuglodontide here represented, are far more
striking than they are in Ornithorhynchus. There can be no
doubt that the afferent impulses reaching the cerebellum from
the anterior end of the body in Zeuglodontidwe were of particular
yalue in supplying these animals with information concerning
disturbances of equilibration. The distribution of the trigeminal
tract within the cerebellum is not fully known, but it is believed
by many investigators that in Mammalia there is a tract for
conveying trigeminal impulses to the cerebellum. The huge
cerebellum in the Zeuglodontide may well be due to the fact
that, by a rapid and hypertrophic development, the trigeminus
provided it with very precise information concerning its position
in space and hence afforded to this primitive creature a ready
solution of the problem of equilibration in a fluid medium.
-Thave said that the distribution of a direct trigeminal tract
within the cerebellum is not fully known; it would be nearer the
truth to state that it is frequently affirmed but is sometimes
denied. Since such denial exists it is valuable to put forward
other supporting evidence. It seems to me that the denial of
the existence of a direct trigeminal connection with the cere-
bellum can only come from an imperfect appreciation of the
developmental history of the cerebellum and a consequent failure
to recognise its extent in the brain-axis—or else from a tendency,
very manifest during the last two decades, to regard the cere-
bellum as an overgrown part of the vestibular apparatus.
As regards the development of the cerebellum it must be
remembered that tho cerebellar ridge first appears in early
embryos much further forward in the hind brain (metencephalon)
than would be anticipated on the ‘“ vestibular” hypothesis.
Even in human developmental history, the nerve which is more
obviously associated with the cerebellum is the trigeminal and
not the vestibular. Ingvar (1918) finds that “Die Basis (cere-
belli) ist frontalwirts gerichtet. Von dem ventralen Rande
dieser Basis laufen die kriftig entwickelten N. N. ‘Trigemini
aus”; or (on p. 343) “ Ventralwirts grenzen die Cerebellar-
wiilste an die Insertionstelle des Trigeminus.”
It is not absolutely certain what actual neuromere of the hind
brain gives rise to the cerebellum, nor is that question pertinent
to the present discussion. The embryological facts significant
here are: (1) that the cerebellum arises in the most anterior
portion of the hind-brain roof, and (2) that the territory of the
BRAIN OF THE ZEUGLODONTID&. 645
neural tube in this vicinity is associated in the vertebrate embryo
with the trigeminus (cf. Wilson and Hill, 1902; Streeter, 1908),
while the acoustico-facialis mass lies entirely posterior to the
trigeminus and is related to a different territory which lies medial
to it. Hence, any encroachment of vestibular fibres from its
territory of the neural tube into the territory which is related
to the trigeminus is a secondary phenomenon whether con-
sidered phylogenetically or ontogenetically, even though such
fibres be direct fibres. In other words, the appearance of vesti-
bular fibres in the cerebellar region is an intrusion just as
certainly as is the appearance of the mesencephalic fibres of the
trigeminal in the mid-brain, or of thalamic fibres in the fore
brain.
The simplest cerebella reproduce more or less faithfully this
condition characteristic of the embryos of all Vertebrates. They
are mere ridges roofing the most anterior portion of the fourth
ventricle and are consequently a link between trigeminal terri-
tories and contain decussations of the trochlear and trigeminal
nerves (Herrick, 1914; Larsell, 1920; Palmgren, 1921).
In the simplest cerebella intrusions of alien fibres are already
found, even as we find alien fibres in the simplest-known fore
brain and tectum opticum. These intrusions occur in the cere-
bellum in such a way that just as we find the olfactory apparatus
is relegated to the periphery of the fore brain and the optic fibres
to the periphery of the mid-brain, so we find that the trigeminal
territory forms the true fringe of the cerebellum. Consequently,
Beccari, Ingvar, and others (Kappers, 1921) have found a direct
distribution of the trigeminus to the cerebellum (¢.9., in
Reptilia).
Topographically, the contribution of the vestibular nerve to the
cerebellum is always surrounded by the trigeminal territory and
the fibres proceeding to the cerebellum therefrom. Ingvar ( 1918)
has shown (in Mammalia) a distribution of the vestibular com-
ponent mainly in the floceulus, lingula, and nodulus, and thus
peripherally to the still more recent intrusions of spino-cerebellar
contributions. On the other hand, spino- and olivo-cerebellar
fibres have not been demonstrated in such peripherally lying
parts such as the lobus floccularis, and it is to be noted (mde
Kappers, 1921) that this region is not dependent upon a cerebro-
pontine contribution; whereas the clinical researches of Winkler
have corroborated the conception that the latest portions of
the cerebellum in the phylogenetic sense are more centrally
situated.
This “laminar” arrangement of the cerebellar constitution, as
it might be termed, although not so clearly defined as the some-
what similar arrangement of successive fibre-intrusions into the
fore brain, is nevertheless present. It is to be expected also
from the fact that those tracts, which later in phylogeny become
incorporated with any region, attain this incorporation by infil-
tration and a spreading apart of the pre-existing mechanism.
42*
646 . PROF. R. A. DART ON THE
The arrangement may be expressed by stating that the vestibular,
olivary, spinal, and cerebro-pontine contributions successively
come to reach the cerebellum by penetrating a territory which
was originally, and so ancestrally, trigeminal.
As has been suggested already, this phenomenon of ‘ pene-
tration” may be illustrated equally well by the infiltrations of the
mid-brain, thalamus, or fore brain, or by the infiltration of the
hippocampal commissure by the corpus callosum in the fore brain
of Mammalia as shown by Elliot Smith. In brief, if we accept
the doctrine of the segmental arrangement of the neural tube
elements, it is to be anticipated that the principle finds illus-
tration in the development of the majority of the inter-segmental
and supra-segmental apparatuses.
The language of neurology is devoid of any term which
describes conveniently this uniformity of behaviour in the laying-
down of subsequent formations in the neural tube. It should
prove of service therefore to descriptive neurology to recognise in
this uniformity the consistent working of a general principle
which underlies the whole architecture of the brain and which we
may term for convenience the Law of Lifiltration.
By an inverse reasoning, if this law is correctly conceived, a
peripheral or fringing arvangement of the trigeminal territory
‘itself, and of fibres known to proceed to the cerebellum from it,
corroborates the identification of the trigeminal apparatus with
the cerebellum and the function of equilibrium, more primitively
even than the vestibular apparatus,
These matters have been neglected by those who assert that
the auricle of the cerebellum, or lobus floccularis (Kappers, 1921),
is the oldest part of the cerebellum. It is not the auricle, but the
anterior medullary velum which would better deserve this appella-
tion. The “vestibular” hypothesis has been attractive because
it has seemed to demonstrate the fact that the cerebellum was
“equilibratory” from the beginning. But this hypothesis tends
to ignore the equilibratory potential of all tactile sense, it fails to
account for the existence of a cerebellum (Cyclostomes) which
has no auricle, and does not account in any way for the striking
relationship which the trigeminal nerve always has to the cere-
bellum in all Vertebrata.
The fifth nerve displays a relationship to that segment of the
neural tube, in which the cerebellum becomes developed, which is
paralleled only by the relationship of the olfactory nerve to that
segment of the tube in which the prosencephalon, and of the
optic nerve to that segment in which the mesencephalon is deve-
loped. The oldest part of the organ is the trigeminal territory
itself just as the olfactory sensorium was the oldest representative
of a fore brain and the visual sensorium of a mid-brain,
I have discovered that Spitzka (1886) put forward a somewhat
similar view concerning the cerebellum nearly forty years ago :—
“Tam inclined to consider it as a homologue of the gelatinous
nucleus of the fifth pair, and as in a primitive relation with that
BRAIN OF THE ZEUGLODONTIDA. 647
nerve; that subsequently the auditory nerve entered into con-
nection with it, and that, increasing in dimension with its in-
creasing neural connections, it attained the high development
found im the human cerebellum through the spinal and cerebral
tracts that are detached into its medullary substance.” 'There-
fore the present statement of this relationship, although more
detailed, cannot be regarded as a novelty.
If we consider the nature of the influences at work in the
production of the early chordate brain, it is evident that the more
anterior segments were concerned in the production of the
olfactory and visual specialisations. ‘These specialisations were
bought at the price’ of successive anterior segments of the
prinitive vertebrate skin and neural tube. When these anterior
segments lost by their specialisation the capacity for the appre-
ciation of pressure, the succeeding segment which retained this
power grew forward invading the territory of the more specialised
segments lying anterior to it. Such an invasion meant an in-
crease or hypertrophy of this (trigeminal) segment and a special-
isation within it of the tactile function. How exuberant has
been its response to this demand is demonstrated by the fact that—
amongst Mammalia, for instance—it subserves the tactile sense
. for the entire region anterior to the second cervical nerve, with the
exception of the vestigial somatic elements of the vagus (Arnold’s
nerve).
The information conveyed by the trigeminal system proved
extremely valuable, because it told the animal go much about
what moved in space around it end also about its own position in
space. ence the central processes from the trigeminal ganglion
assumed a sudden reflex ascendeney over an increasing number
of the succeeding segmental nerves through the radix descendens
and the substantia gelatinosa Rolandi in most Vertebrata. Even
in Man the descending root of the trigeminal represents one of
the most impressive features of the medulla oblongata. Always
a striking feature in Mammalia, this reflex dominance is shown
to its best advantage (in living forms) in Ornithorhynchus ; in
this creature the trigeminal seems to exert a reflex control over
practically the whole of the spinal cord.
Thus there has been a downward encroachment of the trige-
minus upon lower segments of the medulla oblongata in a fashion
comparable with the upward encroachment of the vestibular
nerve into the trigeminal territory or of the mesencephalic root
of the trigeminal upon optic territory. It is a further example
of the Law of Infiltration. Seemingly this encroachment afforded’
the bridge whereby the spino-cerebellar systems were conducted
to the originally trigeminal territory of the cerebellum. For the
spinal fibres naturally lay peripheral to the infiltrating descending
root of the trigeminus, and as the spinal fibres coursed towards
the cerebellum tiiey came to embrace the trigeminus trunk before:
piercing its territory to enter the cerebellum. ‘The differentiation
into anterior and posterior spino-cerebellar tracts emerged from:
648 bit. GC. W. ANDREWS ON “HE
a relationship primarily not to the cerebellum as such, but to the
trigeminus.
. So, in addition to the fringing above mentioned, a graphic
picture of the central role which the trigeminal has played i in the
origin of the cerebellum is afforded by the relation of the anterior
and posterior spino-cerebellar tracts to the trigeminal nerve
(in most Vertebrata) and by the central position of the trigeminus
in the pons (in higher Mammalia),
Considering the nature of the impulses it conveyed, it; is not
surprising that the trigeminal nerve, which has had such diverse
evolutionary possibilities in all vertebrate groups, should have
been very intimately concerned in the emergence and evolution
of the cerebellum in Vertebrata. Nor is it surprising that we
should find illustrated in the Zeuglodontide a group of primitive
mammals taking to a water life at a stage so plastic that the
trigeminal apparatus should become sufficiently specialised and
hypertrophic to determine, through its moulding of the cere-
bellum and the spinal cord, the whole course of evolution (or
devolution) of its central nervous system.
9, Nore on tie SKULLS FROM wuicH THE HNDocRANIAL Casts
DESCRIBED BY Dr. Darr wenn vakun. By C. W. Anbnuws,
D.Se., F.RS. (British Museum of Natural [History.)
(Published by permission of the Trustees.)
Unfortunately, from the nature of the case, the skulls belonging
to the two natural casts above described are unknown, having
been destroyed by weathering, but those from which the remaining
casts were made are preserved and are here discussed. Four such
skulls are known. Of these, two [M. 8150 (a cast) and M. 10228 |
certainly belong to Zeuglodon osiris Dames, from the Qasr-el-
Sagha beds (Upper Mokattam=Bartonian), north of Lake Birket-
el-Qurun in the Fayum. One (M. 9266) is the skull of
Prozeuglodon atrox Andrews, from the lower part of the Birket-
el-Qurun beds in a valley twelve kilometres W.S.W. of the
Gar-el-Gehannem, lying to the west of the lake. his last speci-
men may be regarded as the paratype of the species and is
described and ficured in the British Museum Catalogue of the
Fossil Vertebrata of the Fayum (1906, p. 243). The fourth skull
(M. 10173), forming the middle term of the series, seems from
the nature of the matrix to have been obtained from the Birket-
el-Qurun beds at the western end of the lake. from an horizon
intermediate between those from which the other species were
found. This, however, in the absence of definite information
from the collector is not certain. This skull appears to belong to
a hitherto-undescribed species which has been called Zeuglolon
intermedius Dy Dr. Dart, the characters of which are given
below.
Numerous atihox Zeuglodont remains have been collected from
various horizons in Egypt, and there is some difference of opinion
SKULLS OF TILE ZEUGLODONTIDA. 649
as to their determination and relationship to one another. For
the purpeses of the present paper these may be disregarded and
the series represented by Prozeuglodon atrox, Zeuglodon wuter-
medius, and Zeuglodon osiris alone considered. ‘he characters of
the skulls of these species completely support Dr. Dart’s conclu-
sion, founded on the endocranial casts, that these three species
represent terms in a phyletic series.
The skull of Prozeuglodon has been described in detail in the
catalogue (loc. cit.), so that here it will only be necessary to refer
to a few characters easily comparable with those of the other
forms. ‘The occipital surface (text-fig. 22) above the foramen
magnum is wider than high (if the height* be taken as 100, the
width is 110) and it is only gently concave from side to side. ‘The
foramen magnum and the occipital condyles are both large. The
convexity of the portion of the cranial wall formed by the parietal
Text-figure 22.
Squamous portion
of temporal.
i \gftves
i mG !
SS),
Mastoid —s
process of
petrosal. Foramen
magnum.
Paroccipital process Beppe GEMG
Occipital surface of skull of Prozeuglodon atrox Andrews. M. 9266.
About 4 nat. size.
and squaimosal is greater than in the later forms, while the sagittal
crest is much less developed. The posterior border of the narial
opening is over the space between the first and second premolars
and the snout is less prolonged anteriorly than in the later
forms. Some of the above peculiarities may be dependent to
some extent on the circumstance that the skull deseribed is not
quite adult.
‘The type-skull M. 10173 of Zeuglodon intermedius (text-fig. 23)
is not complete, the zygomatic arch and the terminal portion of
the snout being missing; the teeth also are much broken. In it
the width of the occipital surface (text-fig. 23) is much less
in proportion to its height* (height 100, width 66), and it is
miuch more concave from side to side than in Prozeuglodon.
* In each case the height is measured from the upper border of the foramen
magnum to the point of union of the sagittal and Jambdoidal crests.
650 Dk. 0. W. ANDREWS ON ‘TILK!
But the coneayity is regular and the middle portion of the lamb-
doidal crest is not suddenly reflected backwards as in Zeuglodon
osiis. ‘The occipital condyles and foramen magnum are relatively
smaller than in Prozeuglodon, the occipital crest more developed,
and the convexity of the lateral cranial surface less marked. ‘The
snout was more slender and more prolonged in front of the narial
opening than in Prozeuglodon.
This skull, both in the form of its occipital surface and in other
points, closely resembles that described by Stromer as Zewylodon
osiris (in his paper of 1908) and distinguished by him as Mn.9.
It is very different from that described by the same writer as
Zeuglodon osiris in 1903 and called in his later paper Mn.1.
This latter seems to be the true Zeuglodon osiris, its lower jaw
Text-figure 23.
Squamous
portion
of temporal,
Mastord
process of
Petrosal.
Paroceipital Foramen
process. magnum.
Occipital surface of skull of Zewglodon intermedius, sp.n. M. 10173.
About § nat. size.
agreeing very closely with that of the type-specimen of the
species as described and figured by Dames (1894). Mn.9 is either
the present species (Zeuglodon intermedius) or a form between it
and Prozeuglodon.
The skulls of Zeuglodon osiris from which endocranial casts
have been made are two in number. One (M. 8150) is very
incomplete and reveals only the dorsal contour of the brain.
The endocranial cast of this skull was figured and described by
Prot, Tilliot Smith (1903). The other skull (M. 10228) is a
nearly perfect specimen, and in order that the endocranial cast
might be made it was skilfully cut into sections and the matrix
removed by Mr. L. KE. Parsons (text-fig. 24).
The occipital region of this skull agrees exactly with Stromer’s
SKULLS OF TUE ZEUGLODONTIDA. 651
specimen Mn.1. It is remarkable for its extreme narrowness
compared with its height, the lateral portions of the Jambdoidal
crests being sharply deflected backwards so that the deeply con-
cave occipital surface has the appearance of being pinched in at
the sides. '’his character is also well shown in the cast of the
second skull (M. 8150). If the height be taken as 100 the width
is about 44. The occipital condyles and foramen magnum are
Text-figure 24.
Squamous
portion of
temporal
Mastod
process of
petrosal EI ae
Paroccipital process. Occipita/ condyle.
Occipital surface of skull of Zeuglodon osiris Dames. M. 10228.
About 3 nat. size.
still smaller than in Zeuglodon intermedius and the convex
sagittal crest ismuch higher. The side walls of the skull show
little traces of convexity. The snout seems to have been rela-
tively narrower and perhaps the nostril opened rather further
back than in Zeuglodon intermedius.
The changes occurring in the stages described are :—
(1) Increasing narrowness and concavity of the occipital
surface above the foramen magnum, due mainly to the
folding back of the high lambdoidal crests ;
(2) increased height of the sagittal crest ;
(3) the reduction in the size in the condyles and foramen
magnum ;
(4) decreasing convexity of the lateral cranial walls accom-
panying the depression of the upper portion of the
brain ;
(5) the increase in length and slenderness of the snout and
the probable recession, to some extent, of the narial
opening.
Changes (1) and (2) result in the increase of the surface for
the attachment of the jaw-muscles, and are no doubt correlated
with the increasing size and strength of the mandible.
In Prozeuylodon the two posterior upper premolars have a large
NS
x
652 PROF. R. A. DART ON THE
postero-internal buttress supported by a lar ge distinct root.
In the two species of Zeuglodon this buttress is reduced in size
and probably had no distinct root, though from the state of
preservation of the specimen this cannot be definitely settled.
Some dimensions in millimetres of the skull in the three forms
are given below. These in some instances must be regarded as
approximate only :—
Prozeuglodon.|Z.intermedius.| Z. osiris.
(M. 9266.) (M. 10173.) | (M.10228.) |
i
|
i
i Length from condyles to tip of
|
SHOWA) aiowicues ane, mae eaves 673 | 550 543 app.
Length from lambda to nasal
PMRODENIU preeieamaee we deen ene 403 app. | 453 467
Height of occipital surface from
foramen magnum to lambda... 120 160 156
| ; alk
‘ Least width of occipital surface i
between lambdoidal crests ...... 129 106 69
Width across occipital condyles .| 108 112 92
‘Width across postorbital pro-!
CESSES Nis ade emeneeucreheeeeeaeeeen| 264 _ 283 293
| Width of foramen magnum ...... 56 62 AG
| Length of parietal hortion of
sagittal crest .......ccccce eee eee 125 157 app. 170
|
10. BreuiogRraPny.
Ansrt, O.—* Die Ursache der Asymmetrie des Zahnwalschiidels.” Sitzunghericht.
der Kais. Akad, der Wiss. in Wien. 1902.
“Vine Stammtype der Delphiniden aus dem Miocén der Halbiusel
Taman.” Jahrbuch der k.k. Geol. Reichanstalt, Bd. iv. Hft. 2.
1905.
——- “Cetaceen Studien 1. Mittheilung, Jan. 1909; Cetaceen Studien IT. Mit-
theilung, Mar. 1909 ; Cetaceen Studien Ul. Mittheilung, Jan. 1912.”
Aus der Sitzungsber tichten der Kais. Akad. der Wissensch. in Wien.
—— “Die Vorfahren der Bartenwale.” Denksch. der Math.-Naturwiss.-
Klass. der Kais. Akad. der Wiss. in Wien, Bd. xc. 1913.
AnpreEws, C. W.—“ Extinct Vertebrates from Egypt.” Part I. Geol. Mag. 1901,
p. 400.
-——-—— ‘“xtinct Vertebrates from Egypt.” Part II. Geol. Mag. 1901, p. 436.
—— “Extinct Vertebrates from Egypt.” Part III. Geol. Mag. 1902, p. 291.
“ Purther Notes on the Mammals of the Hocene of Egypt.” Geol. Mag.
1904, pp. 109, 157, 211.
“A Descriptive Catalogue of the Tertiary Vertebrata of the Faytim of
Hgypt.” Brit. Mus. of Nat. Hist. London, 1906.
——— — “The recently discovered Tertiary Vertebrata of Keypt.” The Smith-
sonian Report for 1906, pp. 295-307.
—— — “Zeuglodon wanklyni.” Quart. Journ. Geol. Soc. vol. lxili. pp. 124- 127.
1907,
BRAIN OF THE ZEUGLODONTIDA. 653
Anprews, C. W.—‘ Model of Skull and Mandible of Prozeuglodon atrow.” Geol.
Mag., 1908, pl. ix. p. 209.
“A Description of New Species of Zeuglodont and of Leathery Turtle
from the Eocene of Southern Nigeria.’ Proc. Zool. Soc. London,
1919, p. 309.
\ Antiony, R.— Le Mesoplodon de la Hogue.” Bull. du Mus. Vhist. naturelle,
1909, p. 405.
Breapnet, H. J. L.—“ The Faytim Depression.” Geol. Mag. 1901, p. 640.
Brack, Davipson.—“ Endocranial Casts of the Giraffide.” Journ. of Comp. Neur.
vol. xxv. (1915) p. 341.
—_——_ * Note: on Endocranial Casts of Okapi, etc.” Anatomical Record,
vol. ix.
Boxx, L.—‘ Das Cerebellum der Siiugethiere.’ Jena, 1906.
Brapury, O. C.—* The Mammalian Cerebellum.” Journ. of Anat. & Phys.
vol. xxxvili. 1904, p. 448, & vol. xxxiv. 1906, p. 99.
\ Burmeister, H.—* Bemerkungen tiber Zeuglodon cetoides (Owen’s), Basilosauros
(Harlan’s), Hydrarchos (Koch’s).” Juniheft der allgemein. Lit.
Zeitsch. Halle, 1847.
Carus, C.G., and others.—‘ Untersuchungen tiber das (unter den Namen Hydrarchos
von Dr, A. C. Koch) grosse fossile Skelett.’ Dresden and Leipsic,
1847.
——-——_ “Das Kopfskelett des Zeuglodon hydrarchos.” Nova Acta Acad. Cas.
Leop.-Carol. Nat. Cur. vol. xxii. 1848.
Crark, W. B.—“ Cerebellum of Petromyzon fluviatilis.” Journ. of Anat. & Phys.
vol. x]. 1905-1906, p. 318.
Covz, E. D.—* Brain of Procamelus occidentalis.” Amer. Philos. Society, 1879.
Dames, W.-—Palxontol. Abhandl. N. F. Bd. i. 1894, p. 189.
EprncEr, O.—‘ Nervése Zentral-organe.’ Leipzig, 1896.
Escuricut, D. Fr.—Zool. Anat. Phys. Untersuchungen tiber die nordischen Wal-
thiere. Leipzig, 1849.
Fraas, E.—* Neue Zeuglodonten aus dem unterem Mitteleocan vom Mokattam bei
Cairo.” Paleont. Abhandl., n.s., vol. vi.
____. Jahreshefte Verein. vaterland, Naturkunde Wiirttemberg. Stuttgart,
1905, p. 347.
Grecory, W. K.—* The Orders of Mammals.” Bull. Amer. Mus. Nat. Hist.
vol. xxvii. 1910. E
Hurricx, C. Jupson.—* Cerebellum of Urodele Amphibia.” Journ. of Comp.
Neur. 1914.
Inevar, Sven.— Zur Phylo- und Ontogencse des Kleinhirns, etc.” Folia Neuro-
biologica, Bd. xi. 1918.
Karrers, C. U. Artiins.—‘ Die vergleichende Anatomie des Nervensystems, ete.’
1921.
“Cerebral Localisation and the Significance of Sulci.” Report. of
Anatomy and Embryology Section: Internat. Congress of Medicine.
London, 1913.
Kixentuan, W.—“ Vergleich anat. und entwick. Untersuch an Walthiere.” Jenaische
Denkschriften, vol. iii. Jena, 1893.
Larse1t, Orar.— The Cerebellum of Amblystoma.” Journ. of Comp. Neur.
1920.
Marsun, O. (.—“ Introduction and Succession of Vertebrate Life in America.”
Amer. Ass. for Adv. of Science. Nashville, Tenn., 1877.
Moopts, Rox L.—‘“A Fish Brain from the Coal Measures.” Journ. Neur. 1916.
Osborn, I. F.— Correlation between Tertiary Mammalian Horizons of Europe and
America.” Annals N.Y. Acad. Sci. vol. xiii, No.1, July 1900.
Patmer, R. W.—“ Brain and Brain-case of Anoplotherium.” Proc. Zool. Soc. of
London, pt. iv. p. 878 ef seq. 1913.
Patmeren, A.—‘Studies on the Mid-brain and Cerebellum.” Acta Zoologica,
Hatt 1-2, Arg.2. 1921.
654 THE BRAIN OF THE ZEUGLODONTIDA.
Purrer, O—* Die Augen der Wassersiiugethiere.” Zool. Jahrbuch, Abthieil. fir
Anat., etc. Bd. xxxvii. p. 99 ef seg. Jena, 1902.
Scuutre, H. von W.—“ Skull of Kogia breviceps Blaiuv.” Bull. Amer. Mus, Nat.
Hist. vol. vii. 1917, p. 361.
SHELLsUEAR, J. L.—* Blood-supply of the Dentate Nucleus of the Cerebelliun.”’
‘The Lancet,’ 1. p. 1046. 1922.
Surmazono, J.—‘ Das Kleinhirn der Vogel.” Arch. fiir mikr. Anat. 19.
Kuiior Smita, G.—‘The Brain in the Hdentata.” Trans. Linn. Soc. London,
vol. vii. pt. vil. pp. 360 et seg. 1899.
‘Catalogue of the Roy. Coll. of Surgeons.’ 2nd Kdit., vol. xi. Physio-
logical Series, 1902.
——— “The Primary Subdivisions of the Mammalian Cerebellum.” Journ.
Anat. & Phys. vol. xxxvi., n.s., vol. xvi. p. 381. 1902.
——— The Brain of the Archaoceti.’ Proc. Roy. Soc. vol. Ixxi. 1903, p. 322.
—— “The Morphology of the Human Cerebellum.” Roview of Neurology
and Psychiatry, 1908.
—— “Further Observations on: the Natural Mode of Subdivision of the
Mammalian Cerebellum.” Anat. Anzeig. Bd, xxiii, (1908), p. 368.
Spitzxa, K. C.— Architecture of the Brain.” Journ. of Nervous and Mental
Diseases, 1886.
Srreetrer, G. L.—“ The Nuclei of Origin of the Cranial Nerves in the 10-mm.
Human Embryo.” Anat. Ree. vol. ii. No. 38, p.111. June 1908.
SrromerR, E:—Beitriige zur Palont. Oesterreich. Ung. Bd. xv. 1903, p. 65.
Beitrige zur Palwont. Oesterreich. Ung. Bd. xxi. 1908, p. 106.
—-— “Ueber die Bedeutung der fossilen Wirbelthiere Afrikas fiir die ‘hier-
geographie.” Verhandl. d. Deutsch. Zool. Gesellsch. p. 204, 1906.
(Marburg meeting).
Symineron, J.—‘ Relations of Inner Surface of Cranium to the Surface of thie
Brain.” Edin, Medical Journal.
Truz, EF, W.—“‘'Che lossil Cotacean, Dorudon serratus Gibbes.” Bull. Mus. Comp.
Zool. Harvard College, vol. lii. 1908, p. 65.
Weber, Max.—‘ Die Siugethiere,’ 1904, pp. 680-584.
Winer, H.—‘ Udsigt over Hvalernes indbyrdes Slaegtskab.” Vidensk. Medd. fra
Dansk. naturh. Foren., Bd..Ixx. 1919; (‘Lrans. by G. S. Miller, Jr.,
Smithsonian Miscellaneous Collections, vol. Ixxii. No. 8. Publi-
cation 2650. 1921.)
THE SECRETARY ON ADDITIONS TO 'THE MENAGERIE. 655
EXHIBITIONS AND NOTICES.
May 8th, 1923.
Dr. A. Surrm Woopwarp, F.R.S., Vice-President,
in the Chair.
Lhe Secrerary exhibited, and made remarks upon, a series of
photographs of Big Game from Choma, North Rhodesia.
Miss L. KE. Cutesman, F.E.S., F.Z.S., exhibited, and made
remarks upon, (1) living specimens of Peripatus from Trinidad
and (2) a section of a nest of the Stingless Bee from Australia.
Mr. F, Marrin Duncan, F.R.M.S., F.Z.8., gave an account
of some of the results of his recent work on the microscopic
structure of Mammalian hairs, with especial reference to the
hairs of the Primates, and illustrated his remarks with a series
of photomicrographs taken to demonstrate the character of the
cuticular scales.
May 29th, 1923.
Dr. A. Smiru Woopwarp, 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 April, 1923 :—
The registered additions to the Society’s Menagerie during
the month of April were 367 in number. Of these 90 were
acquired by presentation, 50 were deposited, 215 were purchased,
and 12 were born in the Menagcrie.
The following may be specially mentioned :—
2 Adult male Lions (/elis leo), from Nairobi, presented by
Major-General Sir Edward Northey, K.C.M.G., O.B., F.Z.S.,
on April 9th.
2 Common Wombats (Phascolomys mitchell), from Australia,
and 2 Tasmanian Devils (Sarcophilus harrisi), from Tasmania,
purchased on April 20th.
2 Superb Glossy Starlings (Spreo superbus) and 1 Abyssinian
Barbet (Zrachyphonus margaritatus), from North-East A frien,
new to the Collection, presented by Mr, Alfred Ezra, O.B.¥.,
¥.Z.8., on April 15th.
656 MR. C. TATE REGAN ON A SPOYTED BASKING SHARK.
A collection of animals from Gambia, presented by H.H. Capt.
C. H. Armitage, C.M.G., D.S.0., I.Z.S., consisting of a Wart-
Tfog, 6 Guinea Baboons, Marabou Storks, &e,
A small collection of birds from Colombia, presented by
Mr. W. K. Pomeroy, F.Z.8., containing 2 Prince Albert’s
Curassows and several Tanagers.
A Tail-lined Tree-Snake, Dendrelaphis caudolineatuws, and an
Tridescent Snake, Xenopeltis unicolor, from Singapore, both new
to the Collection, presented by the Singapore Natural History
Society.
Dr. G. M. Vevers, F.Z.S., exhibited, under microscopes, a
series of preparations of the parasitic Protozoon Balantidiwm
coli, and described a case of Balantidiosis in a Brazilian Tapir
now in the Society’s Gardens.
Prof. J. P. Hi, F.R.S., Vice-President, exhibited a series of
photographs of the Australian Lung-Fish (Ceratodus) and of the
rivers which it inhabits, and drew attention to the urgent need
for effective protection of this rare Vertebrate.
Dr. N.S, Lucas, F.Z.S., exhibited, and made remarks upon,
a mucous cyst from the frontal sinus of a Cercopitheque.
Mr. C. Tare Reaan, M.A., F.R.S., exhibited lantern-slides of
photographs of the Spotted Basking Shark (Rhinodon typicus),
and made the following remarks :—
The first of these photographs represents a specimen 38 feet
long captured by Captain Charles Thompson off Miami, Florida,
in 1912. It was set up by a taxidermist, mounted on a truck,
and taken round for exhibition. At the time certain American
newspapers published stories about it that were not strictly
accurate, and to the effect that it was a monster unknown to
science that had been blown up from the depths by a volcano,
etc., etc. The second photograph represents a Lhinodon about
30 feet long, which was run into by a large steamship off the
coast of Brazil and was carried along for several hours, with the
anterior } of the body on one side of the bows and the rest on
the other; when the vessel stopped it floated free and sank
(cf. Gudger, Natural History, xxiii. p. 62, New York, 1923).
Mr. Regan also exhibited some deep-sea fishes taken by the
‘Dana’ expedition, under the leadership of Dr. Johannes
Schmidt. This expedition cleared up the life-history of the
Common Eel (Anguilla vulgaris), but it also nccomplished much
other work of great importance. ‘The fishes exhibited belong to
the very rare and little-known genera Gigantura and Stylophorus,
which agree in having telescopic eyes placed close together and
directed forwards. One Gigantura had swallowed a Chauliodus
considerably larger than itself.
THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 657
June 12th, 1923.
Sir 8. F. Warmer, K.B.E., F.R.S., Vice-President,
in the Chair.
The Sucrerary read the following Report on the Additions to
the Society's Menagerie during the month of May, 1923 :—
The registered additions to the Society’s Menagerie during the
month of May were 214 in number. Of these 149 were acquired
by presentation, 32 were deposited, 22 were purchased, 2 were
received in exchange, and 9 were born in the Menagerie.
The following may be specially mentioned :—
2 Hlephants (Llephas maximus), from Burma, presented by
Dr. Saw Po Min on May 16th.
A collection purchased from the Soudan Government on
May 14th, consisting of 1 female Giraffe (Giraffa camelopurdalis),
1 Wart- Tog (Phacocherus africanus), 4 Arabian Bustards (Zupo-
dotis arabs), and 2 Northern Secretary-Birds (Serpentarius
serpentarius gambiensis).
2 Abyssinian River-Hogs (Potamocherus hassama), new to the
Collection, purchased on May 5th.
Mr. D. Seru-Smiru, F.Z.S., and Dr. P. H. Manson Banr,
DS.0., F.Z.S., exhibited photographs and sketches, and made
remarks upon, the display of Hunstein’s Magnificent Bird-of-
Paradise (Diphyllodes magnifica hunsteint).
Miss Atys— Cunnincuam exhibited, and made remarks upon,
a Cinematograph record of the Gorilla ‘‘ John.”
Mr. E. G. Bouuencer, F.Z.8., exhibited and described models
for the rockwork for the Society’s new Aquarium, made by
Miss Joan B. Procter, F.Z.S.
Mrs. NeEAuE exhibited a living Tree-Hyrax (Dendrohyrax
dorsalis).
Dr. H. A. Bayuis, F.Z.8., exhibited, and made remarks upon,
a Nematode Worm, Zoxascaris leonina, from the Domestic Cat..
Pituitary Gland and Axolotl Metamorphosis.
Mr. HE. A. Spaut, F.Z.8., exhibited specimens showing the
metamorphosis of axolotls induced by injection of pituitary
gland anterior lobe extract, and made the following remarks :—'
Recent studies on the relationship between amphibian meta-
morphosis and endocrine organs have demonstrated the significant
role of both the thyroid and pituitary glands. . The acceleration
of the metamorphosis of tadpoles to frogs when fed with thyroid
658 MR, E. A. SPAUL ON 'THE
tissue and the failure of thyroidless tadpoles to change until
given either thyroid diet or iodine, followed by the conversion of
axolotls to salamanders by thyroid diet, showed conclusively the
importance of the thyroid and thyroid iodine. Although pitui-
tary diet does not induce metamorphosis in either tadpoles or
axolotls, pituitaryless tadpoles fail to transform, whilst axolotls
injected with extract from the anterior lobe of the pituitary
land assume adult characters as rapidly as individuals treated
with thyroid, which is evidence in favour of the significance of
the anterior lobe of the pituitary in metamorphic changes.
The results of experiments briefly outlined here show that the
anterior lobe contains an active principle functioning in a definite
manner in metamorphosis. Previously only young axolotls had
been used, but larger specimens, in some cases beyond that size
at which they change in normal environment, were taken, and by
tri-weekly injections caused to transform to adults. ‘The rate of
metamorphosis depended on the size of the animal and the tempera-
ture. The first signs of change appeared rauch later in the older
specimens, but providing the dose was sufficient the process was
more rapid. Within two or three weeks of the first injection the
change begins and is usually complete by the 40th day in medium
size or the 48th day in the oldest types under suitable conditions
of tomperature, ‘The limits of the latter are 22°-24° C., when the
animal remains quite normal throughout the experiment. Low
temperature retarded the change, and if too high the animal
became sluggish, refusing food, with no noticeable increase in
the rate of change.
The approximate limits of the minimal dose were found, but
it was not possible to give any relation between the minimal
dose within these limits and the weight, as the complexity of
the factors, both internal and external, made the production of
identical conditions each time, and hence uniformity, impossible.
The limits were ‘5 grs. of fresh gland per ‘5 c.c, injection for
small individuals to 1:5 grs. for full-grown animals.
Weights and measurements were taken on days alternating
with those upon which injections were made throughout the
period of the experiment, but no increase in growth was noted,
only a decrease. ‘This does not deny the growth-promoting
properties of the anterior lobe, for the animals used were adult
and sexually mature, so that reduction previous to metamorphosis
was to be expected.
Having ascertained these details, the change resulting from
the injection of the anterior lobe extract was applied as a
biological test in the examination of the purity and strength
of commercially prepared extracts, Quantities up to 1/10 of a
clinical dose could be tested in this manner. Of twelve so tested
only one successfully brought about the change, although as
many as 20 injections were made. The dose was the same in
each case, the quantity stated by the manufacturer being taken
as a basis and equivalent to the minimal doseas found in previous
experiments,
METAMORPHOSIS OF AXOLOLLS. 659
The cause of failure was probably due either to loss of potency
during process of manufacture or presence of impurities having
an inhibitory effect. As regards the first, the strength of the
dose was doubled with no result, so that it was nota question
of quantity. The only impurity, apart from those accumulating
in process of preparation (a very small amount, if present), and
investigation of which was beyond the scope of this work, was
the posterior lobe of the gland. ‘he difficulty in separation of
the two lobes in dissection and diffusion of active principles
during dissection would account for its presence.
It has recently been shown that there is a pigmentation factor
located in the posterior lobe, and by its action on the frogs’
melanophores a quantitative estimation can be made by deter-
mining that concentration which just gives a response. ‘this
method .was successfully applied here and the quantity of
posterior lobe in each extract calculated.
Further, many animals treated refused food, the water in the
container became cloudy, smelling of urine, and some eventually
died, after becoming greatly distended with fluid, which was
found to contain 271 per cent. of urea. As the symptoms were
the same in each case, and-occyrred only in specimens injected
with extracts and not controls, the cause must have been the
same, and introduced by the extract. ‘Cherefore the factor was
apparently contained in the posterior lobe and responsible for
inhibition of activity of anterior lobe secretion. It was possible
to compare roughly the amounts present by judging effects on
specimens.
On comparing these observations with the results obtained
from the pigmentation work, the respective amounts in each
extract did not correspond, which seems to show that there were
two factors at least which were not identical—the one control-
ling pigmentation response of the melanophores and the other
associated in some way with the flow of utine and inhibiting
action of anterior lobe. It is doubtful whether the pigmentation
factor itself had any such action. &
In conclusion, it can be seen that the anterior lobe extracts as
at present manufactured are not pure, containing posterior lobe,
which may completely mask metamorphic action of anterior
lobe, although a small quantity has no effect. The varying
amount in which it may be present shows consistent composition
and is not maintained in manufacture. Again, the stated amounts
of gland present in commercially prepared extracts is not to be
relied upon, as tested by this method.
Proc. Zoo. Soc.—1923, No. XLITI. 43
Hitt barnes
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'
No. 241.
ABSTRACT OF THE PROCEEDINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON.*
May 8th, 1923.
Dr. A. Smiru Woopwarpb, F.R.S., Vice-President,
in the Chair.
The Sxcrerany exhibited, and made remarks upon, a series
of photographs of Big Game eosin Choma, North Rhodesia.
.Miss L. KE. Cuerrsman, F.E.S., F.Z.8., exhibited, and made
upon, (1) living specimens of Per ipatus from Trinidad, and
(2) a section of a nest of the Stingless Bee from Australia.
Mr. F. Martin Duncan, F.Z.8., gave a réswmé of a paper
by Mr. H. Burretu, C.M.Z. a. “Note ona Hibernating Female
Specimen of the Marsupial Acrobates pygmeus,’
Mr. F. Martin Duncan, F.Z.S., gave an account of some
of the results of his recent work on the microscopic structure of
Mammalian hairs, with especial reference to the hairs of the
Primates, and illustrated his remarks with a series of photo-
micrographs taken to demonstrate the character of the cuticular
scales,
* This Abstract is published by the Society at its offices, Zoulogical 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 ‘Procecdings,’ free of extra charge,
to all Fellows who subseribe to the Publications; but it may be obtained on the
day of publication at the price of Sixpence, or, if desired, sent post-free tor
the sum of Six Shillings per annum, payable in advance.
38
The next Meeting of the Society for Scientific Business will
be held on ‘Tuesday, May 29th, at 5.30 p.m., when the following
Communications will be made :—
C. Tare Reaan, M.A., F.R.S., F.Z.S.
The Skeleton of Lepidosteus, with Remavks on the Origin
and Evolution of the Lower Neopterygian Iishes.
Dr. Cuas. F. Sonnraa, F.Z.S.
The Comparative Anatomy of the Tongues of the Mam-
malia.—IX, Edentata, Dermoptera, and Insectivora.
S. Mavuix, I’.Z.8S.
New Cryptosome Beetles.
The following Papers have been received :—
R. Broom, F.R.S., O.M.Z.S.
On the Structure of the Skull in the Carnivorous Dino-
cephalian Reptiles.
N. A. Mackintosh, B.Sc.
The Chondrocranium of the Teleostean Fish Sebastes
Marinus,
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.
39
Communications intended for the Scientific Meetings should
be addressed to
P. CHALMERS MITCHELL,
Secretary,
ZOQULOGICAL SOCIETY OF LONDON,
Reaeenv’s Park, Lonpon, N.W. 8.
May 15th, 1922.
No. 242.
ABSTRACT OF THE PROCEEDINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON.*
May 29th, 1923.
Dr. A. Smirn Woopwarp, F.R.S., Vice-President,
in the Chair.
The Srcreranry read a Report on the Additions to the Society’s
Menagerie during the month of April 1923.
Dr. G. M. Vevers, F.Z.S., exhibited, under microscopes, 2
series of preparations of the parasitic Protozoon Balantidium colt,
and described a case of Balantidiosis in a Brazilian Tapir now in
the Society’s Gardens.
Prof. J. P. Hit, F.R.S., Vice-President, exhibited a series of
photographs of the Australian Lung-fish (Ceratodus) and of the
rivers it inhabits, and drew attention to the urgent need for
effective protection of this rare Vertebrate.
Dr. N. 8. Lucas, F.Z.S., exhibited, and made remarks upon,
a mucous cyst from the frontal sinus of a Cercopitheque.
Mr. C. Tare Reeay, M.A., F.B.S., F.Z.S., exhibited, and made
remarks upon, photographs of the Spotted Basking Shark
(Rhinodon typicus). Mr. Regan also exhibited some deep-sea
fishes taken by the ‘Dana’ Expedition, under,the leadership of
Dr. Johannes Schmidt. The fishes exhibited belonged to the
very rare and little-known genera Gigantenu and Stylophorus,
which agree in having telescopic eyes placed close together and
directed forwards.
* This Abstract is published by the Society at its offices, Zoological Gardens,
Kegent’s Park, N.W., on the Tuesday following the date of Meeting to whicla
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 Sex Shillings per annum, payable in adyance,
42
Mr. ©. Tare Reaan gave a réswmé of his paper on “ The
Skeleton of Zepidosteus, with Remarks on the Origin and Evo-
lution of the lower Neopterygian Fishes.”
Communications by Dr. Cuas. F. Sonntag, F.Z.S., on “ The
Comparative Anatomy of the Tongues of the Mammalia.—
IX. Edentata, Demoptera, and Insectivora,” and by Mr. 8.
Mauus, I.Z.8., on “ New Cryptosome Beetles,” were taken as
read,
The next Meeting of the Society for Scientifie Business will be
held on Tuesday, June 12th, 1923, at 5.30 p.m, when the
following Communications will be made :—
R. Broom, F.R.S., C.M.Z.S.
On the Structure of the Skull in the Carnivorous Dino-
cephalian Reptiles.
N. A. Mackintosn, B.Se.
The Chondrocranium of the Teleostean Fish Sebastes
Marius.
R. I. Pocock, F.RS., F.Z.S.
The Extermal Characters of the Pigmy ITippopotamus
(Cheropis liberiensis) and of the Suide and Camelidee.
Major E. EH. Ausren, D.S O., F.Z.S.
A Revision of the Family Pantophthalmidz (Diptera), with
Descriptions of new Species and a new Genus.
Raymonp Darr, M.Sc., M.B., and C. W. Anprews, D.Sc, F.1.S.,
F.Z.8.
The Brain of the Zeuglodontidee (Cetacea), with a Note on
the Skulls from which the Endocranial Casts were taken.
OxprieLp Tuomas, F.R.S., F.Z.8., and M. A. C. Hinton, F.Z.5.
Hee eee eee eee
On Mammals collected by Capt. Shortridge during the
Percy Sladen and Kaffrarian Expedition to Orange River.
43
The following Papers have been received :—
H. C. Apranam, F.Z.S,
A new Spider of the Genus Liphistius from the Malay
Peninsula, and some Observations on its Habits.
A. Subsa Ran, B.A., M.Sc., F.R.M.S., and P. H. Jonnson,
a NE Oe ee A
B.Sce., F.Z.8.
Observations on the Development of the Sympathetic
Nervous System and Suprarenal Bodies in the Sparrow.
Matcoum A. Smirn, M.R.C.S., L.R.C.P., F.Z.S.
SE ee ER ESOS
A. Review of the Lizards of the Genus Tropidophorus on the
Asiatic Mainland.
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 MITCILELL,
Secretary.
ZOOLOGICAL Society or Lonpon,
Recent’s Park, Lonpon, N.W. 8.
June 5th, 1923.
Nat ee
oe me ye ‘i Ayal we coast fh ;
ot ea
No. 2438.
ABSTRACT OF THE PROCEEDINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON.*
June 12th, 1923.
Sir S. F. Harmen, K.B.E., F.R.S., Vice-President,
in the Chair.
The Sucrerary read a Report on the Additions to the Society’s
Menagerie during the month of May 1923.
Mr. D. Seru-Ssrrn, F.Z.S., and Dr. P. H. Manson Baur,
D.S.0., F.Z.S., exhibited photographs and sketches, and made
vemacks upon, the display of Hunstein’s Magnificent Bird-of-
Paradise (Diphyllodes magnifica hunsteint).
Miss Atyse CunnrenciAm exhibited, and made remarks upon,
a Cinematograph record of the Gorilla “ John.”
Mr. E. G. Bounenesr, F.Z.S., exhibited and described models
for the rockwork for the Society’s new Aquarium, made by
Miss Joan B. Procrer, F.Z.8.
Mrs. Neate exhibited a living Tree-Hyrax (Dendrohyrax
dorsalis).
Dr. H. A. Bayuis, F.Z.S., exhibited, and made remarks upon,
a Nematode Worm, Z'oxascaris leonina, from the Domestic Cat.
Mr. BE. A. Sraut, F Z.S., exhibited, and made remarks upon,
Axolotls in which metamorphosis had been induced by pituitary-
gland injection.
Mr. N. A. Macxinrosy, B.Sc., communicated his paper on
“The Ghondrocranium of the Teleostean Fish Sebastes marinus.”
* 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 Mcllows 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 Six Shillings per annum, payable in advance.
46
Mr. R. I. Pocock, F.R.S., F.Z.8., gave a résumé of his paper
on “The External Characters of the Pigmy Hippopotamus
(Cheropsis liberiensis) and of the Suid and Camelide.”
Major E. E. Ausvey, D.S.O., F.Z.S., gave a résumé of his paper
“A Revision of the Family Pantophthalmidee (Diptera), with
Descriptions of new Species and a new Genus.”
Prof. G. Exuior Suiru, F.R.S., F.Z.8., gave a résumé of the
paper by Mr. Raymonp Dart, M.Sc., M.B., and Dr. C. W.
Anprews, F.R.S., F.Z.S., on “The Brain of the Zeuglodontide
(Cetacea), with a Note on the Skulls from which the Endocranial
Casts were taken.”
Communications by Dr. R. Broom, F.R.S., C.M.Z.8., “On the
Structure of the Skull in the Carnivorous Dinocephalian Rep-
tiles,’ and by Mr. Oxiprretp Tuomas, F.R.S., F.Z.8., and Mr.
M. A. ©. Hinton, F.Z.S., “On Mammals collected by Capt.
Shortridge during the Percy Sladen and Kaffrarian Expedition
to Orange River,” were taken as read.
The next Meeting of the Society for Scientific Business will be
held on Tuesday, October 23rd, 1923, 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 :—
lI. C. Apranam, F.Z.S.
A new Spider of the Genus Liphistius from the Malay
Peninsula, and some Observations on its Habits.
A. Suppa Rav, B.A., M.Se., F.R.M.S., and P. H. Jounson,
B.Sce., F.Z.S.
Observations on the Development of the Sympathetic
Nervous System and Suprarenal Bodies in the Sparrow.
47
Mancoum A. Sairu, M.R.C.S., L.R.C.P., F.Z.S.
SSE eS eS SLO
A Review of the Lizards of the Genus Tropidophorus on the
Asiatic Mainland.
J.G. H. Frew, M.Sc., F.Z.S.
On the Larval Anatomy of the Gout-fly (Chlorops taniopus
Meig) and two related Acalyptrate Muscids with Notes on
their Winter Host-plants.
Arruur Lovreriper, F.E.S., 0.M.Z.S.
a enw oeerrenaeeeeey
(1) Notes on Mammals collected in Tanganyika Territory,
1920-1923.
(2) A List of the Lizards of British East Afvien (Ugan da
Kenya Colony, ‘Tanganyika Territory, and Zanzibar) with
Kveys 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 Mcetings should
be addressed to
P. CHALMERS MITCHELL,
Secretary.
ZOOLOGICAL Society oF Lonpon,
ReceEnt’s Park, Lonpon, N.W. 8.
June 19th, 1923.
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26.
~ 27,
28.
29,
30.
31.
33.
PAPERS.
race
On Mammats collected by Capt. Shortridge during the Percy Sladen and Kaffrarian
Museum Expedition to the Orange River. By Ouprreup Tuomas, F.R.S., F.Z.9., and
Mice nin Ari Os ELNTON SHEA Sees seis Se ta eer ater as ny AR nn ES Rea gg
The Chondrocranium of the Teleostean Fish Schastes-marinus. By N. A. Macxinrosu,
IB Sets (Maxt-forires iO! ioe scysy. cicratelcosre eave a arsternctotacel nailer a 501
The Comparative Anatomy of the Tongues of the Mammalia.—IX. Ldentata,
Dermoptera, and Insectivora. By Cnas. F. Sonnrac, M.D., F.Z.8. (Text
Agumes} OO= Dis) ee senate weretstarate totale sis cyivei hy courses are alee eines ae eee Oy estar ayy ee TS
The External Characters of the Pigmy Ilippopotamus (Cheropsis liberiensis) and of
the Suid and Oamelide. By R. I. Pococx, F.R.S., F.Z.8, (Text-figures 30-46.) . 581
A Revision of the Family Panthophthalmids [Diptera], with Descriptions of New
Species and a new Genus. By Major EH. BE. Ausren, 1).8.0., F.Z.S8. (Text-
figures) IN) ite yeheateya's «Gusta tene caste Sears ettscatateree ca eee ee aaa ae / geo sc0ooa0e 551
New Cryptosome Beetles. By 8. Mauuix, F.Z.S. ere heures 1-8.) ......0+.... 599
. On the Display of the Magnificent Bird-of-Paradise Diphatiodes meanitoe aneteini,
By D. SEES SwitH, F.Z.8. (‘Lext-figures 1-4.) .............---.03..... 00 pedo 609
The Brain of the Zeuglodontids (Cetacea). By Rarmonp ie Daxt, M.Sc., M.B
Ch.M.; with a Note on the Skulls from which the Endocranial Casts were taken.
By C. W. Anprews, D.Sc., F.R.S., F.Z.8. (Text-figures 1-24.).........--- 0000s - 615
1928, Parr III. (pp. 483-659).
NOTICE.
The ‘ Proceedings’ for the year are issued in four parts, paged consecutively,
so that the complete reference is now P. Z. 8. 1928, p.... The Distribution
_ is usually as follows :—
Part I. issued in March,
he el () eae nen? June.
» III. ,, ~ September. -
ly. December,
‘Proceedings, 1923, Part II. (pp. 181-481), was published on July 6th,
1923.
The Abstracts of the ‘ Proceedings,’ Nos. 241-243, are
contained in this Part.
The dates of Publication of ‘ Proceedings’ 1830-1858 will be found in the
‘ Proceedings’ for 1893, page 436,
Tho dates of Publication of ‘ Transactions’ 1833-1869 will be found in the
‘ Proceedings’ for 1913, page 814.
SO
&
rR
Pee
- PROCEEDINGS
| : OF THK
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS
OF TILE
| 1
|| ZOOLOGICAL SOCIETY | |
|
OF LONDON. |
1923.
A oyniat Insz;
| : ee al
x 2 a TERE A 2
| : PART [Ve ee AP Ree che
containine Paces 661 vo 1097; wira 1
Tex aiics
Ppa HI GUMES:
LONDON:
MESSRS. LONGMANS, GREEN, AND CO.,
PATERNOSTER ROW.
8 Prates, any 1 Map. Stional Muse
|
DECEMBER 1923.
PRINTED FOR THI SOCIETY,
SOLD AT ITS HOUSE IN REGENTS PARK.
|
ye [Price Twelve Shillings.) ra
05% ___ 3 hes
LIST OF CONTENTS
1923, Parr IV. (pp. 661-1097).
EXHIBITIONS AND NOTICES.
Page
Tue Secretary. Report on Additions to the Society’s Menagerie during the months of
June, July, August, and September, 1923 ........ 006. ceeee ess eee cece sere veee 1093
Dr. C. W. Anpruws, F.R.S., F.Z.S. Exhibition of phalangeal bone of a clawed Perisso-
CIGAR Sa bocuoedonac SGba0q0000n D0 140090000000900000 50:0900009000000000.090C 1095
Mr. D. Suru-Swirn, F.Z.8. © (1) Exhibition of photographs of Lion-cubs; and (2) of rare
varieties of Black Mangabey Monkey ........0...cceesesecceevesseeeecceceses L095
Dr. Carn ABsonon. Exhibition of drawings and. photographs of Cave Animals from.
g ] gray
the Balkans..... STAVE euawitl stein cl mdecdevarevehettet ch svetsi es emsvariefesaisteiial a haiar oh smenelte Grey mteueten ek cicoreaiemers ee 1095
Mr. Ei. A. Svaun, B.Se., V.Z.8. Remarks on acceleration of metamorphoses of Frog-
tadpoles ee eer ce ee tere tees eens ve ee eo ea ve se Po oe se Core reese sn reste hes eee 1095
ne Skerurany, lUxhibition of photograph ofa Ratel .......5.4...-.--+--5+-nee-- ++ 1096
My. D. Sera-Suern, F.Z.8, Wxhibition of a young Pigmy Hippopotamus, (Lext-figure 1.) 1096
Mr. BF. A. Mirenect-Hevers, F.L.S8., F.Z.8. Exhibition of photographs, and remarks on
. his recent expedition to the jungle-region of Panama ..... WOOHOO RIO COS da G0 GS 1096
Tne Secrerary. Report on Additions to the Society’s Menagerie during the month of
Mclover MOD ee Acie certo ane crsusievonevovaeeieliel set suarenee Toencant acne NEO RAO OOO 1097
Tne Secretary, Exhibition of Autograph letter of Sir Stamford Rafiles, and of a collec-
tion of Autographs recently presented to the Society ......-. 0. ee ee ee eee ee wees LOOT
Mr, R. Tf. Gunrner, M.A., F.Z.S. “Exhibition of (1) Vertebra of Mesozvie Crocodile ;
ancl (24) OF Qa TMEIONS Ole WASH GLGMGUS conan cbond0on00 0b e6 VGN Da sDOUDODOOs On 1097
PAPERS.
34, On the Structuro of the Skull in the Carnivorous Dinocophalian Hep iles: By Rh.
Broon, M.D., F.R.S., O.M.Z.8.° (Text-figures 1-17.) -. 0.6... sc ec ee ce te ee eee soo | HDI
35, Notes on Bast Afvican Mammals, collected 1920-1923. By Arrnur Loyrerinan, I.E.S.,
CObMEYAS (uy) Songun omedodioedddudodadedeusacodsesndcodor ce cadadoogonge Ute)
36. Observations on the Development. of the Sympathetic Nervous System and Supra-
renal Bodies in the Sparrow. By A. Suppa Rav, B.A., M.Sc., F'.R.M.S., and
P, H, Joungon, B.A., B.Se., F.Z.8., ete, (Voxt-figures 1-20.) .. 0... cece ee ee ee ee 741
37. A New Spider of the Genus Liphistius from the Malay Peninsula, and some Observa-
tions on its Habits. By H.C, Asranan, F.Z.S., F.L.8. (Plate I.; Text-figure 1.).. 769
38. A Review of the Lizards of the Genus Zxopidophorus on the Asiatic Mainland, By
MAN CODA AN SMITA MAR CANu alt @ nes ale Za sumanicnetetetleleherchernelelcioneieneteietelcieucieietsiete 775
Contents continued on page 3 of IVrapper.
ZOOLOGICAL SOCIETY OF LONDON,
INSTRUCTIONS TO AUTHORS.
Owine to the cost of printing, the Publication Committee
beg to draw the attention of Authors submitting papers for
publication by the Society to the following Regulations. If the
conditions are not observed, papers may have to be returned :—
1. MSS. submitted for publication must be written in a clear
and legible handwriting, or, preferably, be type-written, and
must be in final form for printing. The MSS. should be written
on one side of each sheet only, and the pages clearly numbered.
All tables of contents, references to plates, or legends to
illustrations in the text, elc., must be in their proper places,
with page and plate numbers left blank; and the position for
the text-figures must be indicated. References to literature must
be exact, with the page number as well as the volume number
and date.
2. ProoFs AND THEIR Correcrion.——I'wo copies of the slip
proof, and a page proof for final revision, will be sent to the
Author, but it is important that all serious corrections be
made on the slip proof. Expenses for proof corrections are very
heavy, but, if the MSS. comply with Regulation 1, should not
exceed 25 per cent. of the printers’ charges. If the cost of
corrections exceeds 25 per cent., a proportion of the excess will
be charged to the Author.
3. Revised proofs must, if possible, be returned within one
week from the date of their receipt by the Author.
4. The Author receives 30 copies of reprints free, and, with
the sanction of the Publication Committee, may obtain leave to
buy extra copies, at the cost price charged by the printers and
engravers to the Society. ‘The number of separate copies
required must be stated when returning the proof.
5. InLustrations.— Drawings intended to be reproduced as
line-block text-figures should be drawn with a good black ink,
such as ‘Process Black” or the best “Indian ink” on white
“ Bristol board.” All lines and dots must be clear and sharp,
and no washes or colours used. Wash drawings are not suitable
for reproduction as text-figures. All necessary lettering must
be clearly, but lightly, indicated in pencil, so that the engravers
2
may insert sharp, well-drawn letters. Hach drawing should be
numbered to tally with the text-figure number given in the MSS.,
and be accompanied by a short, concise, and Teeible descriptive
legend. When making drawings for text- figures, or grouping
drawings, it should be “borne in mind that, when reduced, the
figures must not exceed the dimensions of the printed matter on
the page of ‘ Proceedings,” viz.:—33 x 63 inches. Drawings or
photographs intended for reproduction as plates will have to be
reduced, when necessary, to fit a space of 4545 x 7 inches.
6. Copyrianr.—No use whatever is permitted of the type or
illustrations prepared for the Society’s Publications other than
for the Separate Copies mentioned above, unless the leave of the
Committee has been obtained, and recorded on the Minutes.
Permission is given only with the consent of the Authors of the
papers, if alive, and the source of the text and illustrations shall
be stated in the work in which they are used. The use of
illustrations from the Society’s Publications, if within five years
of the date of publication, will be granted only upon the payment
of half the cost of their production. All communications made
to the Society shall, if accepted for publication, be regarded as
the property of the Society, and not returned to the Authors,
except for the purpose of comparison with the proof when set up
in oye
Note.—The Publication Committee is always ready to give
Authors who apply for it, permission to reproduce matter that
they have contributed to the Society’s ‘ Proceedings” and
“Transactions,” but the Society has been advised that a formal
assignment from Authors is necessary to enable it to protect
copyright against third parties,
ZOOLOGICAL SOCIETY OF LONDON.
o
Turis Society was founded in 1826 by Sir Sramrorp Rarruzs,
Mr, J. Santne, Mr. N. A. Viaors, and other eminent Naturalists,
for the advancement of Zoology and Animal Physiology, and for the
introduction of new and curious subjects of the Animal Kingdom,
and was incorporated by Royal Charter in 1829.
Patron.
HIS MAJESTY THE KING.
COUNCIL.
HIS GRACE THE DUKE OF BEDFORD, K.G., F.RS., President,
Con. 8. Monckton CoPeman,
M.D., F.R.S.
Cuarters Drummonp,
Treasurer.
Aurrep Ezra, Kse., O.B.E.
Hueu 8. Grapstonn, Esa., M.A.,
F.R.S.K.
Tue Rr. Hon.
Grey oF Fattopon,
P.C., Vice-President.
Sr Sipney F. Harmer, K.B.1.,
M.A., 8c.D., F.R.S., Vice-
. President.
Pror. James P. Hitt, D.S8c.,
F.RB.S., Vice-President.
Lt.-Cot. tun Lorp Atastarr
Rosert Innes-Ker, D.S.0.
Str Waurer Rorrr Lawrencn,
Br., G.C.I.E., G.C.V.0., C.B.
Ksa.,
YHE VISCOUNT
K,G.,
Cot. Sir Henry McMaunon,
G.C.M.G., K.C.LE.
E. G. B. Mrapz-Watpo, Esa.,
Vice-President.
P, Coatmers Mrronet, Ese.,
C.B.E., M.A., D.Sc., LL.D.,
F.R.S., Secretary.
THe Eart or Onstow, O.B.E.,
Vice-President.
Masor Atsert Pam, O.B.E.
Lr.-Cot. Sir Davin Pray,
C.M.G., C.I.E., F.R.S.
Tux Lorp Qurnyzonoven.
Harotp Joun Hastinas
Russet, Ese.
Tue Maravess or Sido, F.S.A.,
Ricuarp StepHens Tartor, Esq.
A. Smita Woopwarp, Ese,
’ LL.D., F.R.S., Vice-President.
2
The Society consists of Fellows, Imperial Fellows, Honorary,
Foreign, and Corresponding Members, elected according to the
By-Laws. It carries 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 “Tho 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.R.M.S., F.R.P.S., F.Z.8., is open daily (except Sunday) from
Ten a.m. till Five p.w.; on Saturdays, Ten a.nt. till One p.m.
‘The Library is closed from:Good Friday to Kaster Monday, and
upon all other Bank Holidays. It is also closed annually for
cleaning during the month of September.
The Meetings of the Society for General Business are held in
the Meeting Room at the Society’s Office on the third Wednesday
of the month at 4.30 p.m. except in September and October.
‘The Meetings for Scientific Business are held in the Meeting
Room at the Society’s Office fortnightly on Tuesdays, except in
July, August, September, and December and January, at half-past
Five o'clock p.m.
The Anniversary Mente | is held on the 29th of April, or the
nearest convenient day, at Four P.a.
Tho Society’s Gardens are open daily from Nine o’clock until
Sunset, or 8 p.m. when sunset is later. Dr. G. M. Vevers is the
Superintendent, Mr. D. Seth-Smith, Curator of Mammals and
Birds; Mr, Edward G. Boulenger is Director of the Aquarium ;
Miss Joan B. Procter, F.L.S., is Curator of Reptiles; Miss L. EK.
Cheesman, F.E.S., is Curator of Insects; Dr. C. F. Sonntag
is Anatomist; Dr. N. 8S. Lucas, Pathologist; Prof. Rk. T. Leiper,
F.R.S., with the assistance of a panel of experts, examines
and reports on the Parasites; Prof. G. H. Wooldridge, F'.R.C.V.S.,
is Honorary Consulting Veterinary Surgeon; and Dr. R. W. A.
Salmond, O.B.E., Honorary Radiologist to the Society. Applica-
tions for anatomical or pathological material, or for facilities for
work in the Prosectorium should be addressed to the Secretary.
TERMS FOR THE ADMISSION OF FELLOWS.
Futtows pay an Admigsion Fee of £5, and an Annual Contri-
bution of £3, due on the Ist of January, and payable in advance,
or a Composition of £45 in lieu thereof; the whole payment,
including the Admission Fee, being £50.
No person can be deemed a I'ritow until the Admission Fee
and first Annual Subscription of £3 have been paid, or the
annual payments have been compounded for.
Fextows elected in November and December are not liable for
the Subscription for the vear in which they are elected.
PRIVILEGES OF FELLOWS.
Fittows 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 Hussanp of a Futtow can exercise these privileges
in the absence of the Tellow.
Until further notice, Frtnows will receive 40 undated Green
Cards, available on any Sunday or week-day up to tho end of
February of the year following the ycar of issue, and 20 White
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of the Green Cards may be exchanged for a book containing two
Orders for each Sunday in the year. ‘Twenty White Cards may
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children’s tickets are no longer issued, but the Green and White
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Tttows are not allowed to pass in friends on their written
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Frttows have the privilege of receiving the Society’s ordinary
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Fxttows also have the privilege of subscribing to the Annual
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and Publications relating to Zoology in each year, for the sum of
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.
Frttows may obtain a 'ransrerabcu Ivory Ticker admitting
two persons, available throughout the whole period of Fellowship,
on payment of ‘len Pounds in one sum. A second similar ticket
may be obtained on payment of a further sum of Twenty Pounds.
4
Any Fertow who intends to be absent from the United Kingdom
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pay the balance of the ordinary subscription. After three years a
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ON THE SKULL OF CARNIVOROUS DINOCEPHALIAN REPTILES, 661
34. On the Structure of the Skull in the Carnivorous
| Dinocephalian Reptiles. By R. Broom, M.D., F.R.8.,
C.M.Z.8.
[Received May 7, 1923: Read June 12, 1923.]
(LText-figures 1-17.)
Inprx.
Page
Scapanodon duplessist Broom.........0checcecsecee cee ceeseeseeeneess 663
Dinophoneus ingens, gen. Cb SP. NOV. ..........ee cee eee eee eeeene sss 6866
Dinartamus vanderbyli, gen. et sp. NOV. .......cecce cee eaeeeees 669
‘Tapinocephalia, new Suborder of Dinocephalia .................. 671
Titanosuchia, new Suborder of Dinocephalia...................... 671
Burnetia mirabilis, gen. et sp. nov. ........ HSE OTE
Burnetiamorpha, new Suborder of ihoredonuall Spies OFS
Classification of the Therapsida .............0ccecceeseee 674
Addendum ........... A una, SOD
Mandible of Tubanosuahis Shek siecorn eatin oatsnsaeyetns ees Meee 676
Tinobius strubeni, gen. Cb Sp. NOV. ..eccescecsesseeeesseseeneeseeees 683
Though our knowledge of the structure of the Anomodontia
and of the various suborders of the Theriodontia is fairly com-
plete, the more primitive groups—the Dromasauria and the
Dinocephalia—-are still very imperfectly known. Of the Dino-
cephalia we know a good deal of both the skull and skeleton. of
the Tapinocephaloid group, but of the Zitanosuchus-like forms
we know very little of either the skull or skeleton. The present
short paper will add something to our knowledge of these latter
types.
The ehief reason why so little is known of the giant Dino-
cephalians while the Anomodonts and Theriodonts are so well
known, is that the Dinocephalians in South Africa are in older
beds, where they cannot be easily picked out from a loose shale,
but are usually in masses of extremely hard limestone. If
unweathered out it is only with great difficulty that the bone
can be distinguished from the matrix; and when weathered
usually much is hopelessly gone and probably much still in the
very hard stone. Hence it happens that only a very few skulls
are in any museums, and, so far as I know, only two mounted
skeletons.
The first Dinocephalian remains from South Africa were
collected by Dr. W. G. Atherstone in 1872 and described by
Owen in 1876. The front end of a snout of a large form was
named Z'apinocephalus atherstonei. Some vertebre and other
remains were referred to this species, but there is some doubt
Proc. Zoou. Soc.—1923, No. XLIV. 44
662 DR. R. BROOM ON THE STRUCTURE OF THE SKULL
whether the reference is correct. Nothing was known of the
affinities of the form until many years later.
In 1879 Owen described under the name Zitanosuchus ferox
some jaw fragments that had been discovered by Mr. 'T. Bain.
Many years earlier, fragmentary remains of reptiles which we
now know are allied to the South African Dinocephalians were
described from the Permian rocks of Russia. Seeley in 1888
visited Russia, examined the specimens described, and figured
two genera represented by fair skulls—Deuterosaurus and Rhopa-
lodon. In 1892 he described the first good skull of a South
African Dinocephalian—a skull in the Cape Town Museun
collection,—which he named Delphinognathus conocephalus, and
he founded the order Dinocephalia for the reception of this form
and Vapinocephalus.
For many years nothing further was added to our knowledge
of the Dinocephalia. When in London in 1909 I examined
some specimens which had been in the Seeley collection. The
most important was a very fine skull of a large Tapino-
cephaloid reptile, which I referred to Vapinocephalus atherstonet,
and of which I gave figures. Another important specimen was
the top of the head of a Titanosuchid, which I referred to
Vitanosuchus ferox, In 1910 I gave a re-description of Delphino-
gnathus. Between 1911 and 1915 in various papers I gave
figures and descriptions of the skull and the principal bones
of the skeleton of the small Dinocephalian, Aoschops capensis.
Krom imperfect remains f{ had previously described a considerable
number of Dinocephalians—“ccasaurus priscus, Pelosuchus
priscus, and Tawrops inacrodon of the Tapinocephaloid group, and
Archeosuchus cairnerossi, Titanosuchus cloetet, and Scapanodon
duplessist of the 'Titanosuchid group,
In 1914 Watson described fully all the material in the British
Museum. ‘lhe skull which I had referred to V'apinocephalus
atherstonet he believed to be a new genus and species, and named
it Mormosaurus seeleyi. A second imperfect skull he called
Pnigalion owent. A third very imperfect skull he named Lamia-
saurus newton. ‘This third skull has a snout of the Titanosuchid
type, but an occiput rather of the Tapinocephaloid type; and
when in 1914 I examined the specimens in the British Museum,
I could not feel at all satisfied that the snout belonged to the
occiput. [am therefore pleased to see in Watson’s paper on the
Theriodontia (1921) that he also recognized there may be an
error in referring the snout and occiput to the same animal. ‘To
avoid confusion it will be well to regard the snout as the type of
Lamiasaurus newtoni. Watson refigured the top of the skull of
the Titanosuchid which I had referred to Zitanoswehus ferox,
adding the premaxilla, which he found belonged to it. In 1921
he gave new restorations of the skull of Rhopalodon fischeri, and
finding further fragments of the supposed Z'itanosuchus skull,
discovered that it belonged to a distinct type which he named
Anteosaurus magnificus.
IN TIEE CARNIVOROUS DINOCEPHALIAN REPTILES. 663
In recent years Haughton has described a complete skull of
Tapinocephalus atherstonet and a new Tapinocephaloid, Struthio-
cephalus whaitsi, and the skull of a small and probably immature
Titanosuchid under the name of MJoschosaurus longiceps.
In 1914 van Hoepen briefly described a new Titanosuchid under
the name Jonkeria truculenta.
ScAPANODON DUPLESSISI Broom.
In 1904 I gave under the above name a brief description of a
new type of Titanosuchid reptile. ‘The type consisted of jaw
fragments with teeth. It was distinguished from Titanosuchus
Jerox Ow. by having relatively much smaller molar teeth, and in
those molars having flattened spade-like crowns. ‘The type came
from Zeekoegat, in the Prince Albert district. Nothing further
has been known of this form till recently.
A few months ago I visited Mr. W. van der Byl at Abraham’s
Kraal, also in the Prince Albert district, and looked over the
large number of specimens that he had collected in recent years
on the farm. Among these was the weathered skull of a large
Titanosuchid. On critical examination this proves to be
Yext-figure 1.
Side view of skull of Scapanodon duplessisi Broom. About 4 nat. size.
Scapanodon duplessisi, and reveals to us the main features of the
skull apart from the palate. ;
The skull has the nasal, frontal, and parietal regions badly
weathered, and the matrix where unweathered is extremely hard.
Both sides of the skull are fairly well preserved, and portions of
all the teeth of the upper jaw are to be seen.
The general appearance of the skull will be best understood
from the figures given. Though the bones of the frontal region
are greatly thickened, there is no evidence of great bony bosses
overhanging the orbits as in Antessaurus. The pineal foramen
is very large, and the occiput is very much flatter and broader
than in. Anteosaurus. ‘The whole skull is also relatively much
broader in Scapanodon.
44®
664 DR. R, BROOM ON THE STRUCTURE OF THE SKULL
There are five large incisors in each premaxilla. ‘he teeth
are not well enough preserved to show whether they were
pointed or had, like the front teeth of the Tapinocephaloids, a
large basal keel. But there seems a little evidence in favour of
at least an expanded base. The roots are very long. The fifth
incisor is directed more outwards than the others. Between the
fifth incisor and the canine is a fair-sized diastema in which the
lower canine lies when the jaws are closed. In the very large
majority of the Karroo carnivorous reptiles the lower canine
passes into a deep cavity in front of the upper canine, but inside
the edge of the jaw. In Scapanodon there is no doubt that the
canine passed up outside the edge of the jaw, as portions of both
lower canines are preserved occupying the space between the
fifth incisor and the upper canine.
Text-figure 2.
Upper view of skull of Seapanodon duplessisi Broom. About 7 nat. size.
The upper canine is a large rounded tooth.
The molars are apparently 14 in number. ‘The anterior are
the largest, and they steadily decrease in size in passing back-
wards.
The nasal bones are completely weathered away, and the
frontals badly weathered and largely gone. Tnough is left to
show the upper margin of the orbit. From the structure of the
parts of the frontal remaining it is probable that there was a
considerable thickening of the bones in the frontal region.
Though portions of the prefrontals and lachrymals remain, the
limits of these bones cannot be made out.
The jugal bones are, however, well preserved, that of the right
side being practically perfect. In front it passes well forwards
above the posterior end of the maxilla. It forms the whole of
IN TILE CARNIVOROUS DINOCEPHALIAN REPTILES. 665
the lower border of the orbit. Posteriorly it forms with the
squamosal the short, deep, lower temporal arch. The posterior
process of the jugal reaches to the descending part of the
squamosal. Its lower border is curiously and very coarsely
roughened.
The squamosal is a very large bone with an ascending, a
descending, and an anterior process. The ascending process forms
the back of the temporal fossa, and articulates with the parietal
and the tabular. The descending process passes downwards and
slightly forwards. It has a moderately developed outer side and
a larger occipital face. The occipital face almost completely
covers the huge quadrate.
There is a small quadrato-jugal which must be very largely, if
not completely hidden from external view by the squamosal.
The quadrate is very badly weathered, but must have been a
Text-figure 3.
Oblique section from the nasal region to the largest part of the pterygoids,
in Scapanodon duplessisi Broom. About + nat. size.
large bone. The upper portion lies far up inside the folded
squamosal.
The occiput is not very well preserved, but appears to be
essentially similar in structure to the fairly well preserved
occiput of the genus to be next described.
The parietals are very thick and the pineal foramen large.
The postfrontals and postorbitals are very badly preserved.
Much of the lower jaw can be seen on one or other side. The
general structure appears to be fairly similar to that of the jaw
in the Tapinocephaloids. The dentary has a distinct though
probably short coronoid process. ‘The angular is large, and forms
the greater part of the outer side of the back part of the jaw.
The surangular is a powerful bone with a very thick upper edge,
which, when the jaw is closed, lies outside the descending process
of the squamosal as shown in text-fig. 1.
666 DR. R. BROOM ON THE STRUCTURE OF THE SKULL
A slightly oblique transverse section through the skull shows
something of the structure of the palate. ‘lhe section is illus-
trated in text-fig. 3; it passes from what is the middle nasal
region above to near the middle of the deepest part of the ptery-
goids, and shows the great size of the pterygoids. Inferiorly
they are somewhat parted, but above they unite in forming a
very high median crest. On each side a section of the ectoptery-
goid is seen, and above this the posterior spurs of the palatines.
On one side a considerable portion of the maxilla is cut across;
on the other a part of the jugal and a very small part of the
maxilla,
Owing to the extreme hardness of the matrix, no attempt has
been made to clear the palate; and as in the following type the
palate is fully revealed, it seems unnecessary, for doubtless the
structure will be very similar in Scapanodon to that seen in the
other genus.
DINOPHONEUS INGENS, gen. et sp. nov.
This new genus and species are founded on two skulls discovered
by Mr. M. J. van Wyk on the farm Kookfontein, in the Prince
Albert district. Though the two skulls are apparently the same
species, to avoid any possible confusion specimen B will be
regarded as the type.
Specimen A consists of a nearly complete skull which has the
Text-figure 4.
Side view of skull of Dinophoneus ingens, gen. et sp. nov. About } nat. size.
frontal and parietal regions and part of the occiput beautifully
weathered out, but with the lower half of the back of the skull
completely enclosed in very hard matrix. Most of the snout is
preserved, but weathered above badly and with the front teeth
mostly gone. The greater part of the left mandible is preserved, .
but much of it isin hard matrix. ‘Che front and back parts of
the left mandible ave preserved in fairly good condition. Of
specimen A there is a fine scapula and, presumably, other skeletal
i
IN THE CARNIVOROUS DINOCEPILALIAN REPTILES. 667
remains—another scapula, a humerus, and many vertebr—also
belong to it.
Specimen B consists of the greater part of a second skull of
apparently the same species. This skull was found near the
other, and is made up of a large number of considerably weathered
scraps. Fortunately it is possible to join all the principal
portions, and it is found that nearly the complete skull can be
restored. The palate is practically perfect from the occiput to
the front of the prevomers. Much of both maxille are present,
but very little of the premaxille. Much of the left jugal arch
is preserved, and the whole of the occipital and squamosal regions
are nearly complete with the parietals and frontals above. ‘hese
latter are, however, much weathered.
Text-figure 5.
Upper view of skull of Dinophoneus ingens, gen. et sp. nov. About } nat size.
The details of the frontal and parietal regions are from Spec. A.
From the two skulls it is possible to make out all the more
important details of the cranial structure.
‘The premaxillary region is not very well preserved in speci-
men A, and is lost in specimen B. It is manifest, however, that
there have been four large incisors, but none of these is fully
preserved. As one has the typical Dinocephalian crown, and the
lower incisors preserved also have the Dinocephalian crowns, we
may infer as highly probable that the incisors all have this
peculiar type of crown. The canine is very large, but not well
preserved. Behind the canine are 13 molars, which gradually
decrease in size on passing backwards. The thirteen occupy a
space of about 143 mm. The crowns are not preserved.
668 ° DR. R. BROOM ON THE STRUCTURE OF THE SKULL
In both specimens the nasal region is lost, but in both the
frontal and parietal regions are preserved—in specimen A in
perfect condition ; in specimen B considerably weathered.
‘The shape of the frontal and parietal bones will be best under-
stood from the figures. ‘The frontal is moderately flat in front,
but bends upwards behind to articulate with the elevated
parietal. he interorbital width is 117 mm. ‘There is a fairly
large prefrontal which forms much of the supraorbital margin.
The postfrontal is a small bone wedged in between the frontal
and the postorbital. The postorbital is large. It forms appa-
rently the whole of the postorbital arch, and passing upwards
forms about half of the upper wall of the temporal fossa.
The parietal is narrow, but very massive. ‘The pineal foramen
‘Text-figure 6.
be, yj Fan
MY
Zs0rn6 G000000
oe
SSS
Ue,
Y \ \\ Yj "
re
i 96 eat
it \ ii )
Palatal view of skull of Dinophoneus ingens, gen. et sp. nov. About + nat size.
5 Entirely drawn from Spec. B.
is large and round. Some distance behind the foramen the fossz
are only 5°3 mm. apart.
The jugal is not well preserved, but specimen B has a large
isolated fragment of it. In this fragment is also much of the
anterior part of the squamosal.
The squamosal is a very large bone. It forms most of the
posterior wall of the temporal fossa. It has a large anterior
process which forms much of the subtemporal arch, and a still
larger portion of the bone passes downwards and forwards to
support the quadrate. The squamosal also forms a large part of
the back of the skull, as is shown in text-fig. 7, The quadrato-
jugal is a small bone which lies inside the squamosal for the most
part and also supports the quadrate.
IN THE CARNIVOROUS DINOCEPHALIAN REPTILES. 669
The quadrate is large, and extends far up inside of the
squamosal. It has a large internal and anterior process which
articulates with the posterior process of the pterygoid.
The occiput is fairly similar to that of the typical Dinocepha-
lians such as J/ormosaurus figured by Watson. There is a broad
interparietal, and the tabular is large and extends down between
the squamosal and the supraoccipital. <A. fairly well developed
stapes can be seen in position.
The palate is beautifully preserved in specimen B. ‘The
prevomers are very large bones which extend nearly half of the
whole length of the palate. The palatines run along the inner
sides of the maxille and pass backwards to meet the prevomers,
the pterygoids, and the ectopterygoids. The ectopterygoids are
relatively small, and are situated as shown in text-figs. 6 and 4.
The pterygoids are larger and peculiarly twisted bones. The
Text-figure 7.
Occiput of Dinophoneus ingens, gen. ct sp. nov. About § nat, size.
Entirely drawn from Spec. 13.
anterior part of the palate formed by the prevomers, the palatines,
and anterior parts of the pterygoids passes backwards and down-
wards as far as the huge pterygoid transverse processes. Here
the pterygoids bend back on themselves nearly at right angles,
and form processes which pass upwards and backwards to articu-
late firmly with the quadrates, much as in Sphenodon.
On the base of the skull between the posterior processes of the
pterygoids is a typical Therapsid vomer or “ parasphenoid ” as it
is called by most writers.
DINARTAMUS VANDERBYLI, gen. et sp. nov.
This remarkable new type is founded on some skull fragments
discovered at Abvaham’s Kraal by Mr. W. van der Byl, whose
keen interest in paleontology has resulted in the discovery of
quite a considerable number of new types. Ihave much pleasure
in associating his name with the find.
670 _DR. R. BROOM ON THE STRUCTURE OF THE SKULL
The type consists of most of the preorbital portion of a skull
with much of the left postorbital region and with an incisor
portion which probably belongs to the same skull, but the contact
-has been lost. Besides these portions, which probably belong to
one skull, there is much of the postorbital portion of a second
skull, and much of a lower jaw which may belong to either of the
skulls—more probably to the first.
There are four large incisors, of which the first is of Dino-
ccephalian type, but the other three incisors may have had pointed
crowns. ‘There is some little indication that this may have been
so. ‘The canine is large and somewhat curiously bent. There
ave five well-developed molars which occupy a space of about
54 mm,
In the figure I give (text-fig. 8), some idea of the shape of the
skull will be obtained. The surface of the bone is flaked off, so
that the sutures given may differ a little from those that would
appear on the surface.
Text-figure 8.
Side view of skull of Dinartamus vanderbyli, gen. et sp. nov.
About } nat. size,
The lower jaw which I believe to belong to this species and
probably to the same skull is quite remarkable. ‘here are
apparently three incisors and what look like two large canines.
lt is probable, however, that the first is the true canine, and
that the other is an enlarged first molar. Following the
enlarged first molar are four fairly normal molars. If this
view be correct, the dental formula of Dinartamus will be
1 5
BD Tratthorge
Tt will be seen that Dinartamus differs considerably froin the
typical Titanosuchids such as Scapanodon and Dinophoneus, and
ought to be placed in a distinct family which may be called the
Dinartamidee.
IT do not intend in the present paper to enter into the dis-
cussion of the aflinities of the Titanosuchids with the ‘Tapino-
cephaloids, or the more interesting question of their aflinities with
the other Therapsids. But in the meantime one may suggest
a. Cc.
f
"IN THE CARNIVOROUS DINOCEPHALIAN REPTILES, 671
the advisability of dividing the Dinocephalia into two sub-
orders—the Tapinocephalia and the Titanosuchia. ‘The latter
suborder would have in it two known families—the Titano-:
suchide and the Dinartamide.
BURNETIA MIRABILIS, gen. et sp. nov.
The specimen on which this new genus and species are founded
is a nearly perfect skull, without the lower jaw, found by me
at Water Krantz, in the extreme north-west of the Graaff Reinet
division where it adjoins the Richmond division. At the point
where the specimen was found, fossils are very rare, and this
specimen was the only one discovered in a couple of hours’ hunting
on good exposures. ‘The horizon is most probably’ the lower
beds of the Lystrosaurus zone.
The skull is quite unlike any other that has ever been found
in South Africa or elsewhere; and only now, five years after its
Text-figure 9.
Side view of skull of Burnetia mirabilis, gen. et sp. nov.
A little less than 4 nat. size.
discovery, do I venture to describe it. There seemed a possi-
bility that it might be a late degenerate Titanosuchid, more
especially as it has large bony bosses over the orbits somewhat
like those seen in the British Museum specimen which Watson
has called Anteosauus magnificus. After patiently waiting all
these years, I find it convenient to describe it in this paper,
which includes an account of all the important details of Titano-
suchid structure, because it has some superficial resemblances to
Dinocephalians, although not at all a member of that group.
The skull as preserved is 177 mm. long, and was probably,
when perfect, about 190, and is 170 mm. wide. Unfortunately,
practically all tie outer bones of the skull are composed of a very
fine-grained, spongy bone, from which it is quite impossible to
remove the covering matrix except by grinding, and all trace of
sutures seems to be obliterated. As preserved, all the surface
672 DR, NR, BROOM ON HE SLRUCLURE OF 'THE SKULL
bone is covered as by a skin with a thin layer of calcareous matrix.
Though this obscures the margins of the orbits and temporal
fossw, all of which ave filled with matrix, it does not in any way
hide the real structure of the skull.
When seen from above, the most striking peculiarities of the
skull are the great posterior width, due to a remarkable develop-
ment of what are apparently the squamogals, and the presence
of five bony bosses—one above the nose, ene over each orbit, and
one in each post-temporal region.
The snout is relatively narrow. ‘The front part is unfortunately
missing, but from what is preserved it must have been sharp, as
shown in the figure of the side view, and very possibly it had a
horny covering as in Dicynodon. ‘There is evidence of one small
Text-figure 10.
=<
Upper view of the skull of Burnetia mirabilis, gen. ct sp. nov.
A little less than 4 nat. size.
incisor, a little in front of the canine, but I do not think there
could have been any in front.: The canine is a large, flattened
tooth which passes well forward. Behind the canine are four
small molars, The canine measures 12 mm.x5°5 mm. The
four molars occupy 18°5 mm., and the first is situated 4 mm.
behind the canine.
The orbits are small, and wide apmt and entirely lateral.
Above each is a large oval boss which completely hides the orbit
from upper view. .A deep transverse valley is present between
the back of the nasal boss and the fronts of the two orbital bosses.
There is a fairly wide space between the orbital bosses which is
transversely concave and antero-posteriorly slightly convex.
IN THE CARNIVOROUS DINOCEPHALIAN REPTILES. 673
Behind the frontal region is seen the pineal foramen, which is
relatively small but stands in a little rounded elevation of bone.
Vhe foramen stands in practically the middle of a very wide
parietal region. ‘The back part of the orbital boss is continued
as a less thickened bony avea above the small temporal fossa, and
then rises into the huge post-temporal boss which passes both
upwards and backwards. Whether this post-temporal boss is
formed by the parietal, tabular, or squamosal cannot be made out.
This. huge boss is continued outwards and downwards into what
is manifestly the squamosal region, and forms a huge ‘“ cheek”
not unlike that of a Pareiasawrus. That the cheek is not Pareia-
saurian in structure is manifest from the fact that there is an
undoubted temporal fossa.
In front of the temporal fossa there is a small boss on what is
probably the jugal bone.
The palate is fairly well preserved, but I have not at present been
able to satisfactorily clear it. So far as can be seen, it resembles
that of the Gorgonopsians more than that of other Therapsids.
As in the Gorgonopsian palate, there are a pair of dentigerous
areas partly formed by the posterior and main ends of the
palatines and partly by the pterygoids. The pterygoids have
large transverse processes like those of Therocephalians. There
appears to be, as in the Gorgonopsians, no suborbital opening
in the palate: if there is ove, it must be very small. The
posterior part of the pterygoid is very short; the whole distance
from the occipital condyle to the transverse pterygoid processes
is less than one-third the whole length of the palate. ‘The
quadrate is almost completely lost, but it must have been quite
small, and not likely larger than that in Cynodonts—possibly
smaller.
‘Nhe occiput has not been cleared. ‘here appears to be a small
median part formed by the occipital elements proper with the
paroccipitals, and two enormous lateral portions formed evidently
mainly by the squamosals,
his remarkable skull I propose to call Burnetia mirabilis after
James Burnet, Lord Monboddo, the famous Scottish evolutionist of
the eighteenth century, the friend of Boswell, of Johnson, and
of Burns.
The affinities of Burnetia are manifestly nearer to the Gorgon-
opsians than to any of the other known groups. It is fairly
certain that Burnetia is a Theriodont. The palatal structure
removes it quite clearly from the Dinocephalians and from the
Anomodonts. But while Burnetia has some affinity with the
Gorgonopsians, and may be descended from a Gorgonopsian
ancestor, the-remarkable specializations it has undergone seem
to justify me in placing it in a distinet suborder which may he
ealled Burnetiamorpha. At present it will stand as the sole
genus and representative of the sole family—the Burnetide.
674 DR. R, BROOM ON THE STRUCLURE OF THE SKULL
CLASSIFICATION OF TIE ‘THERAPSIDA.
Classification of fossil animals must always be more or less pro-
visional, and will have to be constantly altered as new forms are
discovered, and as new light is thrown on previously imperfectly
known forms. And yet we must always have a classification of
some sort to work by.
Watson, in his recent paper—-“ The Bases of Classification of the
'heriodontia” (P.Z.S. 1921), discusses at considerable length the
inter-relationships of the various groups of mammal-like reptiles.
The Theriodontia he divides into the following groups :—Gorgon-
opsidi, Therocephalide, Cynognathide, and Bauromorpha, With
this grouping 1am in entire agreement, but I would prefer to
regard the groups not as families, but as suborders. Seeley
placed Diademodon and Gomphognathus in a distinct order—the
Gomphodontia. Few would be willing to follow him thus far,
but one feels that though they are doubtless allied to Cyno-
qnathus, the differences are sufficiently great to justify their
being put in at least a distinct family. In the Therocephalians
also we have quite a lot of genera that can be conveniently
grouped together in families within the suborder. If we regard
the Therocephalians as a Family merely, we cannot classify the
large number of genera unless we resort to Subfamihes. All
living Birds resemble each other more closely than do the known
'Yherocephalians. Yet the living birds are subdivided into
Superorders, Orders, and Suborders. If the ‘Therocephalians are
only entitled to be regarded as a Family, then all living birds
ought to be placed in one Family, which would be rather in-
convenient. If, however, we regard the ‘lherocephalians as a
Suborder, it seems to me no harm will be done, and it will be
much more convenient than looking on the group only as a
Family.
Watson considers that “the three orders Deinocephalia,
Dicynodontia, and Theriodontia may have arisen from a common
stock whose direct conservative descendants are the Gorgonopsids,
and the Dromosauria may represent a more widely separated
stock of the Anomodontia.” 1 expressed a somewhat similar
opinion in a paper I sent to the Williston Memorial Volume
before Watson’s paper appeared. We will probably, however,
require many connecting-links before we will be quite sure of the
lines of evolution of the Therapsids, and I fear we will never get
those links in South Africa. In our oldest Karroo beds that have
land reptiles we get already typical Therocephalians, typical
Yorgonopsians, typical Dicynodons, and Dinocephalians. If we
ever get Ecca beds with good reptiles, we may then get the
common ancestors.
We do not, however, need to know the origins of of the groups
to get a useful classification, and the one which I would propose
is as follows :—
IN THE CARNIVOROUS DINOCEPHALIAN REPTILES. 675
Class REPTILIA.
Subclass SYNAPSIDA Osborn.
Super-order 7 7z£RAPSIDA Broom.
Order DINocEPHALIA Seeley.
Suborder TapinocePHALiA Broom.
~ TrranosucniA Broom.
Order DROMASAURIA Broom.
Order ANOMODON'TIA Owen.
Order THERIODONTIA Owen.
Suborder THrrocEPHALIA Broom.
» Gorconorsta Seeley.
5 BuRNETIAMORPHA Broom.
» BaurrtamMorrHa Watson.
» CynopontriA Owen.
[Received May 22, 1923.]
ApvprenpuM (27th April, 1923).—A few days after I had sent
off the above paper to London, I was very fortunate in receiving
from Mr. van der Byl one or two additional specimens, which
enable me to give a full description of the Titanosuchian jaw,
and thus to complete all the more important details of the skull
structure. On a part of the farm, Abraham’s Kraal, that had
not previously been prospected, Mr. van der Byl discovered the
weathered remains of three Titanosuchians, and collected all that
appeared to be portions of jaws. These, when pieced together,
proved to be the practically perfect mandible of a large ‘litano-
suchian, the badly weathered and fragmentary jaws of another
of the same species, and the dentaries of a huge Titanosuchian,
unlike any other that had previously been discovered. It is not
often that a collector makes at one time such an important haul ;
and when we consider that we owe to his discoveries previously
made a full knowledge of the skull structure in a group of
Mammal-like reptiles hitherto practically unknown, one feels
that Mr. van der Byl must be offered the most hear ty congratu-
lations of the scientific world.
The large jaw has preserved the roots of four fanee incisors,
the root of a large canine, and the remains of 15 molars. As
most Dinocephalians have lost the crowns of the teeth when
discovered, and have been named from the general size and
arrangement of the teeth, it is very difficult to be sure whether
we are dealing with new forms or with species that have
previously been named from very imperfect specimens. Owen’s
Titanosuchus ferox is admitted to be a most unsatisfactor y type,
676 DR. R. BROOM ON THE STRUCTURE OF THE SKULL
consisting of little more than the roots of the teeth. In 1903 I
made an “imperfect dentary in the South African Museum the
type of a new species of Z%tanosuchus, under the name 7’. cloctei.
It differs in size from Owen’s species, but otherwise appears to
agree sulliciently closely to make it seem safer to keep it pro-
visionally in the same genus. In 1915 Haughton added a third
species, 7’, dubiws, which also differs considerably in size from
Owen’s species, and less markedly from 7’. cloetei. Haughton’s
specimen came from Abraham’s Kraal, and had also been dis-
covered by Mr. van der Byl. The type of 7. cloctet is registered
as coming from ‘‘Gamka River.” As Abraham’s Kraal is on
the Gamnka River, it is not improbable that Haughton’s type and
mine came from the same farm, and in any case they are likely
to have come from the same horizon. This new jaw discovered
by Mr. van der Byl is so near to 7’. cloetei that I think it safest
to refer it to this species. 7’. dubius is probably a synonym, the
differences being such as might be accounted for by difference of
age or sex.
Mandible of 'Vitanosuchus cloetei Lroom.
The mandible discovered by Mr. van der Byl is nearly perfect,
but broken into ten portions. Six small portions are missing,
but four of these are quite unimportant, ‘The two others result
in the loss of the contact of the articular region of the jaw with
the front part, but so little is missing that the jaw can be
completely restored with practical certainty. The whole jaw
measures in length 565 mm., and, as will be seen from the figures
given, the depth is so little in comparison to the length that the
jaw appears slender. In reality it is quite a powerful jaw.
The dentition consists of four incisors, one canine, and a series
of 15 molars. ‘The incisors, though the crowns are not satis-
factorily preserved, are manifestly of the typical Dinocephalian
type. ‘I'he four together occupy a space of about 84mm. The
canine is rounded, and measures at the base of the crown 29 mi.
by 28 mm. The 15 molars measure 149 mm. None of the
crowns of the molars are well preserved, but from portions pre-
served it is manifest that they: have been flattened with, on
the inner side, vertical strie at the base. The anterior and
posterior edges are curved somewhat outwards and have a few
coarse serrations. ‘here is clear evidence of dental succession
at least as far as the 10th molar.
The dentary forms more than half of the outer side of the jaw,
but only about a third of the inner side. In front it is fairly
massive to support the powerful incisors and canine. There is a
large symphysis. ‘l'here is no coronoid process,
The splenial is an exceptionally long splint of bone which
forms nearly the whole of the lower half of the inner face of the
front half of the jaw. It forms a small part of the symphysis.
Its relations with the other bones will be best understood from
the sections given.
IN ''HE CARNIVOROUS DINOCEPHALIAN REPTILES. 677
The angular is a very remarkably constructed bone, A. little
in front of the middle region of the jaw the anterior end of the
bone is wedged in between the splenial and the dentary, and on
passing backwards we find the angular becoming steadily larger.
As seen in the figures given, it forms most of the outer side of
the back half of the jaw. On the inner side of the jaw it forins
a relatively small part, being largely hidden by the prearticular.
The very peculiar structure of the bone cannot be readily seen
except in section. It consists in the posterior part of the bone
being split in two, and containing a large cavity between the two
portions.
The coronoid is relatively 1 very small bone, so small that it
might be spoken of as rudimentary. Jt lies above the anterior
part of the prearticular, as can be seen in text-figs.11&12. It is
Text-figure 11.
—~—S>
SSS
-. ——
— =
Right mandible of Pitanosuchus cloctet Broom. Slightly larger than } nat. size.
so fragile that its anterior and posterior limits cannot be clearly
made out, the delicate bone being crushed and crumpled. That
a small coronoid is present is beyond doubt, and its limits must
be very nearly those shown in the figure.
The prearticular is a long, slender bone which runs forward
from the articular to about the middle of the jaw. It forms the
lower border of the long, narrow opening into the jaw. In front
it lies between the coronoid and the splenial. Its mode of
attachment to the articular and to the angular will be understood
from the sections given.
The surangular is a well-developed bone which forms the
upper part of the posterior third of the jaw. Anteriorly it passes
forwards between the dentasy and the coronoid and prearticular.
The specimen does not clearly show how much of the posterior
part is surangular and how much articular. Not improbably.
all the posterior strongly-developed ridge is surangular, and the
articular is entirely internal to it. Its relations with the other
elements are clearly seen in the sections.
Prov. Zoon. 8oc.—1923, No. XLV. 45
678 vit. R. BROOM ON THE SYRUCLURE OF THE SKULL
The articular is a strong short bone, which has on its posterior
fuce a broad concave articulation for the quadrate and possibly
the quadrato-jugal.
descending articular process.
on the inner side clasped by the prearticular.
I have figured six transverse sections of the jaw taken at the
points marked in the figures.
Text-figure 12.
Y
4
Y
AN
Si
SN
aD er
Transverse sections of the jaw of Vilanusuchus cloetei Broom at the points
indicated by the corresponding letters on the figures of the jaw.
fractures, they are so nearly trausverse that they may be regarded
us sections,
Section A, A.—'This section is through the 10th molar, which
is a replacing tooth just becoming functional. ‘The greater part
of the jaw is here formed by the dentary. Below and on the
inner side is seen a section of the splenial, and in the cavity of
the jaw are a few bony spicules, somewhat crushed and displaced,
which represent the anterior end of the angular.
Inferiorly there has been a _ powerful
The front part of the articular is
Though the sections are all
IN THE CARNIVOROUS DINOCEPHALLAN REPTILES. 679
Section B, B.—This section is immediately behind the last
molar. Tere the jaw is still mainly formed by the’ dentary,
which forms the whole of the outer side and about half of the
inner. ‘I'he splenial is still a fair-sized element, which forms
nearly half of the inner side. ‘he angular is, however, in this
section considerably larger than the splenial.
Section C, C.—This section, a little distance behind the previous
one, is near the posterior part of the dentary. ‘The largest
element in this section is the angular, which forms the main
portion of the lower half. On its outer side is a section of the
lower posterior process of the dentary. On its inner side is a
section of the posterior part of the splenial. The upper part of
the section is formed by the coronoid portion of the dentary.
lt is a powerful and curved structure. Underneath it is the
anterior process of the surangular. On the inner side of the
section is seen a large part of the prearticular cut across, and
wbove this is a section of the feebly-developed coronoid element.
Section D, D—This section is through the middle of the
posterior opening into the jaw. Above is a section of the
surangular capped by the coronoid portion of thedentary. Below
is the large angular with articulated to it the prearticular.
Section HE, i.—Between the previous section and this one the
angular splits into two plates, an outer and an inner. As seen
in this section, the inner plate is closely attached to the surangular,
and the outer plate, which is much the thicker, is quite free from
the inner both above and below. The inner plate has attached
to its lower end the prearticular, and between the two is the
posterior part of the opening into the jaw. The surangular is
still a powerful bone.
Section F, F.—This section is only a short distance behind the
preceding. The anterior part of the articular is cut across. It
is formed of soft spongy bone. Round its lower side lies the
prearticular. Above the articular and partly attached to it is
the large surangular. Outside the surangular is the inner plate
of the angular, xcept in its lower part it is quite thin. Above,
it broadens out into a delicate bony cap. In the specimen this
is somewhat crushed, and is restored in the drawing into what is
probably its original condition. Though close to the outer plate
it probably is not united to it. The outer plate is very thick.
Between the two plates is quite a large cavity.
Tn discussing the affinities of this Jaw with the jaws of other
early reptiles, we are hampered by the fact that in only a few
forms is the jaw structure fully known. In fact, as it is only in
the last twenty years that the jaw structure of living forms such
as the Lizard and Tortoise has become fully known, we palonto-
logists may perhaps be forgiven for not having yet been able to
reveal the structure of the jaw in all our extinct types. We do,
however, know the jaw structure fully in three groups of
Therapsids—the Anomodonts, the Gorgonopsians, and the
Cynodonts—and fairly well in the homodont Dinocephalians.
45*
680 DR. R. BLOOM ON THE SLRUCLTURE OF THE SKULL
We also know fully the structure in some of the Pelycosaurs.
In the early reptiles more related to the primitive Diapsidans
—the Ichthyosaurs and the Plesiosaurs—the structure is also
satisfactorily known.
Quite clearly the affinities of the Titanosuchian jaw are with
those of the Anomodonts and Gorgonopsians; and though in
each of those three groups the jaws have some quite charac-
teristic features, the jaw structure shows that there is a close
genetic relationship between the three.
If the Vitanosuchus jaw be compared with that of a Gorgon-
opsian, it will be seen that the two are fundamentally similar.
Text-figure 13,
A. Transverse section of the jaw of Dicynodon andrewsi Broom near the middle
of the jaw, for comparison with section CC of the jaw of Titanosuchus
cloetei Broom, ‘lhe curious overlapping of the prearticular by the splenial
is seen.
B. A similar section a little further back from another jaw of Dicynodon spec.
C. Section through the back of the jaw of a young Gorgonopsian, showing the
splitting of the angular. Compare with section KE of jaw of Titanosuchus
cloetei Broom.
he Gorgonopsian differs in having a much larger coronoid bone,
which serves as a splint to bind the back part of the dentary and
the front part of the prearticular. ‘The dentary has a well-
developed coronoid process which is lacking in Zitanosuchus.
The articular is much smaller in the Gorgonopsian, in keeping
with the very small size of the quadrate. ‘he Gorgonopsian
angular shows the same sort of splitting that is seen in Z%tano-
suchus ; but in the Gorgonopsian the outer plate is much less
fully developed.
The Anomodont jaw also shows marked affinities with that of
Titanosuchus, but these are to some extent: obscured by the
remarkable specializations. There have been one or two points
IN THE CARNIVOROUS DINOCEPHALIAN REPYILES. 681
in the Anomodont jaw concerning which opinions have differed
which must first be cleared up, and of these the most important
is the presence or absence of a coronoid. In 1910 I stated:
‘There is no coronoid process and no coronoid bone.” In 1912
Watson, in his interesting paper on Reptilian lower jaws,
describes the coronoid as ‘‘a thin slip of bone lying on the inner
face of the dentary, separated from it and the splenial by very
distinct sutures in several specimens, and apparently just entering
the symphysis.” And in his figure of the inside of the jaw he
shows what he regards as the coronoid. ‘Two years later he
expressed some doubts about the coronoid, and stated: ‘“‘I have
never been able to see a clear distinction of the bones in section,
and the apparent suture is, perhaps, something else.” In 1914
von Hoepen gave by far the best account of the Anomodont jaw
that has yet appeared. Ile paid minute attention to the question
of the presence or absence of a coronoid, and gives his conclusion
Text-figure 14,
re
Wigs)
.
Right mandible of the Anomodont Kannemeyeria simocephalus (Wicthofer).
very emphatically: ‘‘I can therefore definitely state that the
alleged coronoidal element between the dentary and the comple-
mentary does not exist in Lystrosaurus nor in Dicynodon.”
Sollas and Sollas in 1913 also stated that they could “ find no
trace” of a coronoid. The matter might have been left as settled
that there is no coronoid, but unfortunately the deservedly great
weight of Watson’s authority has resulted in his opinion being
accepted by Abel, and his figure of the Dicynodon jaw, showing
the supposed coronoid, being reproduced in ‘Die Stiimme der
Wirbeltiere.’
1n view of the difference of opinion if may not be regarded as
superfluous to give two new drawings of the Dicynodont jaw and
a couple of views of the jaw of #ndothiodon.
The jaw of Kannemeyeria, like all Dicynodont jaws, shows the
eculiar development of the angular. If the inner part of the
angular had an upper development against the surangular which
682 DR. R. BROOM ON THE STRUCTURE OF THE SKULL
met the upper part of the outer plate, the condition would be as
in Titanosuchus. he Dicynodout angular represents what was
probably the pre-Titanosuchian condition, ‘The inner view of
the jaw shows the large prearticular and the absence of the
coracoid, ‘The whole inside of the jaw is seen to be strikingly
similar to that of Zitanosuchus. The Dieynodon jaw might
almost be looked upon as a degenerate ‘Titanosuchian jaw which
had lost the rudimentary coronoid, lost the teeth, and developed
a new type of articulation.
The Hndothiodon jaw is a primitive Anomodont type. It
differs from that of Dicynodon in being a more loosely put
together jaw. ‘The prearticular is more slender, and the splenial
considerably longer. There is, as in Dieynodon, no trace of a
coronoid.
Text-figure 15,
Right mandible of Madothiodon angusticeps, sp. noy. ‘This is the jaw of the
Sndothiodon mounted in the American Museum which I had referred to
E. aniseries Owen. It differs in the arrangement of the teeth and in the
jaw being longer and narrower.
The Zitunosuchus jaw differs from that of the Pelycosaurs in
the fact that these have only a slight indication of the peculiar
development of the angular, and also that in these latter there is
a long well-developed coronoid. When I first described the jaw
of Dimetrodon, 1 thought there were in the Washington specimen
two coronoid elements. Williston on re-examining the specimen
considered that the supposed dividing suture is a crack. IT have
had no chance of again examining the specimen, but it is quite
likely Williston is right on this point.
Another jaw that may be compared with that of Titanosuchus
is the Plesiosaurian jaw. I do not consider the Plesiosaurs
nearly related to the Therapsids. Some others do, But it must
be admitted that there is considerable resemblance in the jaw
structure. VPelonewstes has a very large splenial, a long coronoid,
IN THE CARNIVOROUS DINOCEPHALIAN REPTILES. 683
and a small prearticular, The apparent resemblance is, in my
opinion, due to the fact that each is but a modification of the
early reptilian type.
I give two sections of the jaw of a Dicynodon' and one of a
young Gorgonopsian for comparison with the sections of the jaw
of Titanosuchus, ‘The first two show how essentially similar
Titanosuchus is to Dicynodon. The third shows how the angular
in the Gorgonopsian is split in a somewhat similar manner to
that seen in Titanosuchus.
Text-figure 16.
ay
7
Ye Sh
B.\s\eahab ee LS
Sp.
A. Right mandible of the Gorgonopsian Allurognathus tigriceps (Broom &
Houghton).
B. Posterior two-thirds of right mandible of the Plesiosaur Peloneustus philarchus
(Seeley), showing the prearticular.
It, is perhaps too early yet to definitely state the lines of
evolution of the Therapsids, but we are perhaps not far from the
truth if we suggest that the Titanosuchians, the Anomodonts,
and the Gorgonopsians have all sprung from a common arngastion
which was what might be regarded as a pre-Gorgonopsian.
I hope shortly to give a full account of the Gorgonopsian
skeleton, and I will “discuss at greater length the probable
affinities of the groups.
Enobius* stRUBENI, gen. et sp. nov.
This new genus and species are founded on a specimen con-
sisting of the two dentaries of a large ‘Titanosuchian reptile,
quite “different from any previously known. The snout of the
animal must have been very short and broad and blunt. In each
dentary there are the remains of four large incisors, and though
the crowns are lost, there is little doubt ‘that they have been of
the typical Dinocephalian structure. The four occupy a space
of 80 mm. About 7 mm. behind the fourth incisor is a very
* From ato Bias—fearfully strong.
684 ON THE SKULL OF CARNIVOROUS DINOCEPHALIAN REPLILES.
large canine which measures 38 mm. in length by about 30 mm.
in width. Behind the canine on the left side are portions of
three small molars. There have certainly been four, and very
probably there have been five. Those preserved are relatively
small, The second measures about the base of the crown
8mm.x7mm. These molars are arranged as shown in text-
fig. 17, On the right side the molar arrangement is different.
There is preserved only one molar, which is close to the canine,
and this is larger than any of the molars of the left side. The
section of the tooth measures 11 mm.x10 mm. Possibly the
first molar on the left side belongs to an earlier set. On the right
side I ean find no trace of other molars.
A considerable part of the posterior portion of the dentary is
preserved, which shows that while the jaw is broader than in
‘Text-figure 17.
Tooth-bearing portion of the dentaries of Hnobius strabeni, gen. et sp. nov.
A little more than + nat. size.
Titanosuchus or Dinophoneus, it is also relatively shorter. ‘The
measurement across outside the canines is about 215 mm., and
with its huge canines and short jaw this new form must have
been a much more lion-like carnivore than any of the previously
known types.
T have named this giant carnivore after Mr. Frederick P. 1.
Struben, the proprietor of Abraham’s Kraal, who for years has
taken a keen interest in paleontology, and as the result of whose
interest Abraham’s Kraal has become the most important, paleon-
tological centre in the Karroo. As Abraham’s Kivaal adjoins the
classical localities Jan Willem’s Fontein and Vers Fontein,
where the first South African Dinocephalians were obtained, it
is likely that ere long, with Mr. Struben’s assistance, we will be
able to clear up the confusion of the early types and specimens.
T'o face p. 685. | Pt. IV. 1923. Loveridge (Mammals).
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TANGANYIKA TERRITORY.
SHEWING PRINCIPAL COLLECTING LOCALITIES
1915 — 1923. i
Based on the R.E.A.A.A.Road Map by kind permission o
THE ROYAL EAST AFRICAN AUTOMOBILE ASSOCIATIG
Scale of Miles. Roads) ee racks 8
O 10 2030 4050 100 miles. eas Jways Swamps :
John Bale.Sons & Danielsson. L‘4
NOLES ON EAST AFRICAN MAMMALS. 685
35. Notes on Hast African Mammals, collected 1920-1923.
3y Antuur Lovarimar, F.W.S,, C.M.Z.S.
[Received August 14, 1923: Read October 23, 1928. |
CONTENTS.
Page
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INtRoDUCTION.
‘The present paper nay be looked upon as supplementary to the
1915-1919 notes *, as localities then recorded are not repeated
(except in one or two instances) or similar observations duplicated.
A new feature of the present paper is the inclusion of care-
fully ascertained native names from only those tribes in whose
district the specimens were collected. I consider it of importance
that these names should be collected as soon as possible for, with
the increasing facilities for travel, many tribes—especially along
the Central Railway— are rapidly forgetting their specific names
for animals and adopting the blanket-names of the town-bred
coast Swahili, who calls every rat, excepting Cricetomys, “ Panya.”
The rat-eating Wagogo and Wanyaturu, on the other hand, have
specific names for almost every species.’
T have avoided using names supplied by other tribes until that
particular animal has been collected in their particular district,
as when a native is asked for the name of an animal he is not
familiar with he will invariably supply you with the name of the
nearest creature in appearance to it that he knows, and at times
these may not be even related (¢.g., lemurs and squirrels, or
mole-rats and blesmols). For these reasons a very large per-
centage of the names collected by travellers and given by
Matschief are quite wrong (e.g., on p. 65, ‘“‘Uma mwitu” is
given for the jackal instead of ““Mbweha”). Umba ya mwitu is
the hunting dog, and means ‘dog of the bush.” Matschie gives
for its Chigogo (not Kigogo) name ‘ Mbua ya porini,” which is
merely the concoction of a bright native, as Iminzi is the correct
Chigogo equivalent. None but a naturalist can hope to obtain
* Loveridee, “ Notes on East African Mammalia.” Tn Jour. BH. A. & Uganda Nat.
Hist. Soc. No. 16, pp. 88-42; No..17, pp. 39-69.
+ Matsehie, “ Die Siugethiere D.O.A,” 1896. |
P.Z.8. Pt. [V. 1923. Loveridge (Mammals),
1'o face p. 685.
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LIwaLe
TANGANYIKA TERRITORY.
SHEWING PRINCIPAL COLLECTING LOCALITIES
1915 — 1923, S
Based on the R.E.A.A.A.Road Map by kind permission of
THE ROYAL EAST AFRICAN AUTOMOBILE AS
Ge
SOCIATION AN
Scale of Miles. Roads » “\ TUNSUROS
© 10 2030 4050 100 miles. - Tracks .._... ‘
Se RAIIWAYS deeetmtat Swamps Seto \
4
John Bale.Sons & Danielsson. L
686 Mh. A, LOVERIDGE: NOTES ON
native names of animals with any degree of accuracy, and only
then by a lengthy crogs-questioning can he be certain there is no
confusion.
In this connection I should like to express my thanks to the
Rev, John Briggs; who very kindly and thoroughly revised my
whole vocabulary of Chigogo names, to Messrs. Thomas and
Kershaw, for identifying all the primates, bats, and rodents, and
i good many in the other groups (most of the ungulates were
not submitted owing to the expense of freighting them); to
Dr. Baylis, Mv. Stanley Hirst, and the late Hon. N. C. Rothschild,
for kindly determining the parasitic worms, ticks, and fleas ; and
lastly, but not least, to my chief, C. F. M. Swynnerton, Esq., for
the encouragement and interest he has shown in my work, and
for permission to publish various notes jotted down whilst on
official journeys.
The material mentioned in the following pages consists of
920 skins and skulls, comprised as follows :—
No. of
Onder. Families. Genera, Species. Specimens.
Rrimates aan see 2 4 10 40
Cliicopteramer eee ae 8 14 22 200
Insectivora .).......... 3 D 13 43
Rodentia: pens, 9 25 47 417
Carnivora eee i) 15 28 120
Waiieulatare, coves: 4) 20 24 100
OS cooosunee 32 83 144 920
As for two years of the time the writer was stationed at Kilosa,
in the office of the Game Department, he hid to rely on native
collectors to a great extent, The success of the collecting is largely
due to the enthusiasm of two brothers, Salimu and Nikola, who
collected over 600 skins and exhibited the greatest keenness in
looking for novelties. It need not be supposed that the skins are
poor on this account ; on the contrary, I think they will compare
very favourably with any made by Europeans.
lt is to be regretted that in the great majority of cases the
taking of the measurements had to be left to Nikola, but after
checking him on many occasions I have sufficient confidence to
believe that they are of sone value, though it should he borne in
mind that a native does not usually see the necessity for such
accuracy as does the mammalogist. Where measurements are
given in this paper they are always in this order:—(i.) Length
from nose to vent. (ii.) Length of tail without terminal hairs.
(iii.) Length of hind foot without claws. (iv.) Length of ear
from tip to notch. In the case of bats a fifth measurement is
added: (v.) Length of wing, sometimes across outstretched wings
as well. All figures ave in millimetres.
Four new species and one new race have already been described
EASY AFRICAN MAMMALS. 687
from this material by Mr. P. 8. Kershaw *, viz., Vycteris maniea,
lephantulus renatus, Steatomys muanse, Arvicanthis tenebrosus,
and Rattus chrysophilus singide.
From Kast to West and South to North the principal localities
mentioned in this paper are :-—
TANGANYIKA ‘Terrrrory.
Dar es Salaam District.—-Dar es Salaam,
Morogoro District.—Bagilo, Mkindo River, Wami River, Mkata.
River, Rudewa, Longa, Chanziuru, Kimamba, Iondoa,
Tindiga, Mbala, Kisanga, Madazini, Miombo, Kipera, Kilosa,
Kidai,
Dodoma District. —Wibakwe, Ueakuyu. Mpaniva-kwa-Sagoi,
Kidenge, Itende, Pwaga, Lukole, Igulwe, Dodoma, Itumba,
Mbonoa, Ndogwe, Suna, Gwao’s, Mbulu’s, Pooma, Singida,
Mdjengo’s.
Kondow JIrangi.—Mlewa’s, Mtali’s, Mkalama, Zengeragusu,
Usshora, Usurwe, Ulugnu.
Labora District.—Vahora, Izikisia, Ndala, 'ambali, Luguo,
Wembere Ilats.
Micanza District.—Simbiti River, Nduguyu River, Sanga,
Lalago, Mwadira, Shanwa, Sagayo, Nyambita, Mwanza.
Bukoba District.—-Bukoba, Kakindu, Kabare.
INewya Conoxy.
Frere Town, Nairobi, Mldoret, Kisumu.
The notes on each animal have been arranged on a definite
plan. TVirstly, the native names, then distribution, habitats,
measurements, variations, breeding seasons, diet, habits, paia-
sites and enemies.
PRIMATES,
CERCOPILTILECGID”,
CERCOCEBUS ALBIGENA JoUNSrONE Lyd.
A. single pair of Johnston’s Mangabey were obtained from
Kakindu, Bukoba. ¢. 548. 740. 150. 30mm. 2. 550, 771.
130. 45 mm.
CERCOPITHECUS DOGGEFTL Poe.
A single pair of this monkey were collected at Kabare, where
they are very common in the great forests according to my col-
lectors. ¢&. 640. 845.50. 40mm. 2. 503. 700, 180, 40mm.
* Wershaw, “On a Collection of Mammals from Tanganyika Territory.” Amn.
& Mag. Nat. Hist. ser. 9, vol. xi. p. 686, May 1923. Kershaw, Aun, & Mag. Nat.
Ifist. ser. 9, vol. xii, October 1923.
688 MR, A. LOVERIDGE: NOTES ON
CERCOPITHECUS ALBOGULARIS RUFILAtTUS Poc.
Senghwa in Kikami and Kisagara, Kima in Kiswahili.
The Blue Monkey was met with at Bagilo, all along the Wami
and Mkata Rivers, common at Kilosa, Miombo, and Kipera, It
shows a marked preference for primary forest and the great trees
burdened with epiphytic growth which fringe the banks of the
larger rivers.
A young specimen which I had in captivity for some months,
and which was subsequently killed hy a leopard, was on several
occasions observed to eat ‘* cuckoo spit” of its own accord, and
with evident relish! ‘Lhe rapidity with which it pounced upon
insects—chiefly grasshoppers—and disposed of them was really »
remarkable. I have watched adults raiding maize plantations
at Tindiga, but they are not so addicted to thieving practices as
their relatives C. p. johnstoni.
A maggot of Caudylobia sp. had to be removed from the fore-
arm of the young monkey just mentioned.
CERCOPITHECUS PYGERYTHRUS JOHNSTONI Poc.
Negederi in Kikami and Kisagara, Numbili in Kinyamwezi,
Niadengwa i in Chigogo, Tumbili in Kinyaturu, Kinyiramba, and
Kiswahili.
Seen or collected at Bogoti, Tindiga, Kilosa, Kipera, Kidai,
Ttende, Kidenge, Mpanira, Ikikuyu, Kibakwe, ‘Lukole, Igulwe,
Usshora, and Ulugu.
‘The largest male measured 543. 650, 130. 35 mm., and female
ATO. 535. 121. 40mm. They probably breed all the year round:
notes of females with young at their breasts were made in
February (Igulwe) and July (Tindiga). A newly-born young
one was found in a shamba at Kilosa, it weighed 14 ozs., and
measured 180. 270, 65, 25 mm. (26, vil. 92),
A large female monkey in a dying condition was brought to
me by a native, who said it had been attacked by a Martial Hawk-
Kagle. As the native carried a bow and arrows I was rather
doubtful of the stor y, but on skinning it later no hole was found
but the skull was dinted and cracked, It had two old breaks in
its tail which were healed. At Kidenge a party of monkeys in
the euphorbious thorn-scrub on the side of the mountain raised a
great outcry and kept it up for half-an-hour; on enquiring the
reason from some natives who were idly watching, 1 was told
that an eagle had dropped down from the sky and was perched on
a rock near the monkeys (Kidenge, 14.11, 23).
The Bogoti specimen had red acarine parasites on its ears.
CrRCOPITHECUS PYGERYTHRUS CENTRALIS Neum.
Numbiri in Kisukuma.
Two from Sagayo and Kikindu, ‘The larger male measured
490. 620, 150, 301nin,
EAST AFRICAN MAMMALS. 689
Pario cynoceruAuus Linn.
{loku in Kikami, Nyabu in Kisagara, Nyaniin Kiswahili.
Hight examples of the Yellow Baboon were shot in the neigh-
bourhood of Kilosa, where it is all too common.
The largest male measured 858. 637. 237. 55 mm., and female
685. 565. 170. 481mm.
I believe they breed pretty well all the year round. Females
carrying young were met with in July and August, 1921, May
and July, 1922. A female was killed on 10,iv.22 with a
@ fwtus weighing 10 ozs. and measuring 171. 136. 52. 22 mm.
The young are sometimes carried on the back but generally, and
especially when very young, they cling to their mother’s breasts,
thus being back downwards when she is walking.
A baboon’s dietary is too large and varied to record ; they
break down the mulberry bushes when in fruit with great reck-
lessness, and exhibit this same lack of foresight in their ruthless
treatinent of paupau trees. In one instance, a tree which was
growing against the kitchen wall had its stem gnawed through
near the ground so that the tree, which was very heavy with
fruit, fel]. ‘Their love for paupau fruit emboldens them to come
within ten feet of the house.
One of my collectors set a snare for a bushfowl, and on visiting
the trap one morning found one leg in the noose and a baboon
eating the rest in a nearby tree. I have been told that in
Bagamoyo District the natives have the greatest difficulty in
keeping fowls owing to the baboons earrying them off, but
whether it is the present species that is concerned I cannot say.
If one is armed they are extraordinarily wary and difficult to
approach, and if suddenly surprised run away uttering piercing
screams, and at other times they treat one with more or less
contempt. Other animals seem to derive benefit from association
with baboon troops: a female bushbuck was seen with a company
in a rubber plantation on many occasions during the spring of
1921, and I saw the same or another running up the road in
company with a troop about 3 p.m. in the afternoon of 11. v. 22.
On 29. vi. 22, some ten wild pigs with a party of baboons passed
within two hundred yards of the house.
Their vitality is both astonishing and heartrending. Some-
body having fired at a baboon near the house one day was sure
that he had mortally woynded it, but I came upon the poor beast
all alone at 6 a.m. two days later. It was crouched upon a sheet
of galvanised iron by a deserted German house in the bush, and
was applying its jaw to the iron with the very evident object of
assuaging the pain, for its lower jaw was torn away on one side.
I shot it through the chest at close range, and yet it ran for sixty
feet before it dropped dead. I firmly believe the native story
that a wounded baboon will clap its hand over a wound and
successfully staunch the flow of blood, whether with the intention
of leaving no blood spoor I cannot say. Several times I have
690 Mh. A, LOVERIDGE! NOTES ON
shot baboons in trees, seen them fall, much blood at the foot, of
the tree, but no trail left.
‘Two instances of Yellow Baboons carrying off their companions
have come to my natice. In one casea local resident, M1. Ixostel-
lesky, shot a eonioen that was picked up by « still bigger
beast, which started down the hill with it. Myr. IKostellesky
raised an outcry which caused Capt. Turnley and his natives to
run out of the house, when they met the baboon with its burden,
which it dropped in fear for its own safety. The wounded
animal was too hard hit to be able to move,
The second case was witnessed by Mr. D. W. Bisshopp, who
shot a female carrying « young one on the Jvinga road. He
called to his natives to run and capture the little one, but as
they approached another female baboon returned —the rest of the
troop were in flight—imenaced the boys at close quarters, seized
the young one and inade off with it.
Mr. Kenny Dillon, of Kissaki, wrote me of a recent and quite
authentic case which occurred in his district in February, 1925,
where baboons attacked and killed the infant child of a native
woman while she was at work in the fields.
Their principal and almost only enemy at Wilosa is the leopard,
which they will mob when opportunity offers. Some notes on
this will be found under the heading of J”, pardus.
T have only once found a flea (Ctenocephalus canis) on a baboon,
and this was upon a healthy male in his prime who had alsoa
good many mites on his breasts and in their neighbourhood.
Another male had a large number of nematodes (Physaloptera
mordens Leip.) in its alimentary canal.
Pario NEUMANNI Matsch.
Mhuma in Chigogo, Pooma in Kinyaturu, Poma in fivamba.
Four specimens of this essentially rock-loving baboon were shot
at Mtali’s, Ulugu, Usshora, and Zengeragusu. They were also
seen close to Mkalama. They were always associated with rocky
kopjes or outcrops, and at Mtali’s a large troop slept on a huge
rock at the sumiit of a kopje.
The largest male measured 830. 500. 200. 55 1mm., and female
700. 390. 190. 50mm, A native brought a young one, which he
had had for two-and-a-half months, into camp at Zengeragusu.
Though kept on a leash it was very tame, and the air of abandon-
ment with which it lay back in its owner’s arms was truly
comical.
At Mtali’s the local medicine man came to me for the dorsal
vertebrz of an old male; these he claimed were a cure for weak-
ness and backache when worn on the lumbar region.
Pario 'ressunLaruM Klhot.
Kuku in Kisukuma,
Five specimens from Sugayo, where these baboons haunt kopje
GAs AFRICAN MAMMALS. 691
cotintry similar to that freyaented by Neumann's Baboon, but
sleep in the great trees fringing the course of the Simiyu River.
The largest male measured 890. 430. 220. 50 uin., and female
710. 440. 200, 50min.
Monkey malaria parasites were found in blood smears taken
from a specimen in whose stomach were many nematodes
(Streplopharagus armatus Blane).
LemMuninvsz.
GALAGO PANGANIENSIS Matseh.
Mwegi in Iikami, Kimwegi in Kisagara, Komba in Kiswahili.
Three specimens from the Rutigi, Ilonga, and Chanzuru were
taken alive. It was also heard along the Waini and Mkata
Rivers at many points, at Kimamba, Tindiga, and Kilosa.
The Honga specimen was only a day or two old when found
(24. 111. 23) by some natives felling trees and cleaving bush; it had
probably been dropped by the mother in escaping from a. tree
which was being felled. I kept it for eight days, feeding it on
neat sweetened condensed milk, and occasionally a little water.
It seemed to thrive on this diet and relieved itself naturally
und regularly. I therefore sent it back to Kilosa in charge of a
native who was looking after my other avimals: it died three
days later (go. 7. 98. 25. 181nm.).
The Chanzuru specimen would be about two months old when
caught (23. iii. 23), and became very tame, showing a great liking
for chicken bones. At the end of a month it was found to be
heavily infested with lice, which were destroyed by a single ap-
plication of parattin and camphorated oil well rubbed in. ‘Che
fur assumed its normal appearance in two days, but an eczema,
which had been present before, gained ground, and would not
react to treatment, so that I had to chloroform the little creature
on 23. v. 23.
The Rufigi lemur was adult when captured, and though I kept
it for three months it was wild and intractable; it had a very
hearty appetite, but having already described the dietary of a
Morogoro lemur there is no object in repeating it.
GALAGO SENNAARIENSIS Less.
Three specimens were purchased at Usshora, Sanga, and
Sagayo; a fourth which was brought to me alive escaped shortly
afterwards. Though adult it did not bite much, but drew blood
when it did. The very red eyes are rather a striking feature of
this little grey lemur.
3. 135, 230. 65. 38 nm. (Sagayo); 9. 180. 250. 65. 38 mum.
(Usshora). ‘The latter, killed by a dog, had a large fetus in utero
very new birth: 3g. 60, 75. 32. 151m.
692 MR. A. LOUVERIDGES NOTES ON
CLLLROPTERA.
Few native languages have specific names for the Bats, though
many distinguish between the Megachiroptera and Microchir-
optera. The general name for bats in Kinyamwezi and Kisu-
kuma is Tunge; in Kikami it is Ndema; in Kisigara, Goim-
barema, though in South Usagara Kipurabutu is often employed,
though it may be specific for Pipistrellus nanus. In Chigogo a
bat is Ibudibudi, in Kinyaturu Tai, in Kiramba Tule.
Fruit-bats in Kibena are Wembelema, in Kisagara Pulagutwi.
Tnsectivorous bats in Kinyamwezi are 'Tuvugulima, in IKibena
Mbudibudi.
Preropvrpa.
Rovuserrus weacnut A, Sm.
A. single specimen from Bagilo on 6. in, = 5 US0S all), AS):
30. 270 min.
BIwoLon wenyum lerr.
A single specimen was found hanging on a tree in a native
maize plot at Mahaka on 27. iii.22. 3. 195. 0. 35, 20. 364 min.
Kpomornorus LAbriarus T'emm.
‘’wo males of this Hpauletted Fruit-bat were shot in a cocoa-
nut paln and mango tree at Kilosa on 8. v.23. At times Fruit-
bats of this genus‘are very numerous, and at others only one or
two can be found after much search. ‘The larger. measured 120.
0. 20. 20. 190 mm.
MMBALLONURIDA.
TAPHOZOUS MAURLTIANUS Geoftr.
A specimen of the ‘'omb Bat was shot from the trunk of a
cocoanut palm at Kilosa, where they are numerous. One was
seen on a baobab at Itende. Parasites in the shape of Nycteribids
were found in the fur of the Kilosa specimen.
LIPONYCTERIS NUDIVENTRIS Cretz.
Twelve specimens from Mtali’s and Shandwa, where they were
found in fissures of the rocky kopjes by my collector. ‘The largest:
male measured 95, 15. 10. 10. 195 mim., and female 100. II. 14.
10. 202 min. ; both from the latter locality.
PRTLALIIDA.
Nycvreris CAPENSIS A. Sin.
Twenty-four specimens from Kilosa, ltende, Igulwe, Suna, and
Gwao’s from culverts, hollow baobab trees, a darkened vacated
EASY AFRICAN MAMMALS. 693
room in company with J/. caffer, and flying in a lighted room.
Quite a hundred individuals emerged from one baobab.
The largest male measured 58. 44. 10. 25. 125 mm., and female
55. 60. 10. 33. 130mm. ; both from Kilosa,
NYCTERIS HISPIDA Schreb.
A male from Madazini, near Kilosa, and a female from Bagilo.
The former flew into a lighted tent. do. 45. 38. 8. 15. 110 mm.
©. 60. 45. 5. 30. 120 mm.
NYCTERIS LUTEOLA Thos.
Nine specimens from Kilosa and Sagayo. The seven Sagayo
bats were brought to me by a native, who caught them in a burrow
occupied by a porcupine. Largest ¢ 65. 56. 11. 31. 160 mm.,
and 2 65, 64. 10. 35. 154mm.; both from Kilosa.
Nycreris Manica Kershaw.
A single specimen, the type, from Tindiga near Kilosa.
MreéaADERMIDA,
Lavi rrons rex Miller.
The Yellow-winged Bat was met with in thorn-bush at Msimba,
Kilosa, Mlewa’s, Mtali’s, Wembere Flats, Nduguyu River, and
Sagayo, but purposely not collected:
CARDIODERMA Cor (Peters).
Two male Big-eared Bats flew into a lighted room (8 p.m.) at
Usshora. ‘They measured 80. 40. 17. 38. 170 mm. and 70. 40. 17.
38. 185-400 mm.
TLirprosipERID”.
Riswonorxus Loparus Peters.
A single specimen flying in a lighted room at Kilosa. ¢. 55.
25. 8. 15. 130 mm.
One of these bats made three attempts to pick off a moth .
(Cylegramma latona) which was bumping along a lighted ceiling.
The third time it was successful (Kilosa, 14. v. 23).
HIprosIDEROS CAFFER Sund.
Thirty-two specimens of this Horseshoe Bat were taken at
Frere Town, Kenya, Kilosa, and Mbala, flying in a lighted room,
in an uninhabited room, and a rock cavern respectively. Of
these, nine were males and twenty-three females. ‘They exhibit
a great deal of variation in colouring, some being nearly as red
as JZ. ruber, others grey or brown. Largest g 59. 35, 7. 15.
130mm., and 9 56. 34. 7. 10. 130 mm. ;
Proc. Zoou. Soc.—1923, No. XLVI. 46
694. MR. A. LOVERIDGE: NOTES ON
One female had three huge maggot-like adult female dipterons
(Ascodipteron sp.) embedded beneath the skin of the elbow-joint
of both wings. Many had Nycteribids (Penicillidia pachymela
Speiser) in their fur, which Major Austen informs me has
previously been recorded from Somaliland but the host was
unknown.
HiPPostpDEROS RUBER Noack.
Five specimens from Kilosa and Mbala, where they were taken
in company with flocks of . caffer in an uninhabited room and
29
rock cavern respectively. Largest ¢ 59. 35. 8.14. 145. 333 mm.,
and 2 55. 33. 10. 14. 152-342 mm.
HiprosipERnos MARUNGENSIS Noack.
A single male of this huge Horseshoe Bat was shot at Iilosa
(8. v. 23) as it was hanging alone from the midrib of a banana
leaf at a height of 12 feet from the ground, the palm being near
. the bridge belowthe boma. gd. 106. 36. 21. 31. 300-642 mm. It
measured 25% inches across the outstretched wings. ‘lhe only
parasite was a mite (larva of a Trombidid) on one of the wing-
membranes.
VESPERTILIONIDA.
PiristRELLUS NANUS Peters.
Known as Kimburugutu in Kikami.
Three male Banana Bats were collected. Two of these were
flying at Kilosa, and one was found between the stem and the
leaf-stalks of a banana palm, so that they are not wholly depen-
dent on bunches of bananas as would appear from Lang and
Chapin’s field-notes. Largest g 40. 37. 8. 10 mm.
PIPIsrRELLUS KUMI FUscatus Thos,
A male from Bagilo on 6. v.22 measured 60. 40. 5. 7. 110mm.
GLAUCONYCTERIS ARGENTATA Dobs.
Sixty-four specimens from Kilosa were killed by two shots
from a ‘410 collecting-gun ; my collector had no idea that he had
killed so many till he began to gather them up. As many
more flew away from neighbouring banana palms, so that they
must be very numerous at certain seasons, Males predominate
in an extraordinary way, as there were only eighteen females,
All were measured, but 49 were almost identical in their dimen-
sions, viz. 50. 45. 8. 10. 140 mm.
The ovaries of all the females were examined for foetuses,
but without result. The stomachs of more than a score were
scrutinised, but the contents were too finely masticated for
recognition of any insect, nor were any internal parasites visible
LAS! AFRICAN MAMMALS, 695
under a hand-lens. Four mites (Pteroptus) were found upon
wing-membranes, but apart from these they appeared free from
parasites.
G{LAUCONYCYERIS VARIEGATA Tomes,
A single male was found hanging to the ceiling boards of the
verandah roof one morning at Kilosa, 18.1.22. ¢. 47. 53.7. 12.
150-330 mm.
Mo.LossiIpD&@.
CiAREPIION LIMBATUS Peters.
Vifteen Free-tailed Bats of this species were obtained from the
roof of my house at Kilosa, where they were a great nuisance on
account of the strong smell caused by the accumulation of their
defecations above the lining boards. ‘hey do not leave the roof
till it is very nearly dark. Between 4.30 and 5.30 a.m. they
return, making a great swishing noise with their wings—like a
ruler struck through the air, Sbis is preparatory to going to
roost for the day, which is done to the accompaniment of much
squeaking and twittering as they take their places and scuflle
along the ridge pole. It is by no means uncommon for thei to
fly against the gauze meshing of the verandah and fall to the
ground. ‘Though //. caffer is as common about the house it does
not do so, but skilfully avoids all obstacles.
he Jargest male mensured 56, 38, 7. 14, 125-282 min., and
female 60. 40. 10, 18, 128 mm.
CHAREPHON EMINI de Winton.
_ My native collector secured ten specimens from beneath a large
slab balanced on a boulder at a height of five feet from the
ground at Sagayo.
The largest of four males measured 73, 34. 8. 17, 146 min., and
of six females 72. 36. 7. 18. 155 mm.
Mors ossorni Allen.
Five specimens of Osborn’s Bat were taken in a hat at Kisumu,
where, to judge by the smell, they are excessively abundant in
many of the buildings. ‘This very distinct species has only been
known from Leopoldville, Belgian Congo, where it was collected
by the American Museum Expedition ; by the present record its
range is considerably extended to the east.
All the bats of this family are a nuisance when they take up
residence in dwellings, and J have found them very difficult to
dislodge. One of the best methods is to fill the space between
roofing-iron and ceiling boards with sprays of thorny mimosa,
whieh presumably tear their wings and cause them to seek shelter
in someone else’s house not so fortified.
The only male measured 84, 47. 13. 16. 189 mm., and largest
female 80. 45. 12. 16. 175 mm.
46*
696 Mr. A. LOVERIDGE : NOTES ON
INSECTIVORA.
HRINAORIDA,
ATELERIX HINDEI sotika Heller.
Kalunguyeye in Kinyamwezi, Kilungumesu in Kisukuma,
Kinenyifuri in Kibena, Sejesi in Chigogo (also in use among the
Wasagara in such parts of their district as the hedgehog occurs),
Kengye in Kinyaturu, Kilongomegia in Kiramba.
Of the five hedgehogs collected at Gwao’s, Pooma, and Usshora,
only one wasa male. Most of these were kept in captivity for
some time as well as others which escaped. They will drink milk
immediately after capture and eat meat readily after considerable
chewing. One was seen crossing the road in the moonlight and
ran with surprising swiftness. The natives of Singida District
stated that they were not at all uncommon. A young one no
larger than a tennis ball escaped from a deep wash-basin.
3. 180. 22. 29. 26 mm. Largest 9. 175. 18. 28. 25 mm.
Stomach of the latter was full of minced iisect remains and
parasitic worms (Physaleptera clause Rud.). At Pooma the body
of one was found disembowelled in the neighbourhood of a kopje ;
it had evidently been killed by one of the carnivora.
MACROSCELIDS,
RityNcmocyon Perenst prTErst Bocage,
Zagari in Kikami, Mbulusanje in Chigogo. All Elephant
Shrews are Sangi in Kiswahili irrespective of species.
A single pair from the Usambara Mountains, which is the type-
locality of 2. p. wsambare Neum., considered a synonym of 2. p.
peter'si.
RHyNCHOCYON SWYNNERTONI Kershaw.
‘Zozo in Kikami, but Konghole amongst the Wakami of
Mikesse.
A single pair of this recently described Hlephant Shrew were
obtained by my collector from the type-locality Kipera on
25. ix. 22, after a great deal of trouble, as they take refuge in the
long grass immediately they are sighted. I also saw one of these
spotted Hlephant Shrews cross the road near Madazini. The male
measured 370, 240, 67. 34, and the female 250. 205. 67. 31 mm.
Prrropromus MArscurE Neum.
Two females, an adult and immature, from Mahaka and Izikisia
respectively, the adult measuring 220. 170. 50. 38 mm.
PETRODROMUS NicrisETA Neum.
Sangi in Kikami and Kiswahili.
Two were collected at Kipera; it was also occasionally seen
at Kilosa, Chanzuru, Kimamba, and Bogoti.
BAST AFRICAN MAMMALS. | 697
ELEruANTULUS PULCHER Thos.
‘Ten specimens from Sanga, Lalago, and Sagayo. Largest male
measured 140, 110. 31. 20 mm., and female 140. 121. 31. 23 mm,
Many ticks (2thipicephalus sp. nymphs) weve taken from the nape
of one specimen.
ELeriuANtutus ocuLanis Kershaw.
BESeuay Yuae Sv
Bulu in Chigogo.
Three specimens from the type-locality Dodoma and from
Kidenge. Seen at Igulwe. Occasionally taken in rat-traps,
which they probably visit for the ants which are attracted by the
bei, cio IB, L2G, SO, 20 war, 25 Iss AS. as AO sso,
ELEPHANTULUS rENAtTUS Kershaw.
Bulu in Kanyaturu.
Six specimens from Suna, Giwao’s, and Mtali’s, of which Gwao’s
was selected as type-locality. It was also seen at Pooma. They
have theiz runways in scattered thorn-bush, the soil is sandy or
reddish loam. At night they retire into burrows from which the
natives dig them in the early morning, as the Elephant Shrews do
not appear to stir until the sun has warmed the chilly morning
air. Suna specimens wero taken in rat-braps.
Largest ¢ 127.118. 35. 23 mm., and Q 140. 130. 32. 21 mm.
SoRICID &.
CROCIDURA MARTIENSSENI Neum.
A single female from Bagilo, Uluguru Mts., 13. vi.22, mea-
sured 150. 100. 20. 5 mm.
CrocIDURA FLAVESCENS Geoftr.
Moonyoonhe in Kikami, Kinyunga in Kisagara.
‘Two specimens from Kilosa were taken in a rat-trap, where they
had probably gone for insects eating the bread bait. ‘Three more
were found clinging to maize- and grass-stalks in a flooded area
during the rains. ¢. 98.60.23.12mm. Q. 95.58. 15. 11 mm.
A skull was found among the pellets of a Barn-Owl (Stria f.
maculata), one was recovered from the stomach of a One-streaked
Hawk (Kaupifaleo monogrammica), and another from a snake
(Ramphiophis oxyrhynchus).
Crocrpura irra Peters.
Kirukangia in Kiswahili, Nzunga in Chigogo, Jungs in Kin-
yaturu and Kirainba.
Six specimens from Dar es Salaam, Gwao’s, Pooma, and
Izikisia. Three of these were taken in rat-traps with meat bait.
The Dar es Salaam shrews were found beneath heaps of grass and
698 Mh. A. LOVERIDGE: NOWVES ON
garden refuse in open spaces in the middle of the town, where
two half- -erown specimens were met with on the same date
(21. vi. 23). Largest ¢ 90. 60. 13. 9 mm. (Izikisia), and Q 85.
50. 10. 5 mm. (Dar es Salaam).
The Swahili name for a shrew is connected with a widespread
belief common to almost all tribes, viz., that a shrew can follow a
path or road but cannot cross it ; if it attempts to do so it will
immediately fall dead. This legend has obviously been invented
to account for the not infr requent finding of dead shrews in the
road, as is also the case in England. The true cause of death
is that they have been pounced upon by genets and mungoose,
who, on discovering the pungent smell of their prey, abandon
them,
Crocipura niNbet Thos,
A female from Nairobi, measuring 80, 41. 13. 10 mim.’
CRrocriDURA BICOLOR ELGONIUS Osg.
A female from Eldoret, measuring 45. 31. 9. 7 mm.
RODENTIA.
ScrurniD4s”.
AVTHOSCcIURUS BYATTI Kershaw. |
Six examples from Bagilo, Uluguru Mts., of this species so
recently described from the Usambara Mts. further impresses me
with the correspondence between the mammal and bird faunas
of these widely separated mountain ranges.
Largest g 240. 160. 60. 30 nm.; largest 9 240. 190. 40.
30 mm.
HrLIOSCIURUS RUFOBRACHIATUS NYANS& Neu.
Vive specimens from Chantwara, of which the largest male
measured 245. 270. ? 2mm., and female 250. 235. 55. 21 mi.
(native measurements),
THELIOSCIURUS UNDULATUS UNDULAtUS 'l'rue.
A male and female only from Bungu, Usambara Mts.
PARAXERUS PALLIATUS SUAHELICUS Neum.
This Red Squirrel is known as Kimhulindi in Kikami, Chim-
weje in Chigogo, Sindi in Kinyaturu, Kitungu in Kinyiramba.
Nine specimens from Bogoti, Mkata River, Mbweui, Kipera,
and Madazini; also seen in dense maiombo serub at Ndogwe. It
is chiefly found in the tangled undergrowth on the banks of dry
water-courses, or in the big trees where patches of primary forest
still survive. A few were seen in thorn-trees.
EAST AVRICAN MAMMALS. 699
The largest male measured 215. 160. 40. 15 mm., and female
210. 240. 48.22 mm. Another female of practically the same
dimensions held a single diminutive fetus (Mkata, 24, viii, 21).
They spend a good deal of time running about among the dry
leaves beneath bushes picking up food, nibbling here and there,
and catching insects, ‘The stomach contents showed vegetable
matter like finely powdered meal, with traces of chlorophyll
probably representing leaves.
One individual had a mite and three lice in its fur, and a
parasitic worm (Streptopharagus sp.) in its stomach.
The bodies were eaten by my Wahehe and Wabena porters,
and I was told the Wakami also relish them.
PARAXERUS OCHRACEUS OCHRACEUS Huet.
This Olive Squirrel is called Kifrooma in Kikami, Kaderi in
Kisagara, Ndabi in Kinyaturu, Kipumbu in Kiramba.
Four skins from Mkindo and Mkata Rivers and Ulugu. It was
also seen by my collector at Mbonoa. At the second locality it
was found in association with P. p. swahelicus, which on two
occasions it Was seen pursuing. Another was observed jerking
its tail and chattering in unison with a bird.
Two males measured independently were 195. 160. 38.18 mm.,
a female 150. 160. 45.17 mm. ‘The bodies were eaten by the
Wahehe and Wabena porters.
Cricevipa.
Divopinitus Lureus Dollm.
Mbadya in Chigogo, Lebwa in Kinyaturu, Munsa in Kinyi-
ramba, Bewa in Kisukuma.
Twenty-seven specimens from Igulwe, Gwao’s, Mbulu’s, Pooma,
Madjengo’s, Simbo, Nyambita, and Sagayo. It appears to be
restricted to dry thorn-bush country, with perhaps a preference
for sandy soil. These specimens were obtained by offering a
reward to the local natives, who dig them out of their holes
with spears, and having killed them, insert the head beneath a
piece of string tied below the knee, and return to camp with the
pendent bodies and tails of the gerbils swinging to and fro. _
Largest 3 80. 90.15.10 mm., and 9 80.92.18.9 mm. The
tails and ears of many seemed to be scurfy and diseased.
TATERONA VICINA VICINA Peters:
Panya in Kinyaturu. This was stated to be specific, but this
is doubtful as it is the blanket name in Kiswahili for all rats.
Twelve Gerbils from Tindiga, Kimamba, Chanzuru, Uliya,
Mbala, Kisanga, Mbonoa, Pooma, and Mlewa’s. They were dug
out of their burrows in grass-country.
700 MR. A, LOVERIDGE: NOTES ON
Largest 3 172. 174. 33. 20 mm., and 2 160, 135. 34, ADT De
both fi om Mlewa’s.
One of this, or the next species, was Foead in the stomach of
a hawk (A. monogrammica).
TATERONA SWAYTHLINGI Kershaw.
Thirty specimens from Dakawa, Ilonga, Chanzuru, Kilosa,
Pwaga, and Ikikuyu. A large series from Kilosa were trapped in
ordinary rat-traps baited with bread. Some of the others were
dug from their burrows, which were unlined ; as many as five in-
dividuals were found in a single burrow. These holes are scat-
tered throughout the maiombo bush, usually in small groups or
colonies.
Largest ¢ 175. 180. 35. 25 mm., and 2 158. 165. 38. 22 mm.
Occasionally they may be seen in daylight; two young indi-
viduals (¢. 70. 80. 26. 14 mm.) were observed by the side of the
road at Kilosa on 16, vii. 21, and one being killed was found to
be swarming with parasites.
One was recovered from the stomach of a Genet (G'. swahelica).
TATERONA MUANS Matsch.
Nangala in Kinyiramba.
How papecimnen® from Dombolo and Sagayo, Largest g 150.
70. 22 mm., and 9 148. 180. 34, 21 mm.; both from the
first iene
TATERONA TABOR [Cershaw.
_A single female from Izikisia near Tahora. 160. 2? 3
20 mm.
RHIZOMYIDA.
TACHYORYCTES IBEANUS Thos.
A pair from Nairobi @. 180. 68. 30. 8 mm.
'TACHYORYCTES DAMON Thos,
A pair from Engare Mtoni near Mt. Meru.
Mvrips,
DENbDRoMUS OcHRoPUS Osg.
AU phir tromyNairobi. or (Osa LomlOlmimene ra iOn ove NG
10 min,
DENDRoMUuS PUMILIO Wagn.
A female was found at Kipera building its nest attached to
grass stems, nest six feet from ground. 9. 62. 85. 10.5 mm,
EAST AFRICAN MAMMALS. 701
SLEATOMYS LOVERIDGEL Thos.
Tulu in Kinyaturu, Ndolee in Kinyiramba.
Seven specimens of this Fat Mouse were taken at Pooma and
Mdjengo’s. ‘This species was hitherto only known from the type
collected by the writer at Lumbo in Portuguese Hast Africa; its
range is considerably extended by these new records. <. 85. 40.
Te sy aan, | SB, ai WES IL sem
STEATOMYS MUANS.E Kershaw.
Negoso in Kisukuima.
Three speciinens of this new mouse from Nyambita and Sagayo
have been dealt with by Mr. Kershaw. ¢. 76. 26. 13. 12 mm.
Q. 55. 35.13. 12 inm. They were found running about on and
beside the footpath between 7 and 10 p.m., and killed with
canes.
GRAMMOMYS SURDASTER SURDASTER (Thos. & Wrought.).
Five rats from Bagilo, Bogoti, and Tindiga. Largest g 115.
150. 30.30 mm. @. 110. 70. 35. 18 mm.
LATTUS RATLUS ALEXANDRINUS Geoffr.
Mbewa in Kikami, Ngule in Kisagara and Chigogo, Koho in
Kinyaturu, Mkinki in Kinyiramba, N goso in Kisukuma.
Twenty specimens of the Black Rat were collected at Nairobi,
Dar es Salaam, Llonga, Kilosa, Dodoma, Suna, Gwao’s, Pooma,
Mdjengo’s, and seen at Sagayo.
The largest male measured 175. 165. 35. 21 mm., and female
145. 162. 30. 20 mm.
Lt seems to be living alongside the local 2. ¢. microdon in per-
fect harmony, both species being not infrequently taken in traps
set in the same native hut. I found both species clinging to the
hack of a meat-safe this evening (Kilosa, 21. xii. 22).
On moving a large cupboard in the office a rat's hoard (it could
hardly be called a nest) was found. Init were three unopened and
alinost ungnawed letters, which had been in an outward dis-
patching tray on May 31st last. One of these communications
was of importance and its loss at that time gave rise to consider-
able correspondence (Kilosa, 31. xii. 22).
Another rat dragged a pair of cycling-stockings from a shat
down a hole; they were there a month and fortunately had never
been micced, or the blame would surely. have fallen on some
hapless native (Kilosa, 24. xii. 22).
On one oceasion a live rat was brought to me with hind feet
enormously swollen, and three cavities in them seemed to indicate
the evacuated holes of larval Caudylobia sp. (Kilosa, 27. vii. 22).
I saw a young vat in the middle of the morning running rather
blindly along by a wall,and on killing it found it to be swarming
702 MR. A. LOVERIDGE: NOTES ON
with fleas (Yenopsylla brasiliensis and cheopsis), of which I cap-
tured twenty-four. ‘The former flea was taken on another
specimen at Kilosa, and .Y. cheopsis from a rat’s nest in a pantry
at Nairobi. Most of the twenty-four were about the rat’s head
and fore parts, quite half-a-dozen around its eyes—perhaps this
accounted for the strange way it ran into and along the wall
(Kalosa, 31. xii. 21).
Black Rats were found in the stomach of a Serval (2. c. hinde?)
and of a Wild Cat (2*. 0. ugande), but their chief enemy at Kilosa
would appear to be the Barn-Owl, in whose nest many skulls were
found. Pufl-Adders also account for a few.
Ravrrus (4dviiomys) WALAMB.E PEDESTER Thos.
A single male from Kakindo on 15.1. 23.
Rartus (ANrnomys) CHRYSOPHILUS sINGIDH Kershaw.
The two types from Gwao’s, and one from Miljengo’s.
Rarrus (PRaomys) DELECToRUM Thos.
A female from Bagilo, on 17. vi. 22, measured 100. 120. 30.
30 mm,
Larrus (Mastromys) coucHA Micropon Peters.
Mbuku in Kikami, Ngobari in Kisagara, Mhanyalusangha in
Chigogo, Bebea in Kinyaturu, Lituri in Kinyamwezi, and Kungu
in Kinyiramba.
Kighty-five specimens from Bagilo; Wami, Mkindo, and Mkata
Rivers; Rudewa, [longa, Kimamba, Kilosa, Kipera, Rumruli,
Mpanirva, Ndogwe, Gwao’s, Mbulu’s, Pooma, Tabora, Izikisia,
Sanga, Sagayo, and Koma.
It is a garden rat, particularly numerous in the cultivated plots
of the natives; when food is scarce it takes to the grain stores in
the roofs of the native huts where, owing to their numbers, they
must do a great deal of damage.
The largest male of this fine series measured 135, 114. 22.
19 mm., and female 130, 112. 22.19 mm.; both from Monga.
Specimens from the thorn-bush steppe seem to average smaller
and ave paler, but can be matched by individuals picked out from
the Kilosa District series, ‘The grey pelage of the younger indi-
viduals tends to become a deep nut-brown as they grow older.
Three females trapped at Kilosa on 25,1.21 had 7, 8, and 11
embryos respectively, A female from [longa on 30. i1i.23 had
six foetuses. At Kimamba on 7, iv. 23 a litter of nine were found
in a burrow with their mother. Of these nine, one had a white
blaze on its forehead.
A flea (Yenopsylla brasiliensis), larvex of Caudylobia sp., and
nematode worms were taken from these rats.
These Rodents form the staple diet of many wild creatures,
EAST AFRICAN MAMMALS. 703
and were recovered from the stomachs of a Civet (C. c. orientalis),
Genets (G@. swahelica and G. d. newmnanni), Mungoose (11. flavi-
ventris), Pearly Owl (4. perlatwm), WRed-necked Falcon
(Pf. ruficollis), and Banded Harrier Hagle (C. fasciolatus). One
was disgorged by a House Snake (J. lineatus) when captured,
and many species of snakes in captivity fed upon them.
Ravrrus (Masromys) couct aA WILDEBRANDtTII Peters.
Two specimens from Bissel Camp near Longido in 1916.
Mus muscuuus Linn.
A single female of the European Mouse with two new-born
young atrived at Kilosa on 8.i. 21 in a box of groceries dispatched
from Dar es Salaam two days before. 92. 68. 76. 15. 11 mm.
LrcGADA BELLA BELLA Thos.
Kidangi in Kikami, Chimhanga in Chigogo.
Six Pigmy Mice from Dar es Salaam, Kilosa, Dombolo, Sagayo,
and Nyambita. One was caught in a tent at Itende, but sub-
sequently escaped. hey are not infrequently found in houses.
Largest $ 55. 45,12.9mm., 2 56, 42.13. 9 mm.
A nest of very simple construction was found beneath a heap
of rotting grass. The nest measured 40 mm. in diameter, and
was perhaps 30 mm. deep in the cup. The dead grass of which
it was constructed was not woven but loosely yet neatly inter-
laced to form a cosy cup, afew stalks formed a skeleton roof, but
it could not be considered properly domed. It held three blind
young ones of a dark fawn colour unlike their parent; one
opened its eyes on the 20th inst. (Dar es Salaam, 18. v_ 23).
What appeared to be one of these mice was found in the
stomach of a Whistling Hawk (A. sphenarus?%) at Wami River on
Wp ips Alls
CRICELOMYS GAMBIANUS oscoopI ITeller.
Nehesi in Kikami.
Seven specimens from Bagilo. There is also one in the Game
Dept. collections killed by Capt. Turnley at WKilosa on 14. ix. 22.
It must be very scarce at Kilosa, as my collectors never met with
it there.
The only male measured 410. 890. 80. 50 mm., largest female
380. 390. 72. 47 mm. They were trapped in native gardens,
where they are said to go to eat the maize.
LopHuroMys AQUILUS AQuiILus True.
Twenty-five Harsh-furred Mice from Jildoret, Mkindo River,
and Bagilo. Of twenty-three specimens from the last locality
the largest male measured 140. 90. 25. 15 mm., and female
150. 90. 20.10 mm. Dipterous larvee found in these rats have
been identified as Caudylobia rodhaini Gedoelst 4 Major Austen.
704 MR, A. LOVERIDGE: NOTES ON
Acomys srLoust de Wint.
Meru in Kikami.
A pair from Mkindo River on 5.ix.21, 9g. 97. 65.16.16 mm.
©. 85. 74.18.14 mm. Another was trapped near Mkata River,
but too damaged to be preserved. Like so many animals with
spinous fur or quills, the skin is very delicate and readily
tears.
PELOMYS FALLAX FALLAX Peters.
Five specimens from Bagilo, Tindiga, and Rumruli. The
largest male measured 180.160. 32, 20 mm., and female 140, 150.
30. 20 mm.
ARVICANTIIIS ABYSSINICUS RUBESCENS Wrought.
Four males from Kakindo and Kabare. o¢. 130. 82. 25.
10 mn.
ARVICANTHIS ABYSSINICUS MUANS& Matsch.
Thirteen specimens from Sanga, Lalago, Shanwa, and Sagayo
in Mwanza. They are very numerous, using their well-marked
) ’ oO
runs in daylight. Largest 140. 110. 26. 10 mm. 129. 105.
: yug ’
25. 16 mm.
ARVICANTHIS ABYSSINICUS NAIROL.® Allen,
‘en specimens of this well-known form were collected in the
garden at Nairobi, where they might be seen foraging near
the servants’ quarters whenever it was quict. |
ARVICANTHIS ABYSSINICUS VIRESCENS Heller.
Five specimens from Eldoret, of which the largest male
measured 160. 122. 31. 23 mm., and female 140. 113. 30. 18 mm.
ARVICANTHIS ABYSSINICUS NEUMANNI Matsch.
Fudi in Chigogo, Puku in Kinyaturu, Kongo in Kiramba.
Fifty-five from Kidenge, Mpanira-kwa-Sagoi, Ikikuyu, Dodoma,
Mbona, Suna, Gwao’s, Mbulu’s, Pooma, Singida, Mtali’s.
A distinct trace of a dorsal stripe in Suna specimens was
noticeable when they were alive.
Fleas (Yenopsylla brasiliensis) were found in the fur of
Dodoma and Mbulu rats.
These rats were recovered from the stomach of an Augur
Buzzard (B. augur) at Dodoma, and also from that of a Kite
(Ml. a. parasitus) at Mtali’s. Two were found in the stomach
of one large House Snake (2, lineatus) at Gwao’s.
EAST AFRICAN MAMMALS. 705
ARVICANTHIS TENEBROSUS Kershaw.
‘Twenty-nine specimens from Mtali’s, Izikisia, and Tabora.
Mr. Kershaw described this rat as a race of A. abyssinicus, but
this cannot be the case, as A. a. newmanni is found at Mtali's.
They are quite distinct. A. a. newmanni haunts roads, paths,
and open thorn-bush, whilst A. tenebrosus was taken entirely
from gardens where the natives wero cultivating mahoga or
sweet potatoes. :
Largest $ 160. 135. 34. 20 mm., 9 160. 135. 34. 20 mm.;
both from Mtali’s, where only three were collected. 'Tabora rats
were all smaller, the biggest being only 145. 110. 29. 18 mm.
Three White-faced Scops Owls (0. l. granti) had each one of
these rats in its stomach (Mtali’s, 19.x.21). Also in that of
an Augur Buzzard (B. augur) on the same day.
LEMNISCOMYS GRISELDA ROSALIA Thos.
A single male from Mkindo River on 5, ix. 21 measured 130.
138. 27. 1]4mm. A new-born young one, with eyes still un-
opened, was found in the roadway at Kilosa on 29. iii. 21.
Yng. 55. 32. 12. 3 mm.
LemNiscomys MACcULUS MAccuLUS Thos. & Wrought.
A-single male from Kabare on 1. ii. 23 measured 120. is On2 62
15 mm.
LEMNISCOMYS ALBOLINEATUS Osg.
Nyagalla in Chigogo, Kuse in Kinyaturu, Kohe in Kiramba.
Seven specimens from Ndogwe, Suna, Gwao’s, Pooma, Mtali’s,
Kadala, and Izikisia. It was also seen at Mbulu’s and Mlewa’s.
They run about on paths and in the roadway in the early
morning, and seek refuge in the manyara hedges. They are not
very numerous.
_ Largest 3 110. 118. 25. 14 mm.; 9 110. 115. 24. 14 mm.
RiuaABDOMYS PUMILIO DIMINUTUS Thos.
Two males from Eldoret and Rumruli. ¢. 100, 82. 20.13 mm.
O1rOMYS ANGONIENSIS ELASSODON Osg.
A pair of Swamp Rats were collected at Rumruli on 17. vii. 22.
OroMYs NYIKa CANESCENS Osg.
Two adults and one immature specimen from Nairobi, the
young one doubtfully referable to this species. ¢. 151, 91.
27. 18 mm.
706 Mn. A. LOVERIDGE: NOTES ON
PEDETIDS.
PEDETES SURDASTER subsp.
The Springhaas is called Kupa in Kiswahili,and Kamandegeri
in Kinyamwezi.
A pair were purchased at Tabora on 18. xi.21. od. 305. 320,
130.71 mm. @. 420. 420. 130,71 mm. The immature male is
in perfect pelage; the female has bare patches on her back,
possibly resulting from scratching sores. Burrows were common
at Mbonoa and Suna; at the latter place one Springhaas was
trapped, but a jackal carried it off. I followed the spoor, and
saw the jackal.
HystTricip4.
Tivsrrrx GALEATA Thos.
The Porcupine is known as Nungu in Kiswahili, Kibena,
Kikami, Kisagara, Chigogo, Kinyaturn, and Kinyiramba; Huna
in Misukuma.
Five specimens only collected, three others seen, and quills
found, at Rumruli, Kipera, Kilosa, Pooma, Shanwa, Sagayo.
The largest male measured 1160, 70. 75. 40 inmn., aud female
827. 65. 75,39 mm. Native measurements; both animals from
Sagnyo.
At Sagayo was a porcupine expert who, I heard, had dug out
four of these animals the week prior to my arrival. I therefore
engaged him for a week, to study his methods. He first sends
his small boys (two at least were his own sons, and wore only the
clothes they were born in) to find tracks and follow them to the
earth where the porcupine is lying-up, They then fetch their
father, who then sticks a horn in each entrance of the burrow.
hese horns (‘lopi, Thomson’s Gazelle, and Roan, also a large
wart-hog tusk) are filled with a pitch-like substance, into which
beads have been stuck before it hardened. ‘The result of this is
medicine said to prevent any animal that is in the burrow bolting
from the entrances, and at the same time robs them of their
ferocity so that they will not attack anyone entering the burrow;
this is very necessary, as at times jackals or a hyena are found in
the earths,
Having done this, one or more of the small naked youngsters
ave sent down the hole and follow the course of the burrow
underground, sometimes to a distance of thirty or forty feet.
I myself have seen a length of thirty feet. On locating the
animal at the terminus of the burrow’ they knock on the roof,
and the rest of the party, who are listening for this, reply, and
mark the spot. A shaft is then sunk, and so nicely judged in
the three instances where I was present, that it breaks into the
gallery within a foot of the porcupine who is at the terminus of
the burrow. While the shaft is being sunk, sometimes the boy
EASY AFRICAN MAMMALS, 707
remains in the gallery, singing and shouting to prevent the
animal breaking back; so it would appear that not too much
faith is placed upon the virtues of the horns.
The youngsters enjoy their work hugely, and have dark rings
chalked round their eyes ‘‘to enable them to see in the dark.”
They are simply smothered in red soil, for, indeed, when not
otherwise occupied, they play hide-and-seek in the galleries, and
pop up unexpectedly from the various entrances, or turn somer-
saults in the dust.
When the porcupine is uncovered he is speared to death, and
being very tenacious of life, does not succumb to wounds that
would kill any other animal. In the ease of those unearthed for
me, I shot them. One of these was shot through the head with a
22, and Isupposed was about dead. I therefore told the “ fundi ”
to bring it out of the burrow, and so demonstrate the truth of
his statement that he could handle porcupines with impunity.
He was very reluctant, but, pressed by the onlookers, took hold of
the animal, which promptly drove four quills into his palm. One
of the youngsters sucked the wound, whilst another fetched some
green leaves, which he chewed into a paste and spat upon the
wounds, with the object of preventing their hurting. Neverthe-
Jess, the ‘“fundi” stated next day that he had been unable to
sleep from the pain in his hand. I came to the conclusion that,
as a inedicine-man, he was a downright fraud. .
When given a coup de grace, a porcupine’s rattle is very active,
and every quill in its body vibrates and rattles.
The fundi told me that Jast month he found two females
occupying the same burrow, each having two young, the normal
number, The nipples are situated on the side just behind the
fore-limbs.
They all swarmed with a very large species of flea (Pario-
dontis riggenbachi), which transferred themselves to my skinners,
to the latter’s great discomfort.
BATHYERGID&.
HIELIOPHOBIUS EMINI Noack.
This Blesmol is known to the Wakami and Wasagara as Fuko.
1t would appear to be far from common, as only a single pair
(¢. 160. 12. 32. Omm., 2. 180. 14. 30. 0 mm.) were obtained
during the three years. Both were taken at Kipera on 5. v. 23.
The presence of these burrowing rodents is made manifest by the
heaps of fresh soil, as large as a mole-hill, which they will excavate
in the course of a single night. The female was obtained in a
mealie shamba by digging some six feet along, the blesmol being
found at the terminus, which was only some eighteen inches below
the surface. My collector said that it was but a poor method of
catching them, as one might dig for hours without overtaking
one, and if the excavated soil was fresh the way to capture them
708 Mn. A, LOVERIDGE: NOTES ON
was to uncover the hole and blow down the entrance, when the
blesmol would return to block it up. You must then prevent its
return by pushing a stick behind it before it can get back. He
stated that he caught the male in this way.
The stomachs contained finely-gnawed greenish and whitish
matter impossible of identification. The natives say they eat
ground-nuts (= monkey nuts).
I was told that only females are found, and that possibly males
do not come near the surface. This idea has probably arisen
from the fact that the testicles are internal, and the penis con-
cealed in a vulva-like opening from which it can be extruded.
The claws of one specimen were sheathed in caked soil, and the
silky fur of both had many blobs of the same attached very firmly
to the hairs, which do not appear to be so well adapted to the
burrowing habits of the blesmol as are those of an English mole.
The only parasites present were some very small mites running
over the white skin.
THRYONOMYIDS.
‘THRYONOMYS SWINDERIANUS VARIEGATUS Peters.
The Kiswahili name of Ndesi for the Cane- Rat is in general use
thronghout the country, and is the only one known to the
Wakami, Wasagara, and Wagogo.
Two specimens from Myombo; it was also seen at Miindo
River and Madazini, all localities in the Kilosa sub-district.
Of three specimens examined the only adult was a male (394.
143. 70. 39 mm.), which was shot when entering a down-pointing
gun-trap baited with a live goat! The largest female measured
340. 112. 60. 29 mm., and was speared by a native as it ran
across the road.
Both my specimens had fleas (Clenocephalus felis) present, and
the head of the female was burdened with numerous grey ticks
(Rhipicephalus simus Koch), One specimen was infested with
two species of lice (Scipio aulacodis Neum, and 7'ricodestes mangos
Stebbe).
LEPORIDS.
Lepus victor Thos.
Hares ave known as Sungura in Kiswahili, Buga in Nikami,
Kisungula in Kisagara, Sungula in Chigogo, and Siyi in
Kisukuma, ;
Five specimens collected at Mbulu’s, Mdjengo’s, Usshora, and
Sagayo. Hares are easier to see than procure, however, and
(spp. non det.) were met with at Kimamba, Kipera, Mpanira-
kwa-Sagoi, Ndogwe, Mtali’s, Lalago, and Mwadira.
The Sagayo male (450, 95. 100. 130 mm.) in the Game Dept.
collection was taken in a gin set in a burrow, which rather
EAST AFRICAN MAMMALS. 709
suggests that hares may make use of burrows at times when the
grass is scanty or has been burnt off. Another hare was caught
in a gin set for a leopard; the latter came along later and ate
the hare—the spoor was very distinct.
The female (480. 73. 100. 101 mm.) from Mdjengo’s held two
foetuses (24, x. 21). A very small leveret (155.30. 46. 35 mm.) was
brought me at Usshora on 6. xi. 21, having been caught by a dog ;
a larger one was brought me at Mbulu’s on 11.x. 21.
Two individuals from Mdjengo’s and Usshora were teeming
with fleas (Ctenocephalus felis and Echidnophaga larina), and the
former had numerous grey ticks (Rhipicephalus simus Koch)
as well.
Lrrus CAPENSIS cRAwsiAyr de Winton.
A single individual was collected at Eldoret on 9. xi. 21.
CARNIVORA.
CANID&..
THos ADUSrUS NorATUS Heller.
A fine skin but in poor condition was purchased from a native
in Mwanza town, the animal having been killed a few miles outside.
A live Side-striped Jackal from Liwale was given the writer
by Mr. C. B. Goss in January 1923, when it had already been in
captivity a couple of months. It was kept on a leash, with «
small native mongrel for a playfellow, they romped together a
great deal. It would allow itself to be carried for miles, and
liked being petted, but was a fearful coward and hated being
shut up at night, when it would howl mournfully as long as it
thought there was some likelihood of being released ; it preferred
sleeping with the native porters under a tarpaulin.
If quietly approached it would allow anyone to stroke it, but if
its coat was combed it snapped in the air several times as a
warning, and would doubtless bite if the combing was not stopped.
It has never bitten the writer, but came very near doing so on
several occasions, and bit a steward on board as well as several
natives, all of whom were probably to blame for tactlessness. Its
teeth are very sharp, and fortunately it only snaps and does not
hold on.
When given food it always menaced the person who brought
it, and is probably dangerous at such a time. It ate most scraps
from the table, meat, raw or cooked, but not tinned or preserved ;
it appeared to prefer fowls and fish to beef and mutton. For
five months it was fed on maize-porridge and soup bones. It
regularly overate the maize, disgorged the surplus and, after
half-an-hour’s rest, would eat it again. It was fond of rice, milk
puddings, cakes, biscuits, bread without crusts, and sweet fruits.
Its only drink was water, even soups being rejected.
Proc. Zoou. Soc.—1923, No. XLVII. AT
710 MR, A. LOVERIDGE: NOTES ON
Its attitude toward other animals, including dogs, was decidedly
friendly ; it endeavoured to make friends with the cat on board,
but she was suspicious. At Dar es Salaam dogs often gathered
round it in the hotel yard but appeared to fear it, as they would
not approach, but growled.and moved away. .
The animal was kept on a leather rein for five months till he
found he could gnaw itthrough, which he did ten times. 1 gave
up the contest and. allowed him to wander about the ship for a
week till, finding his way into the crew’s quarters, he dragged
some clothes from a bunk along the deck, and seizing on a leather
belt carried it off and gnawed it in half. He was then fastened
with very light wire-rope, but after forty-eight hours cut through
that also.
THOS MESOMELAS MCMILLANI Heller.
Mbweha in Kiswahili. It has no name in Kikami, but is
called Kewe in Kisagara, and Nhyewe in Chigogo.
Hight specimens, of which three only were taken by the
writer’s collectors, from Mkata River, Gulwe, Suna, Gwao’s,
Mlewa’s, Sagayo.
The largest male measured 740. 290. 190. 100 mm., and female
680. 295. 150. 102 mm.
Both of these came from Sagayo, and their stomachs were full.
of hyrax fur and meat, whilst one had some larger carrion also.
The Mlewa specimen had grasshoppers and several large black
scorpions (Pandinus cavimanus Poc.) in its stomach. I+ would
be interesting to watch a jackal’s method of attacking a scorpion.
The Mkata specimen’s stomach was well filled with the fruit of
a tree called by the natives ‘‘ Mkongo.” We came to several of
the trees, and amongst the abundant fruit lying on the ground
was a good deal of spoor of jackal and civet.
‘Twelve fleas (C7. felis) were taken from one jackal.
Lycaon pictus subsp.
Mbwizi in Kikami and Kisagara, Iminzi in Chigogo.
The Hunting Dog was not collected nor even seen by me,
though it is. only too common in the Kilosa District, and
was often heard. Whilst I was away from home on one
occasion a pack of dogs drove a bushbuck right through the lower
part of the house at 6a.m. Another time they harried a bush-
bueic past the front of the house at 5 p.m., and Mr. D. W.
Bisshopp shot one of the dogs from the verandah, but it was not
found for three days afterwards, when it was too far gone to
preserve. At Myombo, some nine miles south of NKilosa, they
pulled down a bushbuek almost on the doorsteps of a settler’s
house in the middle of the morning, It is interesting to note
that bushbuck appear to run to a human habitation when pur-
sued by their remorseless foes.
EAST AFRICAN MAMMALS, 711
Orocyon vireatus Miller.
Bee in Kinyaturu, Bele in Kiramba.
Five males (two being cubs) collected at Mdjengo’s, Mtali’s,
and Sagayo. A score were seen at Izikisia in the moonlight.
The largest male measured 556. 275. 120: 95 mm. One cub
died a month after I got it, perhaps it would be six weeks old
when it measured 290, 115. 90.70 mm. Its twin brother was
measured whilst alive as accurately as possible at five-and-a-half
months old: 470% 250% 120? 942%; he then weighed 64 Ibs.
The two lively little cubs, with eyes open and just able to walk,
were brought me at Mtali’s on 20.x. 21, their father having been
killed by a native dog. At this stage their fur was very woolly
and an almost uniform smoky-grey.
At five and a half months the colour was as follows :—Grizzled
buff, a thick woolly underfur of buff, almost concealed by long
whitish hairs tipped with black. Feet as far up as the knees,
black ; the whole leg is darker than the body. ‘lip of tail black,
extending in a wedge-shape on dorsal aspect almost to root of
tail. Muzzle black, extending on the forehead to between the
eyes, on the cheeks to beyond the eyes. ‘Tip of the ears and for
some distance downwards, black. ‘The ears are fringed with long
all-white hairs, a good deal of buff about the base of tte ears.
When young I gave them a great deal of meat; one of them
ate a green locustid at three <sualee old. Another time IXip, as
the survivor was called, pounced upon a Striped Hawk-Moth,
chewed it well, dropped it, and then ate it up. He is passionately
fond of butter and honey, will eat porridge, bread, eggs, and
drink milk or tea with relish. Bananas and paupau are also
eaten readily, but he will not touch mangoes or pineapple.
He spends a good deal of his time “turning over stones in
search of millipedes, which are gobbled up greedily. I offered:
him a black-and-yellow polydesmid which he pawed over for
some three minutes, but would not eat it. Im strange contrast
was his attitude to a Lesser Stink Ant (Paliothyreus tarsatus),
which came out of its hole and waved its antenne right under
KGip’s nose. He looked at it, then seized and crunched it up
despite the smell, w hich was noticeable to me standing a couple
of feet away.
A very big baboon came up to the kitchen one afternoon, and
aroused [Kip’s curiosity, the fox then being five months old. He
followed the baboon with his head down, “and when the baboon,
catching sight of me, cantered away, Kip flew after it for an
hundred yards with only about six feet between. A month later
haboons were in the vicinity of the house daily, and on. several
occasions I saw him playing with them. He will lie crouched
upon the ground, and a big dog-baboon will come walking slowly
towards him till within two yards, when I<ip will spring up and
fly straight at it, the baboon cantering easily away, looking over
its shoulder at alhe small animal which it could so easily kill.
Many other baboons will be close by during this play.
47*
=]
MR. A. LOVERIDGEH: NOTES ON
Kip’s attitude to baboons is very strange seeing that he is
naturally a timid animal, very nervous of approach, especially
when feeding. Like many animals he developed an antipathy to
natives, which was difficult to understand as they were very fond
of him, but at four months old he took to menacing them,
snarling and growling and looking very wicked. Even when
furiously annoyed at being tied up for the night his teeth scarcely
break the skin.
He was six-and-a-half months old when he met his first dog—
a quiet little mongrel Dacnshund which accompanied a visitor,
Kip advanced growling and bristling towards the stranger, and
exhibited for the first time a large rufous patch on the base of
his tail (dorsally).
MUSTELID2.
MELLIVoRA carENsiIs Schreb.
Kibakusi in Kikami; Nyergeri m Kisagara, Kisukuma, and
Kiswahili.
The Honey Badger or Ratel is not often met with in East
Africa, though probably fairly common and certainly very widely
distributed. Four specimens were obtamed from Makindu, Wami
River, Kilosa, and Sagayo, those from the last-mentionaed locality
being skins without skulls, purchased froma native. A young
male measured 480. 160.105. 30 min., an adult 770. 198. 130.
22? mm., and a female 670. 200. 112. 30 mm.
‘fhe immature inale was one of two individuals encountered on
the plains about 7 a.m. one morning, and shot by my native
collector. It appears quite common for them to hunt in pairs,
which need not necessarily be of opposite sexes, for two adult
females were hunting in company on one occasion.
During July 1921, a native complained that an animal had dug
into his strongly-built mud and wattle chicken-house, and taken
four fowls. I set a gun-trap (°22 B.S.A.) over the entrance it
had made, and the following night the gun went off, but so did the
animal without leaving avy trace of being hit. A few nights
later it returned and took three more fowls in one night, effecting
an entrance by digging in a different spot. In neither instance
was a trace of the fowls left, and presumably they were carried
off whole.
A lion trap was set inside the yard at the spot where it had
broken through the fence, and the following morning my boy
informed me that a ratel was in the trap. The poor beast had
dragged the heavy trap twelve feet and torn a hole in the fence,
but couldn’t get the trap through. It had chewed a ground -sheet
to rags as well as other things within reach, including its own
foot, whose claws I recovered from its stomach after havinig shot
it. I have known this happen in the case of another ratel, and
have little doubt that in the fierceness of their rage at being
caught they lose their sense of pain to some extent, else how
could they mutilate themselves 2
EASL AFRICAN MAMMALS. 113
The stomach of this individual contained fowl feet, beaks, eyes,
etc., and the wing of a bird. The other specimen had a little
grass, probably seized after capture. A screw-worm and a tape-
worm were found in the abdominal cavity.
Despite the nauseous stench of the creature, Wazigoor and
Wabeni porters asked for the bodies of two ratels and ate them!
IcTONYX STRIATUS ALBESCENS Heller.
A single female Zorilla (250, 152. 49, 23 mm.) was purchased
from a native, who had killed it in his fowl-house at Usshora,
30.x.21. Though very young it had minute fetuses in utero.
AONYX CAPENSIS HELIOS Heller.
Fisi maji, its Kiswahili name, is commonly used for the Otter
by both Wakami and Wasagara.
A single female (762. 485. 132. 25 mm.) Otter was trap-shot
with a *22 rifle whilst under water at 'Tindiga, 1.ix.22. here
were fish in its stomach and a new Ascarid (Clwoascaris spinicollis
Baylis), and I was somewhat surprised to find a flea (Cteno-
cephalus felis) in the fur of such an amphibious creature. Ticks
were also present but not preserved.
VIVERRIDS.
CIVETYICTIS CIVETTA ORIENTALIS Matschie.
The Civet is known as Fungo in Kiswahili, Kikami, and
Kisagara.
Twelve specimens from Wami River, Mkata River, Kimamba,
Kilosa, and Sagayo. ;
The largest male measured 900. 480. 140. 60mm. (Sagayo), and
female 940. 445. 140. 60 mm. (Wami R.).
The only note on breeding is the record of two litters brought
me on 21. ii. 21 and 29.xi.21 from Kilosa. ‘he first litter con-
tained three and the latter two. The young are almost black.
The first hatch were very fierce, spitting and biting when
approached ; after a month two of them showed signs of taming,
and would come to me at meal times, standing up on their hind legs
or clambering tomyknee. They never liked being handled. Civets
are very dirty feeders, and these little beasts invariably put their
feet into their milk or held jam down with their paws, getting
into a nice mess, They were very fond of mangoes.
Stomachs of wild specimens held the following :—(i.) Bodies of
hirds and rats skinned by my collectors the previous day, chicken’s
leg and mango skins from the kitchen. "This animal was shot at
Sa.m.in bright moonlight from the verandah. (ii.) Flesh of a
baboon, which I had put out asa bait, some other big pieces of
flesh, skin bearing fur of what might be the gerbil (Tateronasp.),
a locust, and a calculus of hair. (iii.) 2. c. microdon, eggs and
chicks of a francolin, fruit of mkongo tree.
714 Mh, A. LOVERIDGE: NOTES ON
On offering the body of a skinned genet to my young civets
they approached it gingerly, and then the male flung himself,
throat foremost, on the carcass, rubbing his throat, chest, and
shoulders in the entrails in an ecstacy of delight; he was shortly
joined by a female, and the two of them pushed the carcass all
about the floor in their efforts to rub their throats upon it.
After a month in the house I turned these three civets loose in
an empty room of a roofless German house, and as I was uneble
to spend much time with them they grew very fierce, though regu-
larly and well fed. After four and a half months, therefore, 1
chloroformed two of the worst and let the other go. The two
chloroformed, which may be assumed to be five months old,
measured: Male 520 310. 97. 46 mm., weight 53 Ibs.. Female
580. 350. 110. 50 mm., weight over 7 lbs. (the seale only weighed
to seven). This male was always rather backward.
The liberated male had his food placed in the room and returned
for it regularly for a fortnight, after which I went away. I fre-
quently met 16 on the path in the moonlight, making its way to
the house, and it did not flee but generally withdrew in a shadow
and crouched there until I passed,
Few animals are so rich in parasites : : one specimen alone had
nine hippoboscid flies (//. capensis v. Olf.), fleas (Ctenocephalus
felis), ticks (Rhipicephalus simus Woch), and nematodes (Milaria
sp., all females, not /. martis Gmel.) were found in its stomach.
Ticks and this species of flea were almost invariably present.
‘Another flea (Mchidnophaga larina) aud tick (Llemaphysalis leach)
were found on a Sagayo civet,
Two instances of civets caught in gin-traps biting off their feet
came to my notice, indeed I found what was probably the remains
of one of them in a dry watercourse. ‘The skull showed that the
animal was very aged, all the molars on the right rami of the
mandible were missing and the bone healed over, though it
appeared to have had a bad abscess at one time.
The bodies of civets killed on ono safari were. eaten by my
Wazigoor and Wabeni porters.
GENETTA DONGALANA NEUMANNI Matsch.
Nghanu in Chigogo, Ndele in Kinyaturu, Nilele in Kiramba,
Kanididi in Kinyamwezi.
Nine specimens of Neumann’s Genet were obtained at Kidenge,
Itumba (skin seen), Ndogwe, Suna, Singida, Mdjengo’s, Mtali’s,
and Izikisia, ‘Three of these were trapped with a down-pointing
22 rifle while attempting to get at a fowl used as bait.
At Singida an adult male (490. 460. 90. 50 mm.) was brought
to me alive with a stri ing round its neck; though freshly causht
it allowed itself to be stroked and was very amenable, possibly ib
was dazed by daylight. When annoyed if raised the fine black
dorsnl mane on end and bushed out its,ringed tail.
At Tvikisia on 15, xi. 21 two kittens were found in a hole in a
EAST AFRICAN MAMMALS. 715
tree in open miombo bush country. The hole was some ten feet
from the ground. J rather hesitate in referring these kittens to
this race as no adults were collected in the locality.
A. rat (2. c. microdon) was. found in the stomach of the Ndogwe
genet. hese genets cause a great deal of mischief among native
fowls, which they appear to kill for killing’s sake when they can
get into a fowl-house.
GENE?TLTA SUAHELICA Matsch.
Kanu in Kiswahili, Kikami, and. Kisagara, Tondolega in
Wabhelie.
Thirteen specimens from Mkata River, Rudewa, Kimamba,
Kilosa, Mbonoa, and Mdjengo’s.
The largest ale measured 500. 440. 89. Ap mm., and best
female 525. 435. 87. 48 mm.
Iwo very young kittens (¢d d. 190. 165. 40. 25 mm.) were
brought to me at Kilosa on 7.v. 21, but only lived till 20. v, 21.
Another, only a few days old, on 28. iii. 21, throve for a month,
but during my absence from home was allowed into a room where
there was a freshly-cured leopard skin on which it went to sleep.
Jt afterwards apparently licked the skin, for I found traces of
arsenic in its stomach, and it died with all the symptoms of
arsenical poisoning on 27. iv. 21.
It was suffering from constipation when preeht to me and on
the second day extruded its bowels, which I washed with warm
water and permanganate, lubricated with vaseline, and returned.
I gave it doses of salts at 2 and 6 p.m., and it obviously recovered
as it lived for a month after and was always very full of life.
I first fed it on sweetened milk and jam; after nine days I gave
up feeding it, and it lapped milk and it attempted scraps of meat.
It very soon gave up spitting when handled, and showed no fear
-of a mungoose or three young civets, ever so much larger than
itself. It spent a great deal of time in sleeping, varied by
climbing, at which it was anadept. Itclambered up the mosquito
gauze to the top of a door and then called to be helped down ;
backs of chairs were a very favourite goal, which when reached
it would rest upon.
When sitting up in trees at night it is a common occurrence
to hear genets hunting through the grass, pouncing here and
there, or clambering into a bush, from whence birds would go off
with a twitter or a whirr according to kind. I waited for one
such for half-an-hour before it appeared in the road. After one
or two short runs it then vanished, but presently reappeared, and
seeing the goat beneath my tree dashed towards it, but on the far
side, where it halted for a second to look at the goat, and I shot it
dead.
In its stomach were grasshoppers, a large black field cricket,
and beetle elytra, together with not a littie green stuff, mostly
grass-blades as far as one could determine. A second individual
716 Mh. A. LOVERIDGE; NOTES ON
had eaten a gerbil (7'. swaythling?) and a rat (22. c. nicrodon).
In the stomach of another was the tail of a rat, the unidentifiable
foot of a small bird, the body and limbs of a frog (Rana sp.), and
many kinds of insects much chewed. Yet another held a single
rat (J. c. microdon).
‘Three species of flea were taken from three individuals (viz.,
Kchidnopsis gallinaceus, Chimeropsylla potis, and Clenocephalus
felis). Ticks (Hamaphysalis leachi) were found on three genets,
and nematodes (Ascaris sp., females) were also taken,
GENETTA STUHLMANNI SI!UHLMANNI Matsch.
Nilili in Kisukuma,
A single immature male (240. 230. 80. 33 mm.) twelve feet up
ina bush on the banks of the Simiyu River at Sagayo in mid-
morning. Fleas (Ctenocephalus felis) and ticks (Hemaphysalis
leachi var.) in its fur, and ascarids came wriggling from its mouth
just after death. These worms were all females and not refer-
able to Belascaris, but may indicate a new species (Spaul).
Adult genets were seen in the neighbourhood of kopjes on two
occasions.
Herpestes (CALOGALE) FLAVIVENTRIS Matsch.
On three occasions examples of this Mungoose were seen
hunting in daylight on kopjes at Sagayo; they reminded one
strongly of ferrets in the way they explored crevices and re-
appeared in unexpected openings.
3. 350. 280. 52.20mm. 9. 332.260.25.15mm. The male
had eaten a rat (2. c. microdon}; the female, which was gin-
trapped among the rocks, is in the Game Dept. collection.
HERpestes (CALOGALE) GrANtI Gray.
A female from Usshora had been killed by a native in his fowl-
house. I saw one of these mungoose in association with J. g.
lademanni Mats. in a ravine at Mbulu’s, another day a pair were
hunting together over a rocky kopje. WhatI took to be a male
had a black tail-tuft, not ochraceous as in the female.
@. 350. 290. 60. 20mm. It had three fleas (Ctenocephalus
felis) in its fur.
THrrrestes (CALOGALE) MELANURUS RUFESCENS Lorenz.
A male killed at Zanzibar on 12. viii. 16 was presented to the
writer by Dr. Aders, F.Z.S.
Herrestes (CALOGALE) GRACILIS LADEMANNI Matsch.
Known as Kiniboo in Kinyaturu, and Chonja (?) in Kisukuma.
_A single male (350, 280. 60. 30 mm.) from Mbulu’s was driven
by my collector out of a ravine between kopjes in thorn-bush
steppe. ‘I'wo were seen on different occasions at Mtali’s and yet
EAST AFRICAN MAMMALS. 717
another at Mkalama, but they are very active and vanish from
sight amongst the boulders of the kopjes which they frequent.
The male’s stomach contained a lizard (V. emini), a skink
(Mabwia sp.), a chameleon (C. _parvilobus), and grasshoppers.
‘There were also some indeterminable nematode parasites. Six
fleas from the fur were identified as Ctenocephalus felis.
Henresres (CALOGALE) GRACILIs IBEZ Wroughton. |
A female was collected at Ngong Forest, Nairobi, 2. x. 20.
TIERPESTES ICHNEUMON FUNESTUS (Osg.).
Four from Kilosa. Two were purchased from natives who had
killed them in their fowl-houses. A skin was also seen at
Mkalama.
One male (532, 532. 100. 37 mm.) was very lean, but its fur
was in yood condition, some of the black hairs in the terminal
tail-tuft were six inches long. ‘There was rat’s fur in its
stomach.
There were many parasites on this one individual. Lice
(Trichodectes rammet Stebbe) swarmed in thousands. Of nine-
teen fleas captured two were Hchidnopsis gallinaceus and the
remainder Ctenocephalus felis. Nematodes from the liver and
intestines were Physaloptera sp. indet.
ATILAX PALUDINOSUS RUBESCENS Holl.
Karasa in Kikami.
Six specimens of the Water-Mungoose from Bagilo, Tindiga,
and Kilosa.
The largest male measured 530. 350. 105. 35 mm. Largest
female 530. 330. 100. 31 mm.
The HO SEBS taken at Tindiga, held two footuses measuring 104.
52. 21. 5 mm., and both together weighed 3 oz. (27. viii. 21).
One was shot at 8 p.m. in bright moonlight as it approached a
dead monkey I had left on the path, a second appeared shortly
afterwards.
One specimen swarmed with lice (Z'richodectes acutirostris
Stebbe, 7’. mangos Stebbe, 7. ?ramme: Stebbe), a tick (hipi-
cephalus simus Koch), and had filarid worms beneath the skin.
In the stomach of another was an ascarid (Clwoascaris spinicollis,
gen. et sp. n. Baylis).
IcHNEUMIA ALBICAUDA IBEANA Thos.
Kanhanga in Kikami, Mbaku in Kisagara.
Six specimens of the White-tailed Mungoose from Tindiga,
Hlonga, Kalosa, and Nairobi.
_ Yhe largest female measured 600. 455. 132.38 mm. An im-
mature female (440. 350. 100. 38 mm.) was disturbed in the
burning of rubbish and run down by a native, from whom J
718 MR. A. LOVERIDGE: NOTES ON
purchased it. It is very dark in colour and the white hairs
of the tail are concealed by longer blackish ones, so that: it
presents a very different appearance from the typical white-tailed
individuals.
A female killed at Tindiga on 10. 11. 2) had two feetuses, those
on 18 and 24, viii, 21 none,
On 1.1.23 I was given a young male by Capt. Turnley, at which
time its coat was grey and very woolly, but long black hairs were
showing thickly on the nape and fore-par'ts. ’
It liked being picked up and petted and would lie on its back
in one’s arms most contentedly. When very pleased it licked one
after the manner of a cat. -One’s nose seemed to be a cause of
offence, for if lying on a sofa or in any position where it could be
reached, the mungoose would attack it fiercely. With this
exception, if never ‘attempted to bite except in play.
It enjoyed. a game—usually started them in fact. Ono
favourite pastime—after having located a safe retreat under a
cupboard or other piece of furniture—was to steal forth with the
greatest caution, then, feigning alarm, to scuttle and scramble back
with much noise. After 1 repeating this performance several times
in the hope that you will chase it back, which it considers great
sport, and failing to attract your attention, it will appr ‘ach * very
silently and bite your heel or give you a, pat with its claws and
then tear madly back to its retreat. If you still refuse to come
and play, it attacks your shoe- laces and ‘rags’ them with great
abandon.
It showed great alarm of sudden noises, and the hairs of its
white tail immediately stand on end as does its back. fur to some
extent; it spits most explosively. It has a wonderful variety of
sounds at its command, one of contentment is almost indistin-
guishable ‘from that made by the young Blue Monkey, and is
something like ‘urrr.’
After I had ‘had it two months I went on safari, and almost the
first night it- was: turned loose in the tent it was scared by the
sudden centr ance of a, native and ran out into the bush, never to
return.
It showed ent foudnbee for the pupe of moths and wasps, and
broke open the mud nests of the latter with its strong claws, but
never in my experience touched the grubs. It often rejected the
bread in its bread and milk, but liked sweetened condensed mill
and eggs. ‘Chicken bones ‘would be picked clean and greatly
appreciated. Rice, masbed potatoes, and porridge were taken
readily.
The stomachs of the wild adults collected contained the follow-
ing :—(i.) Grass, leaves, termites. This individual was trapped
whilst attempting to enter a fowl-house. (ii.) Remains of fowl,
crickets, paupau skin, (iii.) Grasshoppers. (iv.) Termites.
Fleas (Ctenocephalus felis) and ticks were found in the fur,
and cestodes (Spharganum sp.) between skin and flesh. Two
individuals had numerous screw-worms in viscera.
EASY AFRICAN MAMMALS. 719
HELOGALE UNDULATA UNDULATA Peters.
The Lesser Mungoose is known as Kimgalla in Kisagara,
Muloli in Chigogo, Minyirsira in Kinyaturu.
Five specimens collected at Kipera, Mpanira-kwa-Sagoi, and
Mbulu’s. It was also clearly seen near Mkata River, Pooma.
The species is‘ diurnal and very definitely associated in my
mind with the termite hills, to which the little hunting parties
scamper when disturbed. Before disappearing down the holes
they usually sit up, with fore-paws drooping on their stomachs,
and take one more look at the cause of their fright. One was
shot on a sloping tree-trunk.
Largest male measured 240. 175. 45. 17 mm., and female 230.
170. 45. 19 mim.
HELOGALE vicrorina Thos.
Kamsio in Kinyamwezi, Lonzi in Kiramba, and Kijinolo in
Kisukuma. '
Eleven specimens from Mdjengo’s, Sanga, and Sagayo. It was
also seen at Mlewa’s, Mtali’s, Mkalama, Usurwe, and Shanwa.
Largest male 260. 160. 43. 20 mm., female 245. 157.45. 18 mm.
Young of latter in nest 80. 40. 15. 4 mm. cats
One individual held two fcetuses only (Sanga. 16.x. 22).
A slight nest of grass (native collectov’s statement) was found in
excavating a termite heap, where the female was found with four
almost naked young. She and her family were transferred toa
box, but though eating heartily herself, she refused to feed them
so that they died (Sagayo, 1, xi. 22).
Stomachs contained finely masticated insect remains, which
appeared to be chiefly those of grasshoppers.
Muncos munco cononus (Heller).
The Banded Mungoose is known as Ngutchiro in [Kiswahilh,
5 7s "1° . of .
but the Wa-swahili use this name for all other species of mungoose
which they may encounter and many of whichare quite unknown
at the coast. The Wasagara appear to have no other name than
Negutchiro for it. Tukwain Kikami, Nghalasanga in Chigogo,
Kala in Kinyaturu, Kala in Kiramba, Ikala in Kinyamwezi,
Nhala in Kisukuma. pies
Five specimens collected at Dodoma, Pooma, Mdjengo’s, and
Usshora. It is, however, a very widely distributed species and
was also seen at Mlewa’s, Mtali’s, Usurwe, Mkalama, Shanwa,
Sagayo, and all along the road from Ulugu to Tabora. Curiously
enough, [ have never seen it at Kilosa during the two years spent
there, though a company of mungoose crossed a path one after-
noon, but.at too great a distance for me to be sure they were not.
Atilax (which usually goes singly or in pairs), or some other large
species. hc
720 MR, A, LOVERIDGE: NOTES ON
Largest male measured 390, 240. 78. 25 mm., and female 395,
280. 89 2, 20 mm.
« At Pooma I saw ten of these mungoose sitting Bee on a
termite hill, intently watching a flock of guinea-fowl which were
feeding towards them across the open, the nearest being only
twenty feet away. I was able to approach and shoob two
mungoose sitting close together, but one fell down the hole. The
other I found was a very light sandy, or straw colour, excellently
adapted to life in the thorn-bush steppe. On showing it to my
collector, he said that this variety was well known and that
probably all in the thorn-bush country would be of a similar shade.
Unfortunately for this theory, its dead companion was dug out
of the hole and was the typical nut-brown shade, as were all the
others subsequently seen at close quarters in the thorn-bush
steppe.
Just on the outskirts of Tabora, about twenty feet from the
mainroad along which a noisy stream of natives were passing,
eight mungoose were seen sunning themselves in all mannev of
indolent attitudes about 7 a.m. They took not the slightest
notice of my porters, who had dropped their loads, and were
moving about within fifty feet of the bold little beasts.
An amphisbenid (G@. modestus) and two skinks (Lygosoma sp.)
were found in the stomach of an Usshora specimen; a grass-
hopper, large carabid, and beetle larve in another.
Fleas (£ chidaophaga gallinaceus) were taken in the fur of one,
whilst screw-worms were found in the viscera of another. The
Banded Mungoose is killed by the Wanyaturu for food.
PROTELEID A.
PROTELES CRISTATUS TERMES Teller.
I have not been fortunate enough to obtain a specimen of the
Aard-wolf, but the creature being so rare in ‘Tanganyika Terri-
tory it seems worth recording every occurrence. I have only seen
two skins, one obtained near Itumbi in Manyoni District, the
other, which is in the Game Dept. Museum, was obtained by
Mrs, Billinge at Kiganga on the Ruaha, 31, vill. 22.
It was caught in a lion gin set on a path and was a full-grown
male, measuring 30 inches from nose-tip to base of tail, the latter
being another 10 inches according to Mrs. Billinge’s measure-
ments. She estimated the weight as about 20 lbs.
Ty anip2.
Hy#NA HY2NA SCHILLINGSI Matsch.
Two examples of Schillings’ Striped Hyena were obtained at
Gulwe and Mtali’s. Both were males, of which the larger
measured 1190. 290, 200. 158 mm.
In the stomach of the younger specimen were the remains of a
EASY AFRICAN MAMMALS. 721
monkey skinned in camp that day, a great quantity of mealies,
and some wild yellow fruit which might have been in the stomach
of the monkey. ‘There were also a lot of vertebrae of a good-sized
python and the skin of an agama lizard.
A Hippoboseid fly (/7. capensis v. Olf.) was in its coat and also
two species of ticks.
Crocura CROCUTA GERMINANS Matsch.
Three specimens of the Eastern Spotted Hyena were coilected,
others were also examined from the following localities :—Kilosa,
Mtali’s, Izikisia, Simbo, Shandwa, and Mwadira. —
The Kiswahili name of Fisi seems to be universally employed,
in Chigogo it is altered to Mvisi, and in Kisungwa it is called
Fifi. ; .
The largest male measured 1350. 250. 215. 110 mm., the
largest female 1245. 241. 215. 101 mm.
The latter held a single feetus very near the birth, weighing
34 Ibs. and measuring 330. 70. 60. 29 mm.
One which I shot at a waterhole at 1.30 a.m. had come to drink
earlier in the evening, and a movement of mine in the hide-up
had caused it to bolt. It returned with three zebra at its heels,
and tried to get past a flanking zebra without coming too’ near
my hide-up, which it was quite aware was tenanted. It was in
abject terror, trying to crouch past with stomach close to the
ground, when I shot it at fifty yards.
The contents of its stomach and others were as foliows :—
(i.) Goat’s skin, hoof and ox bones, broken fragments of a gourd
picked up near a native hut presumably. (ii.) Goat or small
buck’s hoof, giraffe bones and strips of giraffe skin, pared off by
a sandal-maker, fowl’s feet, bits of gourd. (iii.) Remains of goat
and ox, wild pig, duiker, rat’s foot, fowl’s feet, and two hairy
ealeull.
Three fleas from the last-named specimen proved to be Cieno-
cephalus felis and Echidnophaga larina. ippoboscid flies (//.
capensis v. Olf.) were found on two specimens, also ticks (hipi-_
cephalus simus Koch).
FRELID.
ACINONYX JUBATUS RAINEYI Heller.
Duma in Kikami and Chigogo. Pocho(?) in Kisagara (Pocho
appears to be applied to the Hunting Dogand there may be some
confusion). ‘The Cheetah is not a common animal in Tanganyika
Territory as it is in Kenya so far as my experience goes. The
natives are unfamiliar with it and few seem to know its name,
usually confusing it with either the Leopard or the Serval!
The only local skin I have seen was of quite a large animal,
which was brought for sale by two natives, who said they had
found it dead near Kideti (viii. 22); having been killed by a snake.
722 MR. A, LOVERIDGE: NOTES ON -
which, after an examination of the skin, I think very probable;
most likely a puff-adder.
The Wanyimwezi, Wanyiramba, and Warangi have a saying
that the witch doctors give a potion to the cheetah which endues
it with sufficient courage and fierceness, and causes it to rush off
into the bush, where it lies in wait for children and even adults
whom the witch-doctor wishes to be rid of: the creature pounces
upon them, lacerates them, and laps the blood, but does not eat
thie flesh, and presently leaves the body.
Friis LEO MASSAIcA Neumn.
One specimen collected at Kilosa. Many others seen; the dis-
tribution is so widespread in ‘langanyika Territory that it would
serve no useful purpose to give localities.
Manes as a guide td subspecies? Some lions broke into a hut
300 yards from Kilosa Boma on 4.11.22 and killed four goats:
traps baited with dead goats were set the following night and one
lion was killed; the other, wounded in the nose, is the “ specimen”
referred to above. ‘his individual had a fine dark mane, and
was a full-grown male in its prime with splendid undamaged
teeth. ‘The other was a very old male, maneless, with worn and
broken teeth, very ill-nourished, and with the spotting usually
associated with youth or females, very noticeable.
Do lions climb trees? At Kipera on 26. xii. 22, I was coming
through a lot of rank sedge ‘almost shonlder-high when I saw a
fine waterbuck on a slight eminence some two hundred yards
away. I fired, and the bullet struck a tree close behind him and
just below his head.* I fired again, and ‘he went down with a
roar. At the same moment a lioness sprang from a tree and
bounded away in full view toa thicket. My boys, who were some
little distance behind me, said that at the first shot they saw the
lioness put her head out of the foliage and peer about, and they
supposed I was firing at her, but with my attention rivetted on
the buck I never saw her till she sprang from the tree. An ex-
amination showed her claw marks, and [think she was at a height
of 12 feet from the ground and about 10 feet from the buck, upon
which she was doubtless just about to spring. ‘The tree was
not quite vertical, and the part she was sitting on was almost
horizontal.
Tt is often said that only old toothless or injured lions tum
man-eaters! ‘Ihe following incident is a good commentary, A
native woman at Tindiga, afew miles from Kilosa, venturing out-
side her hut 8.ii.21, was sprung upon by a lion and eaten in the
mealies scarcely 50 feet from the hut, On:11.11. they visited
a cattle boma and paraded round and round it. On 12.11. they
squatted one on each side of a jumbe’s door, as was evidenced by
their spoor; they then went off and pounced for a man’s foot
through a reed wall: the occupants scared them off by making a
noise. On 13.ii. one of them tore outa bundle of grass from
EASD® AFRICAN MAMMALS. 723
the side of a hut, in an effort to reach the Sage esa who also
scared them off with ai outery. On-15.ii.21 we heard their
persistence had been rewarded, for they killed a man at Kivungu
and ate him except for the head. They appeared to be changing
their beat and making towards Myombo, where they mauled a
man a fortnight ago who was rescued by his friends.
A scout had been sent to Kivungu to set traps and was in a
hut, when the door was burst open by a lion: a woman was
sleeping close to the door. He ran outside and saw the lion
standing not twenty feet from -him, but could not fire as it was
in line with another hut. On 17.ii. they broke into another
hut at Kivungu, five hundred yards from the one forced the night
before. The only occupants were a woman and child, and she
scrambled to the flimsy reed shelf above the door and screamed.
She told me that the lion stood in the doorway but was afraid to
enter. The neighbours then beat tins and lit torches. so that she
was able to run across to an adjacent hut. When they had gone
to sleep and all was quiet the lions returned, dragged out the bed
to the edge of the clearing sixty feet away and ate the plaited
cords, which doubtless veeked of human beings, and smashed up
the framew ork, which I myself saw early the following morning.
On 18. 11. they broke into a hut between Kivungu and Myombo
and dragged out a basket of beans and ate the basket ;, they then
crossed some five miles of country to Tindiga, broke into a hut
there, and nearly got a man. On 19.ii. one of them entered
a hut before it was dark—they were obviously hungry—it was
frightened off and a trap set, which caused its death at dusk when
it returned. Its companion entered a second trap a few hours
later, and the man-hunting automatically ceased.
I measured these lions carefully and independently, and found
them exactly alike with the possible exception of a half-inch
difference in the length of ears. Both were tawny, maneless
males in well-nourished condition, presumably brothers of the
same litter. ‘The one had the remains of a wild-pig and some
grass 1n ifs stomach; the other had nothing but some parasitic
worms. Length of head and body, 67 inches. Tail 31 inches.
~ Another instance of man-killing occurred on the night 8-9. v.21,
when a.lioness appeared to have become desperate from hunger.
‘Che circumstances as I reconstructed them from the spoor and
native statements were thus :—The lioness first appeared (unseen),
in some rank grass outside a native kraal, where it crouched,
wriggled about, and watched a large party sitting around 2 fire
inside the stockade. She then rose up and, with a “ woof, woof,”
trotted through the doorway and seized a youth of fourteen by
the thigh, nin started back for the doorway. The jumbe, who
was a very old man and decrepit, pursued the lioness, striking
her with his bare hand on her quarters as she made for the
doorway—the compound was an unusually large one. When she
reached it the body stuck crosswise, and after a couple of attempts
and with the persistent old man stiJl spanking her, she dropped
724 MR. A. LOVERIDGH; NOTES ON
it, Sprang over it and made off. The boy was quite dead. I could
see very few marks on the body except where she had carried him
in her jaws. |
She then crossed the piece of waste land and came out ona
path where she clawed up the ground considerably, presumably
to clean her claws, or in a fit of annoyanee. She followed the
path for half-a-mile, then cut across another fifty yards of rank
weeds, and came to the edge of a little clearing where a woman
was sitting under the eaves of her hut shelling maize into a dish ;
her baby was slung on her back. Again the lioness lay down,
to take her bearings I suppose, then sprang upon the woman,
whom she carried off to beneath a tree fifty yards away. The
woman screamed, and beat the lioness about the face with her
bare hands; neighbours seized firebrands, tins, ete. with which
to make a noise, and sallied forth. They caught sight of the
lioness crouching over the woman beneath the tree, where it had,
without doubt, taken her to eat; at the sight of the rescue party
the lioness bolted. ‘This was about 7.30 pm. The baby was
unhurt, tiie mother horribly mauled.
The animal next made for Kilosa, and on reaching the village
of Mkwatani broke intoa hut where two women lived together.
Tt smashed down the door, which was only made of matama stalks,
dragged the body ten feet from the door and ate half of it.
There were six other huts within a hundred yards, and when the
neighbours rose next morning they caught sight of the lioness
crouching over her prey, she also saw them and cleared. A
messenger brought me the news soon after daybreak, and I sent
him back with instructions on no account to move the body. I
set off on my cycle, and when close to Mkwatani another native
met me with the information that the lioness had returned in
bright sunshine at 8 a.m. and carried the body off into the
matama. I followed the trail (which was well marked with
various items such asa bit of gory rag, some toes, bits of fat and
the like) for a hundred yards, when the matama became so thick
one could not see ten feet away, and had perforce to make con-
siderable noise in forcing our way through it, so we returned—the
boys to make traps, while I went on to investigate the other
“kills” already referred to. At 5 p.m. an askari on his own
initiative wormed his way up to the lioness as she was feeding
under a mango tree, and riddled her with bullets from a few feet
awny.
A Wippoboscid (Z/. capensis. v. Olf.) was taken on one specimen,
With Lions at their Nill.
One morning, in October ’21, I was sitting some fifty yards from
a, water-hole, when I noticed a little group of animals wending
their way through the thorn-bush towards the water, from which
they were still a hundred and fifty yards. At the first glance
I thought that they were a little group of buck, but a second later
EAST AFRICAN MAMMALS. 725
saw that the leader was a maned lion, followed by another full-
grown and four nearly full-grown cubs. At the first shot these
made off, but concluding that they must come to water some
time, I had a little stockade—perhaps ten feet long by five feet
wide—constructed between two small trees about fifty yards from
the water. At £ p.m. in the afternoon I took up residence in
this little cage and awaited events.
All was very peaceful except for some fifty or more doves assem-
bled around the water-hole ; at intervals of ten minutes these took
fright and with much clatter flew to the surrounding trees, but
soon returned to the waterside. A large flock of guinea-fowl fed
up to within fifteen feet of our hiding place. Little else occurred
to break the monotony of our wait until 6 p.m.: three fine eland,
but with very average horns, emerged from the thorn-bush
scrub and came within fifteen to thirty feet of the hide-up, the
nearest was certainly not more than fifteen feet away. The hoy
had set two gin-traps by the water’s edge and had been told to
remove the pins, which he said had been done: fearful lest the
antelope should get caught in these, I serambled up the palisade
(some nine feet in height), shouted, shoo'd, growled and wildly
waved my arms in an effort to scare them away. The stupid
creatures stood stock still for fully two minutes before wheeling
and cantering off.
From 7 to 9 p.m. I slept a good deal, though constantly aware
of the presence of hoofed animals in the vicinity: these ap-
proached the water with great caution and then stampeded, indi-
viduals passing within a few feet of our stockade on several
occasions. I imagine it was shortly after 9 o’clock (the boy had
mislaid the matches, so we could not have a light, even had we
desired one) that I suddenly became wide awake with a strong
sense of something wrong. A lion was padding round the
stockade and snufiling at times ; he was within five feet of me, but
vainly I sought to pierce the darkness to get a shot. The foot-
steps would cease, but at what angle to fire I could not decide,
and I was above all anxious not to leave a legacy of a wounded
lion behind for the local natives, twelve of whom had been carried
off by lions in this district in the preceding month. ‘These deaths
had occurred for the most part along one stretch of road about
twonty miles in length in broad daylight, so that the road was
now taboo. |
Gradually I realised that, more than one lion was paying us.
attention, as heavy breathing could be plainly heard at several
different points, and at times it seemed that some were lying
down whilst their companion tried to scare us out. The smell of
them was very strong in my nostrils; my companion, who had
wept himself ta sleep at the prospect of an awful end, still slum-
bered peacefully. With startling suddenness a loud grunt was.
given perhaps twenty feet away, it was followed immediately by
a noise of animals rushing straight towards us. I waked the bey
just as I was able to distinguish the hammer of hoofs, and a herd
Proc. Zoon. Soc.—1923, No. XLVIII. 48
726 MR. A. LOVERIDGE: NOTES ON
of eland came sweeping down the hillside through the thorn-bush
in.a most reckless manner. ‘The lions lying on either side of us,
however, rose up and headed the mob with. grunts, so that they
wheeled at exactly twenty feet from our hide-up and swept on
with the magnificent abandon of a cavalry charge. ;
Confused noises about a hundred yards further on followed by
a roar (such as I have heard a felled bullock in a slvughter-house-
give), announced that an eland had been downed by other lions
lying in wait, whilst the remainder of the herd crashed away
through the scrub. Right on the heels of the herd as they
passed us came at least three lions, whose easy swinging gallop
sent a shiver down my spine; they passed so close that we could
hear them panting, and on reaching the kill they made a
variety of noises, mostly grunts of satisfaction. ‘hen one gave
two roars, which were answered by lions grunting all up the
valley. .
T was greatly impressed by the splendid organization of the
drive—lions posted all along the valley to head the herd off. ‘The
final coup from the signal grunt to the roar of the beast, which
had been sprung upon us, was scarce a minute, I should imagine,
and took place with such a. startling suddenness after. the
former silence that it rather put one’s nerves on edge. ‘They
soon began rending and feeding upon the carcass in a very.
amicable way to the accompaniment of many small sounds, which
made me think that they were really a great deal nearer than.
was the case, for daylight revealed the kill between a hundred
and a hundred and fifty yards off, so that I concluded the animal
must have carried its lion for the best part of fifty yards after it
was sprung upop-—and what a beast it was! A fine female eland,
with horns 273 inches long (front-edge measurement), heavy in
calf, which must have been very near birth to judge by its hoofs,
which were all that remained of it. All the meat excepting the
head and legs had been eaten, the legs being still attached to the
skin with some vf the backbone and broken basal portions of.
the ribs.
To return to the lions, however. Following the roays and their
replying grunts, came the lons themselves in ones and twos,
grunting all along the way, and many of them passing close to my
place of concealment, Though a gentle rain was now falling they
presumably smelt me, for several would pause for a moment before
padding on to join their comrades at the kill., What astonished
we was their good behaviour, an almost entire absence of growling
and snavling, merely the munching, lapping and tearing of food,
and the small grunts of satisfaction already alluded to. About
an hour later—so it seemed to me, but I may have been wrong—
a party went down to the water, which was only fifty yards
away, and the sound of lapping was very loud; then some indivi-
dual found the traps (these were set but the pins not removed),
which they growled at and dragged about, together with the heavy
tree-trunk to which they were chained, in the most astonishing
EASY AFRICAN MAMMALS. 727.
way. On hearing the continuous rattling of the chains and the
growling, we congratulated ourselves on having got one lion at
least.
They presently returned to the kill to discuss dessert, and dis-
cuss it they certainly did, growling, grunting and snarling as
they crunched the bones, which they would drag away and eat by
themselves: some walked about and grunted. ‘This disconcerting
commotion continued till a couple of hours before dawn: there
was one interruption when two lions came along the ridge
forming our side of the valley and gave a peculiar sing-song call
that I have never heard from a lion before, and strongly reminded
ine of the ery of a hunting-dog ; this was answered by a lion at the
kill, which left the kill and walking past us, calling the whole time,
joined the others on the ridge above, when they all went off in
the direction taken by the mob of eland. I imagine the peculiar
ery of the lions was a call to come and join another hunt. Not
long after this a lion came back to us and pattered round, lay
down and breathed hard. Though we were having occasional
flashes of lightning and a good deal of thunder the darkness
was absolutely impenetrable, and while listening for further move-
ments on the part of our visitor, I fell asleep.
When I awoke dawn was breaking, but still the light was too
poor to see a rifle sight. I jumped up, and crept silently to the
side of the stockade nearest the kill. Tora moment I saw nothing,
then in the grey light I made out a lioness standing with her
whole flank exposed not fifty yards away, she was looking towards
the water. IJ turned round and whispered to the boy to hand me
the rifle, our quarters being somewhat cramped: in doing so, he
made some small sound which caused the lioness to spring round
facing me and growl, at the same moment a very small cub came
acing up from the water's edge, and the pair of them trotted off
immediately and obliquely. I had scarcely time to thrust the
muzzle of the rifle through the fence and take a hurried shot
after them, before they were lost to sight in the maze of thorn-
bush. The boy scrambled up the tree which formed one end of
the stockade, and said that he could see a big maned lion making
off from the kill, which was hidden from our view by intervening
thorn-bush; he was already a long way off, being doubtless startled
by the shot. So, after all, we returned to camp empty-handed,
but not regretting a very interesting experience.
FEeLis PARDUS SUAHELICA Neum.
Two specimens gun-trapped with :22 at Kilosa. Like the last,
the Leopard. has such a wide distribution in Tanganyika that to
record localities seems but a waste of time. :
Native names seem to be generally derived from the Kiswahili
Chui; in Kinyaturu, Mui; in Kisukuma, Sui; in Kikami, Duma;
in Chigogo, Suwi.
Both the above-mentioned specimens were immature—viz.,
48%
728 MR. A, LOVERIDGE: NOTES ON
male 462. 436. 82. 44mm. and female 1092. 737. 229. 77mm.
A larger male measured 1240. 660. 240. 73 mm., and female 1141.
659. 215. 76mm.
The latter, being also a comparatively young animal, had only
two foetuses (Kilosa, 20.ix. 22), of which the male weighed
pound and measured 217, 110. 42. 10 mm., and the female
206. 107. 39. 10mm. ‘The coloration and spotting most distinct
in front of ears, where hair was quite well developed, spotting (no
coloration) distinct all round area between front and hind limbs,
both belly and back, also on tail. No spotting on fore-limbs, and
only on the outer basal portion of hind-limbs.
At Kilosa the staple food of the leopards would appear to be
baboons. Mys. Turnley was fortunate enough to see a leopard
carry off a baboon at 3 p.m., only 200 yards from her house.
The baboons mobbed the leopard, which made off towards the
hills. By day, however, the baboons are generally masters of the
situation. On one occasion, a native reported baboons mobbing
a leopard within a quarter-of-a-mile of the house, and on arriving
at the place I saw the leopard was 60 feet up a tree. I had two
shots but missed, the second striking tha limb on which the
leopard was crouched among the foliage.
Another time I had sent out two collectors in the morning,
when one returned reasonably excited to say that they had heard
a great outcry of baboons on the opposite slope of a small rocky
ravine, and on going nearer to see what the uproar was about,
they saw fonr old baboons surrounding a leopard and striking at
it with their hands: as surely as the leopard turned on one he
would be fiercely attacked by those behind, so that he was very
much at a loss to know what to do and sought refuge beneath a
bush. - The one boy had remained to watch the proceedings
whilst the other brought the news.
We hastened to the spot, which was fully half-an-hour distant,
only to find that there was no boy or baboons there. My com-
panion was inclined to think that they were romancing, but
having come so far, he crossed the ravine or donga lower down
and went to the place where the alleged fight had taken place,
T remained on the near side. He called out that the spoor spoke
for the truth of the boy’s statement, and easily traced the leopard
from the bush to the donga, on the edge of which the baboons
had collected and torn up the turf in their fear, wrath, and
excitement. My companion said it was obvious that the fight
had gone on, as the baboons would not have left the leopard in
the donga, bat just to make sure, he ordered the boys to throw ina
shower of stones. ‘The next minute I called out something, and
there was an angry snarl in the bushes on the edge of the donga
not ten feet from me, and the grass swayed. The leopard, dis-
turbed by the noise on the far side and the shower of stones, had
been coming up when my calling out had turned it. We beat.
down the grass and found its spoor on the spot where the grass.
had been set in motion. The donga we subjected to a very
HAST AFRICAN MAMMALS. 729
thorough search, but owing to the amount of cover the animal
got away.
Dogs would seem to be the next favourite article of diet, one
was taken off the verandah of a house near mine. At 5 a.m. one
morning I was awakened by a very horrid noise, rather like cats
quarrelling, but dying away in a moan followed by several other
moans, It was only sixty feet from the house, and I ran out
with a lantern, but could see nothing.
My neighbour’s dog, a biggish animal, was in the habit of
coming to ‘this. spot on my path to relieve itself every morning,
and the leopard had no doubt become aware of this. My boy
took up the spoor and traced it for two hundred yards into the
rubber plantation, where he found the dog with only one haunch
eaten. Mr. C. I. M. Swynnerton sat up by this and saw the
leopard as it returned at dusk, but the leopard seeing him also
made off before he had time to ‘fire,
Goats make a very effective bait for leopard traps, but on one
occasion, the setting of the string being too slack, a leopard was
shot dead just as its jaws closed into the goat’s skull so that they
both died simultaneously, the leopard not even opening its jaws
which had penetrated to the brain. Remains of goats were found
in the stomachs of several leopards.
Bushbuck are often killed by them; in the case of one old male
killed in a donga (21. xii. 20), it could be plainly seen that the
leopard had sprung on its back and clawed its throat. It only
ate from the haunches, but the next night returned, dragged the
body twelve feet away, and made another meal from it. On
another occasion a gin was set on a path much frequented by
leopards; a female bushbuck was caught in the gin, and the
leopard killed and ate part of her. A native going to examine
the gin in the morning brought the remains of the bushbuck
hack to eat himself, but was made to return it. The leopard
returned, adroitly avoided both gins which had been reset, and
dragged the carcass half-a-mile away upadonga. The boy took
up the spoor in pouring rain and came on the leopard, which gave
a snarl and made off.
On the Wembere plains I came on two instances where leopards
or a leopard had kalled impalla; in oné case a mae and im the
other a female. At Kipera, between 5-11. ix. 22, leopards kiiled
a male reedbuek, a calf eland, and a Lichtenstein’s hartebeest in
calf, a few days later another reedbuck. In the case of the
hartebeest it might have been the work of lions.
At times they will tackle porcupines. An almost full-grown
Jeopard entered a hut at Tindiga, where it was shot by a native,
who brought it to me in the flesh. It was in a most frightful
condition, covered with sores, from one of which I. recanered a
broken portion of a porcupine quill. On its neck was a bare
patch a foot in length and two inches broad in its widest part ;
the patch was hard dried skin and may have been made by the
leopard clawing at some quill-stumps left in the skin.
730 MR. A, LOVERIDGE: NOYES ON
This was the second leopard in a hut at ISilosa during the
week, fora native awaking to hear something moving about in
his hut, jumped up and got a cuff from a leopard as it made off.
Tt had doubtless entered in search of fowls. ‘T'wo days later, on
the same Otto Hstate, another man heard something in a bush
near his hut, and calling some friends began poking about, with
the result that a leopard sprang on him, clawe ed his enouldexe and
breast quite unpleasantly, and made off as suddenly as it arrived.
T imagine that the man anticipated a bushbuck and surrounded
the bush with his friends, so that the leopard acted in self-defence
and the man got more than he bargained for.
Unless cornered, or wounded, it is unusual for ‘Tanganyika
leopards to molest people. When at Mdjengo’s (7.x. 22), tho
jumbe came to me and said that a leopard, which had been
carrying off a lot of fowls of late, had the night before sprung
on a sleeping child just inside the door of a hut. The animal
‘dropped the youngster almost immediately an outcry was raised ;
it seems possible he mistook it for a goat, though I heard of
another well-authenticated case where the family were sleeping
outside the hut on account of mosquitoes, and a leopard carried
off nnd ate a child. In both instances, the leopards, were killed
by Game Department Trappers. At Kisanga, i in Kilosa District
(11. 22), I heard of a woman who was working in the fields being
sprung on by a leopard and so badly mauled that she died shortly
afterwards: the lecpard was killed by a man who came to her
rescue.
Parasites taken from the one young leopard included a fly
(Hippobosca capensis v. Olf.), fleas (Clenocephalus felis), two
species of ticks (a@maphysalis leachi and LIC aa sumus
Koch), and worms (Physaloptera preputialis v. Linst.). ‘The
second leopard had nematodes (Onchocerca sp. females) in its
neclx.
Friis CAPENSIS HINDEI Wrought.
Kizongoduma in Kikami and Kipogoro, Kijongo in Kisagara
and Kiswahili, Nzuli in Chigogo, and Nduri in Kirvamba.
Four specimens examined from Tindiga, Kilosa, and Sagayo.
The largest male measured 900. 300. 158. 80mm., and female
670. 260. 170. 77 mm.
At Kilosa on 12. vii. 21 some natives cutting grass disturbed a
serval, which bolted up a tree leaving a large kitten at the foot
of the tree. This was brought to me and ‘commenced to take
milk at once, first from a spoon, and then going to a saucer of
its own accord. It spat and clawed a good deal when approached,
but seeing that it had been dragged along with a cord round its
neck, it is not to be <padlonsal ah A mmernelh later, however, it
was still implacable, rolling on its back clawing, spitting and
biting, so [ chloroformed it.
One male was trap-shot through the spine with °22 when
EAST AFRICAN. MAMMALS. 731
attempting to reach the bait of hyrax meat only a hundred feet
from my tent. I got.up at 4a.m. without having heard the shot
and sent a boy to remove the gun, giving him alamp. Je found
the gun had gone off, and stooping over the ground followed the
spoor, when (he said) an animal menaced him just as he caught
sight of its wounded mate—the lamp chose this moment to go
ae ! He cried out, and came running back very frightened under
the impression. that there were two. leopards. 1 then went and
shot it through the head, it SEX dragged itself some fifteen
feet from the trap.
Its stomach contents eiuiisted of a great many house-rats
(R. 7. alexandrinus), and the remains of a skink (4. ferrandi).
My captive serval showed a marked preference for chicken meat,
though it would take vegetables at times. The stomach of
another wild serval held nothing Lut grass, possibly seized after
if had been trapped, as it was gin-trapped. Another was killed
entering a gun-trap baited with a live goat; yet another was
transfixed by a native’s spear as it was eating the native’s fowl]—
it was in very lean condition, which may account for it hunting
fowls in daylight at 5 p.in..
I encountered a serval at Chansuru at mid-day on one occasion,
the sun being at full strength at the time. he animal was in
the road, aro was hedged in by aloes so that it could not easily
escape, and il chased it for sone way on my cycle. Serval were
several times disturbed in long grass country, but they are very
active and get away almost before one has time to shoot.
Twenty fleas (Clenocephalus felis) were taken from one specimen,
which was also afilicted with ticks (//emaphysalis leachi).
FE vis OCREATA UGANDA Schwann.
Kilenga in Kikami, Kimburu in Kisagara, Mvugi in Chigogo,
Titu in Kinyaturu and Kiramba; Paka wa pori in Kiswahili.
Ten specimens from Jlonga, Kondoa, Tindiga, Kilosa, Dodoma,
Mdjengo's, and Mtalis
The largest male imensured 583. 285, 133. 61 mm., and female
530. 308. 120. 601mm. Tail-lengths seem to vary considerably
and not in even ratio with the body-length.
In the sandy thorn-bush country (Dodoma to Mtali’s) the Cats
were of a very pale type, and I was inclined to think, after
comparing the four skins with the six from Kailosa district, that
they indicated a pale type. Just before I left Hast Africa, how-
ever, Capt. Godman shot a very pale example at Kilosa, almost
in the spot where I had obtained some of the dark ones:.it was
quite indistinguishable from the thorn-bush type.
The dark wild cats are typical wild tabbies, and undoubtedly
interbreed with domestic cats at times. At Kondoa I was shown
v litter of six blind kittens under a pile of logs on the outskirts.
of an estate. Two of the kittens were all black, and four tawny-
tabby. ‘The house cat of the owner of the plantation was a black
on, MR, A. LOVERIDGE: NOTES ON
tom. I was called upon to shoot a brother of this black tom at
Kilosa, which had been out fighting one night, and came home
with the brain exposed and very much seratched ; presumably he
had had a dispute with a wild cat as there were no tame ones
anywhere near.
Most of these cats were trapped whilst neeipone to get at
fowls. ‘The stomach contents were :—(i.) fowl and grass ; Cia) lot
of grass ; (ili.) bodies of skinned birds; (iv.) two rats and a bird’s
feather; (v.) a rat (2. r. alecandrinus) and the remains of a fowl.
There were screw-worms:-oen intestines and in viscera of one
Tindiga specimen, but these were not preserved.
Lynx CARACAL NuBICUS (Fischer).
‘Simba wagi in Kikami; Simba mweng'we in Chigogo.
‘The Hast African Lynx is generally referred to the above race,
so I have ventured to put down the subspecies, though no speci-
men was obtained. I saw one skin from Itumba in Manyoni
sub-district in the possession of a friend, and the only one seen
besides was one offered for sale at Shanwa, which had presumably
been killed in the Mwanza District. The natives state that these
animals hunt in parties of five or six but that they ave rarely
seen.
UNGULATA.
BoviDs&,
@ Bos CAFFPER RADCLIFFEL Thos.
Two specimens, a male from Myombo and female from Uluguru
Mts. Very common at Kilosa and throughout the district, they
frequently enter the Otto Plantation and have been seen within
a few hundred yards of the houses.
My native collector, Salimu, shot the cow with a 12-bore gun
in circumstances of sufficient interest to justify my repeating
them here. Buflaloes had been doing damage in «a shamba on the
mountain-side, and the owner of the plot spent several nights
guarding his crops. He was only armed with a spear, and on one
night was chased by ayoung cow (he may have attacked the animal
first), the native clambered on to a rock, and the angry animal
fumed at the base of it attempting to reach him: he leaned over
and stabbed it with the spear about six times along the spine,
ths spear bavely penetrated below the skin. ‘The animal did not
leave the sbamba with the rest of the herd next morning but
Jay up in the mtama near a path, It charged two natives,
throwing them down and sticking its horn into the side of one,
but the horns being short not much damage was done.
Knowing nothing of this, Salimu was returning from collecting
birds the following morning when Jumbe Magoma (the local
headman) came running towards him along the path and un-
ceremoniously clambered into a tree. Salimu called out to know
EAST AFRICAN MAMMALS. 733
what was the matter, and hardly had received the reply ‘ Buffalo’
gasped out by the somewhat winded jumbe, when round a bend
in the path came the infuriated buffalo. Salimu states that he
jumped up on a rock some eighteen inches high close to the path,
and as the animal came for him he fired the only charge of S.S.G.
which he had full in its face. The animal stopped and shook
the blood from its face, and profiting by this interval Salimu
essayed to clamber into the jumbe’s tree, but before he had made
good his seat the buffalo butted the tree and nearly succeeded in
dislodging him. As soon as he was firmly seated he took the gun
from the jumbe and loaded it with No. 3 shot, which he fired in
the animal’s face every time it came back. In all he fired at it
six times before it drew off, swayed, and finally fell. :
An examination of the skull shows that the 8.S.G. smashed
a large hole in the frontal bone; the sho$ was very close, so that
the animal must have been nearly on to him when he fired. The
skull was pitted with No. 3 shot in every direction, most of which
cid not penetrate, but I imagine some entered by the hole already
made by the 8.8.G., and the animal mainly succumbed to loss of
blood from the first charge, -
BUBALIS COKEL WEMBAERENSIS Zuk.
A single male (1860. 370. 520. 200 mm.) from the Wembere
Plains near Dombolo, 9.xi.21. I use the above, possibly invalid,
subspecific name as this Hartebeest is from the type-locality of
this alleged race.
BuBALIS LICHTENSTEINI Peters.
A male (1900, 482. 513. 210 mm.) and a female were shot at
Kipera on 8.ix.22. ‘The latter held a female feetus (630. 220.
260. 90 mm.), and I collected twenty maggots of a_bot-fly
(non det.) from the nostrils of the adult. The species is quite
common at Kipera, the largest herd numbered perhaps twenty,
but they usually go in parties of five or six. When a suitable
thicket is found in which to rest, they return to the same spot
day after day.
DAMALISCUS KORRIGUM JIMELA Matsch.
A male (2230. 370. 435. 190 mm.) and female (1890. 360. 435.
190 mm.) Topi were shot at Sagayo on 3. xi. 22. This must be
very near the type-locality of the subspecies. There were two
other adult females with calves running to heel with this bull;
the female shot was the smallest of the three. In its stomach
were grub-like parasites.
ConNOCH&TES TAURINUS TAURINUS Lyd.
Nyumbu in Kikami, Kisagara, and Kiswahili.
A single male Wildebeest (2380. 530. 480. 225 mm.) on the
Mkata Plains, 13.ix.21. It was accompanied by four females. -
734 Mh, A.’LOVERIDGE: NOTES ON
.CEPHALOPHUS MELANORHEUS SCHUSTERI Matsch.:
Sesi in Kikami.
A pair of Blue Mountain Duiker (¢. 610. 190. 88. 50 mm.
2. 670. 180. 60.55 mm.) were killed by natives in their gardens
at Bagilo, Uluguru Mts.
CEPIHALOPHUS GRIMMI SHIRENSIS Wrought.
Funo in Kikami, Kisagara, and Kiswahili.
A single pair of Duiker were shot on the hills behind Kilosa
during December 1920 (d. 825. 100. 225. 100 mm.). They
are quite common, but owing to the long grass and the quickness
of their hearing, very difficult to procure.
A fly (Zippobosca capensis v. Olf.) was taken on one of these
Duikers.
OREOTRAGUS: OREOTRAGUS SCHILLINGSI O. Neun.
Ngulungulu in Kisukuma and Kiramba. The Kiswahili name
for all Klipspringers, viz. Mbusi mawe—the goat of the rocks—
is in general use however.
Schillings’ Klipspringer was met with at Mtali’s and Sagayo,
v pair being obtained (fg. 630. 35. 190.80 mm.). When feeding
among the bushes at the foot of their kopjes in the early morning
they appear boulder-grey, and may easily be mistaken for Duikers
if not expected.
OUREBIA corroni Thos.
A single male of Cotton’s Oribi (770. 70. 200. 150 mm.) was
shot at Sagayo on 6.xi.22 in grassy maiombo and thorn-bush
country.
RAPHICEROS CAMPESTRIS NEUMANNI Matsch.
The subspecific name is giyen on geographical grounds, for my
specimens combine key characters of several races of Lt. campes-
tris as given on page 149 of vol. il. of the ‘Catalogue of Ungu-
lates.” They have (i.) a dark coronal mark, (1i.) a dark brown,
triangular nasal spot, (ili.) a white ring round the eve and much
white on the oral margin, lips, chin, throat, and inner surface of
the limbs: Having compared them with skins in the British
Museum, it seems very doubtful if some of these races are
sound,
Three specimens of Steinbuck from Shanwa and Sagayo, viz.
male adult (909, 50. 209. 105 mm.), female adult (855. 49. 205,
105 mm.), and female immature (540, 40, 190. 91 mm.). ‘he
latter was kept in captivity for three months and made a
charming pet. It was only given milk, and grazed about for
itself in the vicinity of the camp. When it wanted milk it used
to attack the backs of the native’s knees with vigorous pushes
accompanied by loud sucking noises, and would give no peace till
EAST AFRICAN’ MAMMALS. 135
attended to. Its death was solely due to the stupidity of a
porter, who was carrying it and took no notice of the little
creature strangling itself.
RHYNCHOTRAGUS KIRKI NYIK#& Heller.
Kizimba in Kisagara, Chizimba in Chigogo, Sala in Kisukuma.
Paa for all Dikdik in Kiswahili.
Hight males and six females were shot at Mbala, Igulwe,
Pooma, Mtali’s, Usurwe, Usshora, Ulugu, Mkalaina, and Sagayo.
The largest male measured 725. 41. 200. 80 mm., and largest
female 700. 50, 210. 80 mm.- They show a good deal of variation,
particularly as to the presence or absence of white spots upon the
legs.
At Igulwe a female was shot on 7.ii.23 which held a small Q
foetus, measuring 85. 4. 22.6 mm. The Usshora specimen was
caught by a dog, and I imagine may only have been dropped that
day (30.x.21). It lapped milk readily, but died on the night of
the second day though excreting normally. At Mbala on 27. 11. 23,
an immature male, whose horns were just sprouting (d. 580. 33.
175. 70 mm.), was running with its mother (9. 690. 50. 195.
75 mm.); who appeared to be still nursing him.
These little buck are quite common throughout the Dodoma
thorn-bush country, often in places where there is not a drop of
water and the vegetation apparently dried up. They have a habit
of resorting to the same spot to relieve themselves, as a civet does
also. At Singida I came upon a place where both these animals
had been using the same spot for several days at least.
The Pooma specimen was heavily infested with fleas, which
have been identified as Otenocephalus felis and rsidis.
ReEDUNCA REDUNCA ToHI Heller.
Tohi in Kikami and Kiswahili; Nhohe in KKisagara.
Though Reedbuck were comparativély common a few miles
from Kilosa, I never shot one, and the only specimen obtained
was the skull of a male shot by Capt. Turnley on 28. xii. 20.
Reedbuck were several times found which had been killed by
leopards.
IXoBUS ELLIPSIPRYMNUS Osg.
Kuru in Kikami and Kiswahili; Nhulu in Kisagara.
A male Waterbuck (2110. 390. 490. 210 mm.) at Kipera on
26. viii. 22, and a female (1750. 340. 470. 200 mm.) from Mkata
River on 1.ix.21. The latter had a very minute foetus. One
was found that had been killed by lions near the Simbiti River.
ZEPYCEROS MELAMPUS SUARA Matsch.
Palla in Kinyaturu and Kiswahili; Mpalla in Karamba.
A male Impalla from Zengaragusu on 2.x. 21, a female (1040.
736 MR, A. LOVERIDGE: NOTES ON
138. 142, 130 mm.) from Mbulw’s held a large foetus on 14. x. 21,
whilst another female shot at Sagayo on 31.x.22 held a very
minute one. Large herds were met with on the Mkata Plains
during the dry season. ‘I'wo instances of Impalla being pulled
down by leopards on the Wembere Flats during the dry season
came under my notice,
GAZELLA THOMSONI Giinther.
Lala in Kisukuma and Kiswahili.
A single male Thomson’s Gazelle (1165. 245. 3850. 130 mm.)
from Sanga, and six females from Nduguyu River, Sanga, and
Mwadira, of which the largest measured 1160, 200. 350.
120mm. Of the two females from Mwadira, one had a back-
turned horn nearly growing into the skull between the ears,
whilst the other had a horn missing. This latter held a foetus
(19. x. 22), as did one of the Sanga specimens shot on 16. x. 22.
HiprotTraGus EQUINUS LANGHELDI Matsch.
Kolongo in Kinyamwezi, Kisukuma, and Kiramba. Iorongo
in Kiswahili, obviously a borrowed and corrupted name.
Roan were met with at Izikisia near Tabora (‘Tabora being
type-locality of the subspecies), Mlewa’s in Mkalama, and Sagayo.
A pair were shot in the last-named locality, where the species is
quite common ; herds numbered from five to thirty individuals,
one or two herds being met with daily. The male measured
2240, 480. 600. 290 mm., and the female 2280. 430. 480. 282 mm.;
the latter held a foal nearly ready to drop (¢. imm. 760. 180.
330. 138 mm.), which was somewhat of a surprise as the males
were running with the does. :
In the stomach of the female were tapeworms (Stilesia hepatica
Wolffhugel) and nematodes (Setaria labiato-papillosa Aless.),
whilst microfilaria of the unsheathed variety appeared in blood-
films.
HIprorraGus NIGER ROOSEVELTI Heller.
A. single female (measuring 1900. 350. 350. 210 mm.) was shot
on the hills behind Kilosa on 20. xii. 20. There were some thirty
animals in the herd. A larger herd was met with at Mkata,
whilst single individuals were of common occurrence at Kipera.
Tabanids were present in large numbers on the animal that was
shot, but no tsetse.
TRAGELAPHUS SCRIPTUS MASSAICUS Neum.
Mbala in Kikami and Kisagara.
Bushbuck are very numerous at Kilosa, but only a pair were
shot, the male with a shot-gun from the verandah of the house.
A female was shot on the banks of the Simiyu River on 31, x. 22,
EAST AFRICAN MAMMALS. 137
which has been referred to this race though the two skins are
very different.
Measurements of Kilosa @ 1170. 180. 340. 130 mm., and
Sagayo @ 1180. 150. 335.140 mm. In the latter was a fetal
female, 480. 68. 180. 76 mm.
A male was found dead in fine condition, and I attribute its.
death to the bite of a puff-adder, there being a puncture on the
haunch, the blood was fluid and much hemorrhage about the vent
(Itilosa, 26. iii.21). Another was found in a donga that had been.
killed by a leopard, which had sprung on its back and had heen
carried some yards by the buck—a very old male. Setting gins
for leopards on paths frequented by leopards is not altogether
satisfactory, as I know of two instances in which bushbuck
stepped into the traps and were caught; in one of these cases
the leopard added insult to injury by carrying off the buck.
SuIDS.
CHorROPOTAMUS CHOIROPOTAMUS D&HMONTS Major.
Nguruwe in Kikami and Kiswahili; Ngubi in Kisagara.
A young male (730. 305. 235. 120 mm.) Bush-pig was shot by
ny collector in rank grass into which the native had forced his
way to pick up a monkey. A large pig which accompanied it
(? mother) attempted to urge it away by rending its belly with
her tusks, and inflicted no fewer than twenty-three large cuts as
she endeavoured to push it along (‘Tindiga, 28. i. 22).
Mrs. Billinge, of Ivinga, wrote me that the wild pigs of
tumruli appear to be a smaller race, very fierce, and that “ they
play havoe with dogs.” She sent in a half-grown one which,
being confined in a room, sprang up aruund the walls to a height
of two feet clear of the ground and eventually killed itself
(3d. 800. 240. 180. 120 mm.). “Three other captive young ones
died as a result of being exposed to the sun, and one of these
which I measured was ¢. 400. 135. 110. 65 mm. (Kilosa,
1. ii. 22).
PHACOCHGRUS HZtHIOPIcUS MAssaIcus Linnb.
Ngiri in Kikami, Kisagara, Kisukuma, and Kiswahili.
Four Warthogs were shot at Sagayo, of which the largest male
measured 1110. 360. 210. 120 mm. and was plumbeous in colour.
A female (1220. 350. 195. 119 mm.) shot the same day was
brickish red. A few days before (2. xi.22) a plumbeous female
was seen accompanied by three bright rufous young ones.
On the Mkata Plains one day I stood face to face with a Wart-
hog at about a hundred yards distance and mistook it tor the
gnarled roots of an overthrown tree; a second Warthog standing
immediately behind the first, but with its head down, helped to
foster the illusion of a fallen tree-trunk.
The Sagayo specimens were swarming with tsetse (Glossina
738 MR. A. LOVERIDGE: NOTES ON
swynnertonit Austen), and had many worms (Ascaris lumbricoides
L., young 2; Wsophagostomum sp. n.) in their stomachs, but
blood-smears taken from them gave negative results.
Equip.s,
Equus QUAGGA CrAwsHAYI de Wint,
Punda milia in Kiswahili, Sangeri in Kizungwa, Nhyenie in
Chigogo, Nguo in Kiramba and Kisukuma.
Three specimens were shot at Mtali’s, Simbiti River, and
Sagayo respectively. The Mtali specimen was a very old male
(2470. 470. 525. 190 mm.), the Sagayo female measured 2400.
430, 520. 170 mm., and was carrying a foal very near birth
(2. 1030. 220. 390, 105 mm.) on 2. xi. 22.
‘There were maggots in her stomach, a large tapeworm (Ano-
plocephala rhodesiensis York. & South.) nearly an inch wide, and
nematodes (Setaria equina (Abildg.), 9). The Mtali male had
worms (Crossocephalus viviparus (Linst.) Q and Strongylus vul-
garis Less. 2 9, and S. asini Blgy. § ¢), with numerous Cistrid
larve in its nostrils, and scores of ticks (Amblyomma sp., [yal-
loma wgyptium var., and Ithipicephalus evertst Nn.) about the
anus. x
ELEPHANTIDA,
ELEPHAS AFRICANUS KNOCKENHAUERI Matsch.
Ndoo in Kinyaturu.
A fact worth recording was the appearance of four Elephants
some 200 yards from the Boma at Singida at daybreak on 6. x. 22.
The country is for the most part open rolling downs with
occasional rocky kopjes here and there. It was surmised that
the animals wandered from the western side of the lake west of
the Boma, where Elephant are known to exist in the dense but
low thorn-bush scrub. ‘Vhey drank at the lake east of the Boma
and lost their way. ‘They were fired on by the native police at
the Boma, and the male was wounded, he was followed up and
killed in a native garden some five miles north. He measured
ten feet at the shoulder, and his tusks weighed about 40 Ibs.
apiece. One ear had a curious long tail about 8.inches in length,
doubtless caused by some injury. ‘he other ear, which I pre-
served and which is now in the Game Dept. Office, had two holes
in it as if made by bullets at some time or other. Presumably
IT am correct in referring it to this race.
PROGAVIIDA.
ProcAayia BRUCEL PRITLWITZI Brauer.
Mhimbi in Chigogo, Pimbi in Kinyaturu. Pimbi is also the
Kiswahili name for all species of Hyrax.
Eleven specimens from Gwao’s, Mbulu’s, Pooma, Singida (seen),
EAST AFRICAN MAMMALS. 739
Mdjengo’s, Mtali’s,and Tabora. Skins from Dodoma and Tabora
referred to P. fromm in my last notes have since been referred
to this race.
Largest male 450, 0. 64. 35 mm.; largest female 470. 0. 60.
32 mm. Preenant female with two minute foetuses at Gwao’s
on 10.x.21. Another also with two foetuses at Mdjengo’s
on 24.x.21, and two very large feetuses in a Tabora female on
INS}, Hal Alls
Like all the other hyraces their stomachs contained finely
masticated, bright green leaves of the shrubs which grow about
the bases of the rocky kopjes where these creatures dwell.
Fieas (Xenopsylla isidis) were very numerous on the Mbulu
specimen.
PROCAVIA BRUCEI MATSCHIEt Neumann.
Twenty specimens from Shanwa and Sagayo, where they are
very abundant on the rocks, and at the former place ‘ barked’
loudly when approached.
The largest male measured 530. 0. 75. 30 mm., and female
545. 0.67.30 mm. ‘Three of the Shanwa females held minute
embryos on 20.x.22. ‘Chere were three in one and two in the
others. ‘The stomachs of those examined were clean.
T had a young male alive for a fortnight, which fed readily on
green stuffs and allowed itself to be stroked, but was very nervous
of shadows and of being suddenly approached. It succumbed to
a severe hailstorm.
PROCAVIA TERRICOLA SCHUSTERI Brauer.
Mhelele in Kikami.,
Six specimens from the Uluguru Mts. Largest male 605. 0.
77.35mm. The largest female does not exceed the measure-
ments of the one recorded under Procavia sp. on p. 67 of my
earlier paper. The youngest female taken this time measured
940. 0. 45. 20 mm.
NERVOUS SYSTEM AND SUPRARENAL BODIES IN THE SPARROW. 741
36. Observations on the Development of the Sympathetic
Nervous System and Suprarenal Bodies in the
Sparrow. By A. Suppa Rav, B.A., M.Sc., F.R.MLS.,
Derby Scholar, Department of Zoology and Compara-
tive Anatomy, University College, and P. H. Joun-
son, B.A., B.Sc., F'.Z.8., F.R.M.S., Demonstrator in
Zoology, Department of Zoology and Comparative
Anatomy, University College, London.*
| Received May 29, 1923: Read October 23, 1923. ]
(Text-figures 1-20.) ;
INTRODUCTORY.
In the course of an investigation suggested to us by Prof.
J. P. Hill, as to the origin and development of the suprarenal
bodies in the Sparrow, we have been led from a consideration of
the source of the chromaffin cells in those bodies, to an exami-
nation of the development of the sympathetic nervous system.
The origin of this system hag been a much-debated question for
many years, and the excellence of the material provided for us
by Prof. Hill has tempted us to put forward an account of our
observations on this subject, more particularly as concerns the
relations of the so-called primary and secondary sympathetic
cords. !
While confirming on the whole the view maintained by the
majority of modern embryologists as to the ectodermal origin of
the sympathetic system, the investigation has brought to light
certain interesting facts which do not appear to have been
recorded by previous observers, while we venture to think that
the confirmation of previously recorded observations may be of
some value,
Our observations have been made on a series of nearly a
hundred Sparrow embryos collected during the years 1919-21.
Unfortunately little is known as to the exact age of these
embryos, but by a careful comparison of the stages of develop:
ment reached we have been able to arrange them in a series.
Keibel’s ‘Normal Table of Chick Embryos’ has been of con-
siderable assistance in this task.
The embryos were fixed by various methods, the most successful
being that of Gerhardt. Prof. Hill also placed at our disposal
some chick embryos treated by Bielschowsky’s method. The
best staining results were obtained by the use of Heidenhain’s
iron hematoxylin.
* Communicated by Prof. D. M.S. Watson, F.R.S., F.Z.S.
Proc. Zoou. Soc.—-1923, No. XLIX. 49
742 A, SUBBA RAU AND P..H, JOHNSON ON THE NERVOUS
As excellent historical reviews of this subject can be found in
the papers of Kuntz (11), Goormaghtigh (6), and Da Costa (8),
and in the ‘'Text-Book of Human Embryology’ of Keibel and
Mall, we do not attempt a further summary, but confine our-
selves to a restatement of previous observations on the primary
and secondary sympathetic chains in birds and in such other
. vertebrates as have been observed to possess them, .
Wn. His, Jun, (8), was the first to describe the occurrence in
chick embryos of primary and secondary sympathetic chains.
“He observed, in a 4-day chick, a sympathetic chain similar to
that found in other vertebrates, 7. e. situated immediately behind
(i.e. dorsal to) the carotids and on each side.of the dorsal aorta.
At the 8th day, however, he found a second cord immediately
in front of the anterior’ (ventral) roots of the spinal nerves.
This second cord, he says, soon: outgrows the former in size and
is enclosed in the vertebrarteria] canal. He was unable to trace
the subsequent fate of the primary cord.
In a later paper (9) the same author again discusses the rela-
tions between the primary and secondary cords. He is of opinion
that the primary cord in the neck region atrophies, whilst in the
thoracic and abdominal regions it forms the aortic plexus and
thence migrates along the mesentery to the organs situated in
those regions, He regards the primary sympathetie of the trunk
region as only a step on the path of the sympathetic cells from
the spinal ganglia to the periphery, and lays considerable stress
upon the resistance offered to the migratory nerve cells by the
mesodermal elements.
The secondary sympathetic cords he describes as the latest
formation of the spinal ganglia, and suggests that the formation
of the former is due either to a loss of “migratory power in the
sympathetic cells or to an increased resistance offered by the
mesoblast. P
Miss W, Abel (1) adopts the views put forward by His, Jun.,
but has observed a small contribution from the primary to the
_ secondary sympathetic ‘cord, although she lays little stress on
her important observation.
Kuntz (11) and Ganfini (5) support the main findings of His.
They have also observed in Chelonia the existence of a secondary
sympathetic. All these writers are agreed in describing the
secondary sympathetic as a new for mation quite independent
from the primary § sympathetic system.
A, Relations of the Sympathetic Primordium to the Neural Crest.
While Held (7) derives the sympathetic primordia exclusively
from the dorsal root ganglia, and Kuntz (12) is of opinion that
the chief part is derived by way of the ventral roots, our obser-
vations lead us to support the view put forward by Streeter (12).
He writes: ‘As the neural crest becomes detached and _ its
segmenting part invades the space between the myotomes and
SYSTEM AND SUPRARENAL BODIES IN THE SPARROW. 743
the neural tube, certain ganglion cells separate’ themselves from
its ventral border and independently migrate ventralward into
the neighbourhood of the aorta. It is these cells that form the
connected chain of errant ganglia which we know as the sympa-
thetic system.”
Thus in Sparrow 16 of 6-7 somites [= 32-hr. chick] we find
a well-marked neural crest, reaching from the dorsal border of
the spinal cord nearly to its ventral edge. From the distal end
Text-figure 1.
Transverse section of Sparrow 16 (6-7 somites), showing spinal cord and developing
neural crest. Krom the ventral end of latter cells (c) are proliferated towards
the carotid artery.
C. Migrating cells. ©.A. Carotid artery. IJfed.C. Ependymal cells,
N.C. Neural crest. Néec. Notochord. S.C. Spinal cord.
(Sp. 16, slide 3, row 3, section 12.3 .
of the crest, cells appear to be proliferated into the mesenchyme
between spinal cord and aorta (text-fig. 1). ‘The crest at first
consists, in section, of a single row of cells with deeply staining
cytoplasm and with large, oval, lightly staining nuclei.. They
have usually one large nucleolus, but two are sometimes present.
The cells of the crest show mitotic figures especially at its ventral
edge. Between the latter and the dorso-lateral angle of the
49%
744. ‘A. SUBLA RAU AND P, H, JONSON ON TUE NERVOUS
aorta there are isolated cells with identically the same charac-
teristics as those of the crest, and clearly distinguishable from
the surrounding mesenchyme cells. The latter have much
smaller and spherical nuclei and are distinctly branched, their
processes being united so as to form a cell network. Their cyto-
plasm stains much lighter than that of the crest cells. ‘he cells
proliferated from the neural crest are not found in every section,
but are most marked in the inter-somitic regions, where later we
find the ganglia of the primary sympathetic cords.
B. Relations with the Darsal Root Ganglia.
With the expansion of the neural crest to form the dorsal root
ganglia the process of proliferation of sympathetic cells increases,
and as is shown in text-fig, 2 from Sparrow 15 (= 42-hr. chick)
Text-figure 2.
‘Transverse section of Sparrow embryo 15 (= 42-hr. chick), showing a succession of
cells (Sy.C.) migrating to the dorso-lateral angle of the aorta.
do. Aorta, Sy.C, Sympathetic cells. D.R.G. Dorsal root ganglia.
My. Myotome, Née. Natochord. (Sp. 16, 2, 4, 16.)
there is a succession of cells apparently migrating to the dorso-
lateral angle of the aorta. These cells occur between the myotome,
now considerably developed, and the spinal cord. ‘The cells still
SYSTEM AND SUPRARENAL BODIES IN THE SPARROW. 745
possess Jarge clear oval nuclei above described, but the sur-
rounding cytoplasm appears to lose its individuality, with the
resulting formation of syncytial masses or strands, in which
there is evident mitotic activity. ohn (12) figures such strands
in the Rabbit. Streeter (12) considers this syncytial character
to be only transitory, and associates it with the period of migra-
tion. Contemporaneously with this migratory activity of nervous
elements the primordia of the segmental arteries and veins are
being laid down. ‘Thus in text-fig. 3 we see that the endothelium
of the dorso-lateral angle of the aorta has proliferated to form
a distinct strand of cells, which will ultimately develop into a
Text-figure 3.
‘Transverse section of Sparrow embryo 15 (= 42-hr. chick), showing development of
segmental artery and migration of sympathetic cells (Sy.C.).
Ao. Aorta. C.&. Celomic epithelium. Jfyt. Myotome. S.C. Spinal
cord. Seg.Aré. Segmental artery. Sy.C. Migrating sympathetic cells.
Nétec. Notochord. (Sp. 16, 3, 1, 1.)
segmental artery. Similar vascular strands may be seen in other
sections arising from the cardinal veins. The nuclei of the cells
in these vascular strands are very irregular in shape, and thei!
cytoplasm is considerably lighter and less granular than that of
the nerve strands. In view of the fact that some authors have
been led to consider such strarids as of nervous significance, we
venture to direct special attention to this differentiation.
‘That the cells migrating from the dorsal ganglia are neuro-
blasts is confirmed by silver preparations of chick embryos in
which the ventral roots have not yet appeared, i.¢. of less than
746 A. SUBBA RAU AND P. Ii. JOMNSON ON THE NERVOUS
60 hrs. Text-fig. 4 is drawn from a section of such an embryo.
It shows neuroblasts isolated between spinal cord and aorta
Text-figure 4,
Z
30 77,
z Omm.
S
N
Transverse section of chick of 2} days (Bielschowsky preparation), showing amid
the mesenchyme cells isolated neuroblasts (V.C.) between spinal cord (S.C.)
and aorta (Ao.).
possessing distinct nervous processes. The neuroblast has a large
oval nucleus, and its processes are directed towards the aorta.
C. Relations with the Ventral Roots.
Whilst all investigators are not agreed as to the origin of
sympathetic cells from the neural crest before its differentiation
into the dorsal root ganglia, most are agreed that the ganglia
when formed give origin to the sympathetic cells. Held (7) goes
so far as to derive the whole of the sympathetic ganglia from the
dorsal root ganglia. [is evidence is based on the examination
of Selachian material, but he is so satisfied with those observa-
tions that he applies his deductions to the case of chick embryos,
although he confesses that he has not been able to discriminate
the roots satisfactorily in them.
Kuntz (11, 12), on the other hand, attaches by far the greater
importance to the ventral roots as the path by which the cells
migrate, a view which he supports by both morphological and
experimental evidence.
SYSTEM AND SUPRARENAL BODIES IN THE SPARROW... 747
In collaboration with O. V. Batson he experimented on chick
embryos. By electrolysis they destroyed the dorsal part of the
neural tube of the embryos, and in four successful cases found
that the sympathetic primordia were formed to a greater or less
extent. Kuntz later carried out further work on the same lines
and arrived at similar results. Of similar experiments carried
out on tadpoles, by actual dissection, he writes: ‘“ As far as the
study of the development of the sympathetic trunks was pursued
in these embryos the findings.corroborated those in the operated
embryos of the chick.”
The experimental investigations of E. Miiller and Sven Ingvar
(13), on the other hand, support the conclusions of His and Held.
Text-figure 5.
Transverse section of Sparrow embryo 11 (= chick about 78 hiys.), showing cells
leaving the spinal cord by the ventral nerve root.
D.R.G. Dorsal root ganglion. 2.C, Migrating cells. S.C. Spinal cord.
V.R. Ventral root. (Sp. 11, 4, 2, 17.)
‘They carried out their experiments on Amphibian material. By
destroying the ventral part of the neural tube they found that
the sympathetic was still formed, wnile the removal of the dorsal
portion resulted in its failure to develop. They thence conclude
that the primordia of the sympathetic nervous system arise from
the spinal ganglia. -
The conflicting results obtained by these workers illustrate the
pitfalls inherent in the experimental method as applied to this
particular problem. ° cae
Our observations certainly do not support Held’s view as to
the exclusive origin of the sympathetic system from the dorsal
748 A. SUBBA RAU AND P. H. JOHNSON ON THE NERVOUS
root ganglia, but they lead us to attach somewhat more im-
portance to that source of origin than Kuntz would seem to
allow.
Text-fig. 5 shows the Taira root of a Sparrow embryo, No. 11
(= 78- hr. chick), The ventral roots are well developed, and
the one figured shows a group of cells close to the origin of the
root, and evidently i in course of migration, ‘That the migration
is outwards seems clear from the. observation of a liane stage,
such as is reproduced in text-fig. 6 of Sparrow 51 (= 84-hr.
chick). Here we find some of these tnierene cells leaving the
ventral root and making their way to the dorso-laters al angle of
the aorta, at which point they join the previously described
Text-figure 6.
0
toe OG
“itn
dé. ~
OX 2"
Transverse section of Sparrow 51 (= 84-hy. chick), showing cells leaving ventral
root in the direction of the dorso-lateral angle of the aorta.
O.S. Cell strand, S.C. Spinal cord. V.R. Ventral root.
(Sp. 61, 3, 2, 4.)
sympathetic cells, and with them form diffuse masses of cells,
which are the primordia of the primary sympathetic chains.
At this stage the chains are not formed, but the diffuse cell
masses occur metamerically at intervals corresponding with the
somites.
This inigration of cells along the ventral roots, and thence to
the dorso- lateral angle of the aorta, is visible to a greater or less
extent in all the embryos examined at about this stage (= 3-34-
day chick). Owing to the curvature of the embryo it is obvious
that the occurrence of a transverse section passing along one of
these migratory paths is quite fortuitous, though it can 1 always
SYSTEM AND SUPRARENAL BODIES IN TUE SPARROW. 749
be shown by reconstructions from consecutive sections, a method
which Held employed for some of his illustrations. ‘Text-fig. 7,
however, shows a fortunate section of a chick embryo (D. 4 of 34
days), in which the curved track of darker cells is practically
continuous, thus furnishing a striking picture of the path of
migration. This path seems to be markedly affected by the
Text-figure 7.
a Xf} re
Pie AGE,
Myer BO §
pie a He a, i
ATE)
tin
Transverse section of chick. embryo (3% days, D 4), showing an almost continuous
strand of sympathetic nerve cells from the point of exit of the ventral
nerve root to the dorso-lateral angle of aorta. i
Ao. Aorta. C.S. Sympathetic cell strand. Néc. Notochord. Mzyt. Myo-
tome. S.C. Spinalcord. Sym.C. Group of sympathetic cells. V.R. Ventral
root. ;
segmental arteries and veins, for it is between these vessels that
the sympathetic cells move to their primary destination (see text-
fig. 3). It is possible that the intersegmental region offers a
path of least resistance to this migration.
A track of nerve fibres in the same position is clearly marked
750 A. SUBBA RAU AND P. H. JOHNSON ON THE NERVOUS
in silver preparations, such as that shown in text-fig. 8, and it
seems probable that this path would agree with that of the
migrating nerve cells.
Text-figure 8.
ny
May,
Transverse section of chick (m-4}) from a Bielschowsky preparation, showing nerve
fibrils pursuing the same course as the sympathetic cell strand in fig. 7.
The mesenchyme and other cells are much distorted, but some of those near
the dorso-lateral angle of the aorta show evidence of being sympathetic in
character.
Ao. Aorta. S.C. Spinal cord. V.R. Ventral root fibres. Sym.C. Sympa-
thetic cells. Néc. Notochord. Né¢r. Nerve fibres in the track of migrating
sympathetic cells.
While it appears that many of these cells are sympathetic in
nature, the possibility must not be excluded that some of them
will develop into the sheath cells of Schwann.
D. The Primary Sympathetic Chain.
In their earliest stage the primordia of the sympathetic
chains consist of two parallel series of cell groups situated at the
dorso-lateral angles of the aorta. ‘These are arranged meta-
merically and show a more advanced stage of development in the
anterior thoracic region, Knsuing stages show their gradual
t
SYSTEM AND SUPRARENAL BODIES IN 'THE SPARROW. (51
consolidation by the dispersed cells gathering into compact masses,
and the simultaneous growth of longitudinal cellwar strands, which
unite them into continuous moniliform chains. Silver prepara-
tions show scattered fibres among these cells. A graphic recon-
struction of the consolidated cord in the anterior thoracic region
at this stage is shown in text-fig. 9 A.
So far as we have been able to observe, the cells constituting the
chain at this stage do not show any differentiation of chromaftin
cells among them, Their’ nuclei are oval, their cytoplasm darkly
stained and full of minute granules, cell boundari ies are faint, or
may still be absent. We have constantly observed the presence
of clear spaces surrounding the groups of sympathetic cells near
the dorso-lateral angle of the aorta. Kuntz has noticed these
clear spaces. They may be spaces filled with lymph during life,
or they may be due to shrinkage phenomena. We have followed
the development of the cords by means of graphic. reconstruc-
tions from a series of Sparrows, separated by very s slight differ-
ences of development, and the result is striking. The series of
reconstructions is shown in text-fig. 9.
We have chosen for purposes of reconstruction the anterior
thoracic region of the embryo, where its development is most
advanced. ‘Text- -figs. 9 A & B are reconstructions of the primary
syinpathetie nervous system of Sparrows 7 and 2 (= 78-hr.
chick). They show that at this stage the sympathetic chains in
the anterior thoracic region consist of two approximately parallel
cords, formed by masses of cells united by their connecting strands.
Anteviorly the strands are much thicker, but not sufficiently SO
to hide the metamerism, which corresponds exactly with that
of the dorsal root ganglia. In both the text-figures, but more
markedly in text-fig. 9 B, it will be seen that the ‘swollen parts of
the cord send out lateral processes. These processes do not run
in the plane of the parallel cords, but are directed dorso-laterally.
The apparent differences in the directions to which they point
are due to the curvature of the embryo.
Text-fig. 9 C is from Sparrow 6 (= 84-hr. chick). The monili-
form character of the strands is more marked and gives the
impression that the swellings have increased at the expense of
the connectives. ‘The distances between the ganglionic swellings,
e.g. in text-figs. 9 A & 9C, areroughly equal, rout the connectives
in the latter have become more slender, a result which could he
attained either by a process of degeneration, which seems im-
probable at this early stage, and of which no evidence was
obtained in the sections, or by migration of their elements into
the ganglionic swellings.
Similar migration seems to have taken place into the lateral
processes, which are now connected by broad bases to the primary
ganglia, and taper off to a point dorso-laterally to the cords.
This certainly suggests that their growth is outward from these
ganglia, a view supported by oe iD -equent occurrence of mitoting
52 A. SUBBA RAU AND P. Hi. JOIUNSON ON THE NERVOUS
. Text-figure 9.
Graphic reconstruction of the sympathetic primordia in the anterior thoracic region
of Sparrows. ‘The cord is viewed from the ventral surface and the meta-
merically arranged dorsal root ganglia are indicated by dotted lines. Tho
lateral projections are not in the same plane as the cords, and allowance must
be made for the necessary foreshortening in their projection.
A, Sparrow 7 (= 78-hr. chick). Cords continuous with slight swellings
and short dorso-lateral processes.
LB. Sparrow 2 (= 78-hr. chick). Cords continuous with slight swellings
and short dorso-lateral processes.
C. Sparrow 6 (= 84-hr. chick). Sweilings and processes much more pro-
nounced.
Notice four mesial commissures and the proliferation at the posterior
end which is the region where there exists a close relation between the
sympathetic and the suprarenal bodies.
SYSTEM AND SUPRARENAL BODIES IN THE SPARROW. 753
cells in both ganglia and processes. The posterior part of the
chain in text-fig. 9C still shows considerable dispersion of the
sympathetic elements, and it is from this region that contribu-
tions are sent to the suprarenal bodies. In the anterior part the
chains ave united by transverse commissures.
Text-fig. 91) represents a reconstruction of Sparrow 13
(= 96-hr. chick). The distinction between ganglia and con-
nectives is much more marked, and an important change is also
visible in the lateral processes. The four anterior pairs are still
united by a broad base to the primary ganglia, but instead of
tapering to a point they are now of practically the same thick-
ness throughout. In the three posterior pairs the bases show
signs of constriction, while the distal portion is marked by a
eranio-caudal elongation, Now when we examine the cords
themselves we find that they are thinnest just in the region
where the lateral processes possess these cranio-caudal exten-
sions, while in the anterior region, where the processes have
not yet reached this stage, the cords remain of considerable
thickness. We are again driven to the conclusion that the
cranio-caudal extensions are formed at the expense of the
primary cords.
Text-fig. 9 KH, from Sparrow A (= 53-day chick), shows a con-
tinuation of this development. ‘The lateral processes are now
solid masses, united to the primary chain by short connectives,
and all of them show cranio-caudal extensions, the caudal being
more marked. The correlation between the increased size of the
iateral process and the thinness of the primary sympathetic chain
is again very obvious.
The solid lateral masses are now situated in close relationship
with the spinal nerves, being located just mesial to the junction
of dorsal and ventral nerve roots. They may now be termed the
primordia of the secondary sympathetic ganglia. Subsequent
growth of their cranio-caudal extensions leads by their union to
the formation of the definitive sympathetic chain (secondary
sympathetic chain).
D. Reconstruction of Sparrow 18 (=96-hr. chick). Ganglia and comnec-
tives more marked. Lateral procosses thickened. Posterior ones con-
stricted to some extent.
1. Reconstruction of Sparrow A (54-day chick). Lateral processes now
solid masses united to the primary chain by short connectives. All
show cranio-caudal extensions.
. Reconstruction of Sparrow 60 (= 6-day chick). Cranial-caudal extensions
meet to form secondary sympathetic chain.
G. Reconstruction of Sparrow B (=older than Sparrow 60). Secondary
sympathetic chain formed. Connectives between primary and secondary
degenerating.
ry
C. Connective. D.R.G. Position of dorsal root ganglion. G.S. Gan-
glionic swelling. P.S.C. Primary sympathetic cord. M.C. Median connec-
tive. C.L.P. Constricted lateral process. 2.P.C.C. Lateral processes with
cranio-caudal extensions. J. Junction of cranio-caudal extensions. S.S.C.
Secondary sympathetic chain. I22G. Median ganglia. D.C. Degenerating
connective.
154 A, SUBBA RAU AND P, If, JOINSON ON THE NERVOUS
KE. The Secondary Sympathetic: Nerve Cords.
An interesting example of this union is shown in text.-fig. 9 F,
a reconstruction of Sparrow 60 (= about 6-day chick). We here
see that at two points the secondary chain is actually completed,
while in other places marked caudal prolongations are apparent.
‘The completion of the formation of these definitive sympathetic
chains is found at a stage corresponding to that of the seven-day
bal Text-figure 10. (a & b). d.
Diagrammatic figures showing relations of sympathetic cord to surrounding tissues
as seen in transverse sections of Sparrow 13 at points marked a-a, B-2 in
fig. 9 D.
10a. Passes through a ganglionic swelling of the sympathetic chain (Sym.C.).
The ganglionic swelling lies dorso-lateral to the aorta (D.A.) and between
the segmental artery (Seg.A.) and segmental vein (Seg.V.). There is a dorso-
lateral projection pointing towards the position of the dorsal root ganglion.
106. Passes through a connective (P.8.C.) between two ganglionic swellings.
The two sections are ouly 56 apart.
D.A. Dorsal aorta,’ D.R.G. Dorsal root ganglion. yt. Myotome.
‘Nte. Notochord. Sp N. Spinal nerve.
chick. A reconstruction of this stage is shown in text-fig. 9G,
Sparrow B. | ; ae
The condition of the primary sympathetic chain at this stage
presents very interesting features. The two cords are now much
diminished in size and, gradually approaching one another, are
for some distance almost in-contact. ‘Their moniliform character
is lost, except that in the thoracic region we find three large
SYSTEM AND SUPRARENAL BODIES IN THE SPARROW, 755
median ganglia, partly or completely fused (text-fig. 11). These
ave joined to the secondary ganglia by connectives of nervous
fibres. We have also found these median ganglia in a much
more advanced embryo. The ganglia lie above the aorta imme-
diately ventral to the vertebral column, the centra of which are
in process of development. On either side of them there occur
mesodermal masses, such as give rise to muscle in the adult, in
Text-figure 11.
tebe G8 (6
5 R99 ag.
c Uy ql! :
Transverse section through Sparrow B at point marked a~a in fig. 9 G. The secon-
dary sympathetic ganglion is shown at Sym.C. (2) and is connected with the
median fused ganglia of the primary sympathetic (Sym.C.) (1) chain,
Ao. Aorta. Cp. Capillaries associated with primary sympathetic.
(Sp. B 9, 4, 7 and 8.)
this case probably to muscles attached ‘to the vertebrae. The
connectives between the ganglia are associated with a rich
capillary plexus dorsal to the norta. «
Posterior to these ganglia the primary sympathetic cords
diminish in size, until in the abdominal region it is no longer
possible to trace them. 9 2
Tyvansverse sections throw: light on the processes of develop-
ment above described. ‘Text-figs. 10a & 106.show transverse
756 A. SUBBA RAU AND P. H. JOHNSON ON THE NERVOUS
sections of Sparrow 13 at the points marked a~a, B—-B in text-
fig. 9D. The sections are only 56 » apart, but the difference is
very marked, The section shown in text-fig. 10 @ passes through
the dorso-lateral processes of the primary sympathetic chain, and
shows their relations to the surrounding structures. They lie
between the segmental artery and vein, and extend dorso-
laterally from the aorta to a point about halfway towards the
-Text-figure 12.
Nec.
piel X)-0'3-0
Ga in OU... Bb
‘Transverse section through Sparrow B at point marked =n in fig. 9G. The secon-
dary sympathetic chain (S.Sc.) is closely apposed to the spinal nerve.
origin of the ventral roots. ‘he section in text-fig. 106, on the
other hand, shows the primary sympathetic as a comparatively
small stratid close to the aorta, ‘This section passes through the
myotome and the dorsal root ganglion, while that shown in text-
fig. 10 @ passes between the somites.
Lext-figs, 11 & 12 drawn from sections of Sparrow B, at points
SYSTEM AND SUPRARENAL BODIES IN THE SPARROW. 757
marked A-A and B-8 in text-fig. 9G call for some remark. Text-
fig. 12 shows the primary and secondary sympathetic chains
apparently quite disconnected. ‘Lhe primary has its usual posi-
tion at the dorso-lateral angle of the aorta, the secondary is
situated close against the spinal nerve, and lateral to the noto-
chord. Text-fig. 11, however, shows a section through the con-
nective between primary and secondary chains. ‘The connective
at this stage is composed chiefly of nerve fibres, but along them
there are scattered cells, which are probably destined to form
Sheath cells. The two primary ganglia in this section will be
seen to have completely fused.
F, Rami Communicantes.
The foregoing description of the development of the sympa-
thetic nervous system will explain how there has arisen a con-
siderable amount of confusion as to the rami communicantes in
birds, for it will be seen that in this group there are no less than
three distinct sets of connections associated with the central
nervous system and the outlying sympathetic. These are :—
1. The strands of nerve cells which give rise to the primary
sympathetic cords. A
2. The constrictions of the dorso-lateral processes by which
the primary cords are attached to the secondary ganglia.
3. The definite connections by which the secondary ganglia
become attached to the spinal nerves. . -
Strictly speaking the term ramus communicans should he
restricted to the last of these three groups, but previous papers
and illustrations show that all these three forms hare been thus
designated. Ganfini (5) goes so far as to use the terms primary,
secondary, and tertiary rami communicantes for the three classes
of connections, but in view of their significance in neurology it
would seem preferable to apply the name only to those fibres
by which the definitive sympathetic is linked up with the mixed
spinal nerve. As the definitive ramus does not arise till about
the stege corresponding to that of a seven-day chick, it is evident
that any connection between the sympathetic and the central
nervous system, existing in earlier stages, must be looked upon
ag an example of either the first, or the second, of the types of
connections above mentioned. As the history of their develop-
ment shows, the connections of groups 1 and 2 have only a
transient existence, with the possible exception of the connec-
tions in certain plexuses.
The three types of connections are characterized more or less
definitely by their morphology. ‘those of the first group are
formed of discontinuous cells, though silver preparations show
that scattered nerve fibres occur in them. Connections of the
second type, on the other hand, are formed by compact masses
of cells, especially at their first appearance. In the. course of
their development they become elongated and may be narrowed
Proc. Zoor. Soc.—1923, No. L. 50
758 A. SUBBA RAU AND P. H. JOHNSON ON THE NERVOUS
down almost to a single row of cells. In their final stages their
cellular character disappears by migration of the cells to the
secondary chain, and a fibrous connection is left (text-fig. 11).
With the exception of certain connections which are maintained
with the chief plexuses of the trunk, these also ultimately dis-
appear, usually after the appearance of the definitive ram.
These last are from the beginning purely fibrous. ‘Their develop-
ment consists merely of a growth in length, as the secondary
sympathetic ganglion moves away from its early position at the
point of union of dorsal and ventral roots. The range of move-
ment varies in different regions, being very small in the neck.
Thébault (14) figures the rami communicantes of Corvus as some-
times quite distinct in the neck region, but also describes a fusion
of the sympathetic ganglion with the spinal nerve, and remarks
that all the intermediate stages may be found. A ramus com-
municans can always be found in our material by microscopic
examination.
That the sympathetic ganglion, while making its own contri-
bution to the mixed spinal nerves, carries with it fibres from
both roots, has been abundantly proved by previous observers.
Rossi (16), in his recent paper on the afferent paths of the
sympathetic system, demonstrates by direct anatomical observa-
tions, that cells occur in the dorsal root ganglia which send their
processes into the rami communicantes. These he regards as the
cells of origin of Kdlliker’s fibres, and considers them to be
afferent sensory neurones.
G. The Suprarenal Bodies.
Our observations lead us to agree with those who ascribe to
these bodies a dual origin: e.g. Balfour, Poll, Mitsukuri, Gia-
comini, and Goormaghtigh.
In Sparrow 7 (= 78-hr. chick), text-fig. 13, there are already
signs of the grooves in the ceelomic epithelium from which the
cortical portion of the suprarenal bodies will arise. ‘These
grooves are situated on either side of the mesentery and close to
it. Laterally to them will be found the internal Wolffian vein
and the mesonephros, and in certain cases the external glome-
rulus of the pronephros can also be seen. In text-fig. 14 these
relations are clearly visible, and the proliferation of cells has
already started at the region described. The cells of the coelomic
epithelium are cubical with large nuclei. Mitotic figures are
numerous and the products of division appear to be moving
inwards. In Sparrow 8, text-fig. 15, the proliferation of cells
becomes more marked, and a ridge is formed internally. ‘These
cortical primordia appear to be metamerically arranged. In
Sparrow 11, text-fig. 16, the cortical primordia are very clearly
marked, but still closely attached by a broad base to the ccelomic
epithelium. Groups of cells appear to be moving towards their
later position between the aorta and the mesonephros.
SYSLEM AND SUPRARENAL BODIES IN THE SPARROW. 759
Sparrow 5, text-fig. 17, shows the separation completed ; the
primordia are now compact and contain a few cells from the
primary sympathetic chain, which have migrated down the sides
Text-figure 13.
Transverse section through Sparrow 7 (= 78-hr. chick), showing the formation of
the coelomic groove (C.G.).
C.H. Celomic epithelium. D.A. Dorsal aorta. 2f.N. Mesonephros.
M.S. Mesentery. JI.W.V. Internal Wolffian vein. (Sp. 7, 5, 3, 3.)
Text-figure 14.
M4.N, : yh
Transverse section of Sparrow 8, showing the deepening of the groove (0.G.) and
proliferation of cells (P.C.) to form the primordium of the suprarenal cortex.
Letters as in fig. 13.
of the aorta. It is this contribution from the sympathetic chain
which eventually forms the medullary tissue of the suprarenal
body.
5O*
760 A, SUBBA RAU AND P. Il. JOHMNSON ON 'THHK NERVOUS
C.P M.S.
The suprarenal is now invading the internal Wolffian
Transverse section of Sparrow 11, showivg a further development of
Letters as in fig. 13.
Yo)
rok
2
lant
2 DOK
én ae
“Ft RAG O10) |
e @ (c@ S
i Oe Qs S
any AS NS
al Ach @e @ S
2G 95) 8
6
oO
=
a
a)
ct
op
=)
i
3
wm
Lon
~
oO
sl
Text-figure 15.
Transverse section of Sparrow 8, showing grouping of cells to form the
primordium of the suprareual cortex (C.P.).
vein.
SYSTEM AND SUPRARENAL BODIES IN THE SPARROW. 761
Text-figs. 18 & 19, from Sparrow 58, show the relations of the
sympathetic and the supravenal body. The latter now surrounds
the internal Wolffian vein, while the sympathetic chains dorso-
lateral to the aorta are largely developed, and give rise to masses
of cells which pass from them to the cortical primordia. The
point of entry of these cells seems usually to be on the mesial
face of the suprarenal body (text-tig. 18), but in text-fig. 19
there is apparently a dorsal entry.
Text-figure 17.
Transverse section of Sparrow 6, showing first signs of invasion ot cortex by
medullary colls (J£.C.) from the primary sympathetic. Lettering as in text-fig. 13.
It is noteworthy that these contributions of the sympathetic
to the suprarenal all arise from the primary sympathetic cords,
which apparently proliferate them at about the same time as
they are giving rise to the definitive cords, by the outgrowth
of dorgo-lateral branches, as described above. We have not
attempted a detailed description of the changes which occur in
the development of the suprarenal bodies, as these have been
Text-figures 18 & 19.
76
A. SUBBA RAU AND P.
Hi. JOHNSON ON THE NERVOUS
smpathetic to suprarenal body.
Transverse sections of Sparrow 53, showing contributions of primary sy
In fig. 18 the contribution is dorsal while in fig. 19 it is mesial.
(Sp. 53, 4, 4, 3. Sp. 53, 4, 5, 4.)
SYSTEM AND SUPRARENAL BODIES IN THE SPARROW. 763
admirably set forth by Goormaghtigh in his recent paper. The
only point on which our observations lead us to differ from him
is as to the origin of the medullary substance. This is derived,
according to our observations, from the primary sympathetic
cords and not, as Goormaghtigh maintains, from the mesodermal
cells of the sclerotome.
HL. Discussion.
Since Irancis Balfour (2) in 1877 described the development
of the sympathetic nervous system in the Selachii, many theories
have been put forward supporting, or refuting, his statement as
to its origin. Without going deeply into the history of the con-
troversy, we may summarize the chief suggestions as follows :—
A. Origin ectodermal.
a. From the medullary tube.
b. From the spinal nerve roots.
ec. From the dorsal root ganglia only.
d. From ventral roots chiefly.
B. Origin mesodermal.
Differentiation in sitéw.
C. Mixed origin.
From the medullary tube and sclerotome.
Supporters of the mesodermal origin of the sympathetic nervous
system have based their conclusions chiefly on the presence of the
sympathetic primordia, at the dorso-lateral angles of the aorta
and the carotids, before the appearance of the nerve roots.
Although we agree with them in this observation, we are yet of
the opinion that the cells in question have not differentiated
in situ, but that they have been proliferated from the neural
crest (see text-fig. 1).
We have not found a trace of support for the interpretation
recently put forward by Dart and Shellshear (4), who state that
“4 mesodermal origin of these neuroblasts (2. e. neuroblasts in
relationship with mesodermal structures) must therefore be postu-
lated and is demonstrable.” All nerve fibres which we have
observed could either be traced from the central nervous system,
or, in certain cases, from free cells closely associated with it, and
in no case occupying a position such as would support their
view.
With regard to Goormaghtigh’s (6) contention for a mixed
ovigin, we may state that we have kept a careful look out for the
strands, which he describes as arising from the sclerotome and
contributing to the formation of the primary sympathetic system,
and have been entirely unsuccessful in our search. The paths of
the migrating cells destined to form the sympathetic primordia
certainly pass close to the sclerotomes, but the boundary between
the two structures in our material has always been clearly
marked, and the cells on either side of that boundary sutfticiently
764 A, SUBBA RAU AND P. H, JOILNSON ON TILE NERVOUS
differentiated to distinguish them. It is unfortunate that the
photographie illustrations, which Goormaghtigh puts forward in
support of his view, are far too indistinct to serve as evidence
for it.
_ We are thus led to support the entirely ectodermal origin of
the sympathetic nervous system, and from the observations
already given, it will be seen that we do not confine that origin
to any one part of the central nervous system. ‘The statement
by experimental physiologists that the sympathetic contains both
afferent and efferent components, while it does not afford a proof
of a diverse origin, yet supports the possibility of such being the
case. We have seen that some cells originate very early ‘from
the neural crest, and we are convinced that both dorsal and
ventral roots contribute towards the formation of the sympathetic
primordia.
The observations of His (8) and Held (7) support the origin
from the dorsal root ganglia. These writers describe the
primordia as arising by cells, migrating from the spinal ganglia
by way of the spinal nerve, from which they diverge a little
beyond the point of their junction, and thence make their way
to the dorso-lateral angle of the aorta. The sympathetic cells
thus leave the spinal nerves at a point subsequent to the union
of dorsal and ventral roots, and so may originate from either of
those sources. ‘The assumption that all have arisen from the
dorsal root ganglion is based simply on an analogy with Selachian
material, and has no foundation on observations of avian embryos.
The descriptions of Held and His lead one to the supposition
that this process is continuous throughout the development of
the sympathetic; this, however, is quite contrary to the results
of our investigations. If the sympathetic ganglia were derived
directly from the spinal nerve, one would naturally expect, the
cells composing them to bear a distinct resemblance to those
from which they are said to originate; but this we do not find
to be the case. ‘The cytological characters of the dorsal root
ganglia, and of the cells of the ventral horn of grey matter, are
so different from those of both primary and secondary sympa-
thetic ganglia, that there seems no possibility of one being
derived immediately from, the other (see text-fig. 20).
Ouwr description, as given in §H, of the morphological dleve-
lopment of the sympathetic, allows ample opportunity for this
differentiation to arise; for we discover that though the ganglia
of the dorsal roots and the definitive sympathetic ganglia lie
close to one another, yet the latter have had oppor tunity to
undergo considerable changes in the course of their extensive
migration, jirst, from the neural crest, &c., to the aorta, and,
secondly, from the aorta to near the spinal ¢ ganglion.
_ ‘The reasons for .this complicated migrator y movement have
yet to be discovered, but the evidence for it is, we think, sufti-
ciently definite. ‘The situation of the definitive sympathetic
in the vertebral canal in birds,was pointed out as long ago as
SYSTEM AND SUPRARENAL BODIES IN THE SPARROW. 765
1810 by Tiedemann, but the occurrence of a primary chain in a
position analogous to that of the definitive sympathetic in other
vertebrates was not discovered till many years later by His.
Subsequent writers have all supported his findings, but hitherto
ull have failed to discover any genetic continuity between the
two formations. Ganfini (5), in his paper on the sympathetic in
birds, seems to have observed the relations more successfully
than his predecessors, but he failed to discover the origin of the
secondary ganglia from the dorso-lateral processes of the primary
ganglia. He follows his predecessors in deriving them imme-
diately from the dorsal root ganglia, for he writes: “ Alla forma-
zione del cordone simpatico primario segue la formazione di un
Text-figure 20.
>
5
at
>
<
S
EN |
nea
sS
Veoees = Say
Transverse section of Sparrow 69 (= 8-day chick), showing the differentiation of
cells of the dorsal root ganglion (D.R.G.) and of the secondary sympathetic
ganglion (Sy.G.). Sp.N. Spinal nerve.
cordone simpatico secondario. Questo é formato da una serie di
gangli situati ognuno a livello della parte del tutto prossimale
del nervo spinale ed originati da neurociti derivanti esclusiva-
mente dai gangli spinali. . . . Da ogni ganglio simpatico
secondario si diparte un ramo che dirigendosi ventralmente e
medialmente connette il cordone simpatico secondario col pri-
mario.” From the last statement he appears to have observed
connections proceeding from the secondary to the primary
ganglia, while our graphic reconstructions show exactly the
opposite. !
The nearest approach to correct observation seems to have
been made by Miss Williainina Abel (1), though curiously enough
she attached little importance to it. In her paper we read:
766 A. SUBBA RAU AND P, H. JOHNSON ON THE NERVOUS
“At 144 hours the permanent ganglia appear as small clusters
of nerve cells, lying close to the mixed spinal nerves and con-
nected to them by fibrous rami conumunicuntes. hese ganglia
increase rapidly in size and are formed largely from the sympa-
thetic cells which have accumulated between the spinal nerves
and the temporary (t.e. primary) sympathetic chain. Some of
the cells of the temporary sympathetic chain are also incorpo-
rated in the permanent (7.e. secondary) ganglia. Besides these
the permanent ganglia are formed from the cells which have
migrated directly to them along the spinal nerves. That portion
of the temporary sympathetic chain which is not incorporated in
the ganglia of the permanent chain gradually atrophies and
disappears.”
Our observations therefore agree with Miss Abel’s in so far as
they confirm the importance of the contribution from the primary
to the secondary chain. They differ, in assigning a definite form
to the accumulations of which she speaks, and in demonstrating
definite paths by which this contribution is effected, i.e. the
dorso-lateral prolongations from the primary ganglia. More-
over, from the marked differentiation of the spinal nerve cells
from those of the sympathetic ganglia already described, we
question the existence of small clusters of nerve cells lying close
to the mixed spinal nerve, in so far as she derives them from the
spinal nerve directly. At the time when such clusters do occur
they will be found to consist of small cells with dark nuclei
containing many granules. We cannot see how such a meta-
morphosis should occur in the short passage from the dorsal
ganglia, all the cells of which are large, with pale nuclei and
containing many granules. It seems more probable that the
whole detinitive sympathetic ganglion arises from the primary
sympathetic chains by migratory cells, and by active mitosis,
which latter can always be observed in these ganglia.
As to the fate of the primary sympathetic, Miss Abel is in
accord with previous writers, all of whom express the opinion
that in the course of further development it atrophies and dis-
appears. Our material does not attain a sufliciently advanced
stage for us to express a definite opinion on this question, but
there are clear evidences of its presence in the oldest stage we
have observed, Sparrow 69 (= 8-day chick). The definitive
sympathetic chain in the cervical region of this embryo runs in
the vertebrarterial canal. Between the vertebrie of the hinder
part of the neck, small bundles of fibres emerge and run ventrally,
but closely apposed to the cartilaginous vertebral column; these
fibres enter large and distinct medial ganglia, lying just under
the vertebral column, and above the carotids. In Sparrow B, a
little younger, three such medial ganglia are specially noticeable
(see text-fig. 11). Longitudinal fibres run from these ganglia to
the tissues lying between them, which are highly vascular, being
traversed by a capillary plexus linked up with the carotids. The
ultimate destiny of these ganglia awaits further investigation.
SYSTEM AND SUPRARENAL BODIES IN THE SPARROW. 167
Apart from them the primary cords are so reduced, by the
migration above described, that they may be considered to be
wholly absorbed by the definitive sympathetic nervous system.
It should be stated that the primary sympathetic chain contri-
butes also to the formation of ceeliac and other ganglia, thus the
degeneration of the primary sympathetic chains, described by
previous writers, should rather be termed a migration from the
primary location.
We are unable to assign a cause for this migration of the
primary sympathetic chains from the dorso-lateral angles of the
aorta to their definitive position, as secondary sympathetic chains
in the vertebrarterial canal. Thébault, in his description of the
sympathetic system in adult birds, suggests that the long thin
neck of the bird renders the sympathetic liable to lesions; hence
he explains its inclusion in the vertebrarterial canal. If this be
the case, we may regard the definitive location as an adaptive
modification. It is perhaps worth notice that the reptile in
which primary and secondary sympathetic chains have also been
described, viz. the tortoise, is also endowed with a Jong and
flexible neck.
In conclusion, we would express our gratitude to Professor
J. P. Hill, F.R.S., for his generosity in supplying us with such
an abundance of excellent material and for much assistance in
the course of our work and in the preparation of this paper.
We are indebted to Professor D. M. S. Watson, F.R.S., for
friendly criticism and for aid in final preparation for the press.
I. Summary.
1. The primary sympathetic primordia originate from the
spinal cord, and are hence ectodermal in origin.
2. The primordia of the ganglia of the primary sympathetic
chain are laid down metamerically at inter-somitic points.
3. The constituent cells of the primordia are derived from
a. The neural crest ;
b. The dorsal root ganglia ;
c. The neural tube by way of the ventral root.
4. The primordia extend cranially and caudally to form a
continuous chain on either side.
5. The primary ganglionic swellings extend dorso-laterally.
The free ends of these extensions enlarge and becoming sepa-
rated from the primary ganglia give origin to the definitive
sympathetic ganglia.
6. As in the case of the primary ganglia these definitive ganglia
extend cranio-caudally and unite to form a continuous chain.
7. The primary sympathetic gives origin to
a. The secondary sympathetic chains ;
b. The chromaffin cells of the medulla of the supra-
renal body ;
ce. The celiac and other ganglia ;
d. The unpaired ganglia in the cervical region.
768 NERVOUS SYSTEM AND SUPRARENAL BODIES IN THE SPARROW.
K. bibliography.
A very full bibliography of the development of the sympa-
thetic nervous system up to the year 1905 will be found in
Hertwig’s ‘ Handbuch der vergleichenden und experimentellen
Entwickelungslehre,’ Band ii. ‘Veil 111.
More recent contributions to the literature are given by
Goormaghtigh and also by Da Costa in the papers cited “below.
The following are the works to which reference is made i
this paper :—
1. Asset, W.—‘ Further Observations on the Development of the
Sympathetic Nervous System in the Chick.” Journal
of Anatomy and Physiology. 1912
2. Batrour, I’.—‘‘'The Development of Hl: ismobranch Fishes.”
Journal of Anatomy and Physiology. 1876-78.
3. Da Cosra, A. C.—‘‘ Origine et développement de Vappareil
surrénal et du systéme nerveux et sympathique chez les
Cheiroptéres.” Mem. Soc. Portug. Sci. Nat. iv. 1917.
4. Darr & Suaptisnear.—‘* A New Interpretation of the Mor-
phology of the Nervous System.” Abstract. Anat.
Record, vol. xxi. p. 54.
5. Ganrint, C. “Lo sviluppo del sistema nervoso simpatico
negli uccelli.” Arch, Ital. del Anat. 1916-17.
6. Goormaaurien, N.—‘‘ Organogenése et Histogentse dle la cap-
sule surrénale et du plexus ceeliaque.” Arch. de Biol.
1921.
7. Hexip, H.—Die Entwickelung des Nervengewebes. 1909.
8. Ths, W., Jun.—“ Die Entwickelung des Herznervensystems
bei Wirbeltieren.” Abhand. der kon. siichs. Gesell.
der Wiss, 1893.
9. His, W., Jun.— Ueber die Entwickelung des Bauchsympa-
thicus beim Hiihnchen und Menschen.” Arch. fiir
Anat. und Phys. 1897.
10. Kunrz, A.—‘“ Development of the Adrenals in the Turtle.”
Amer. Journ, of Anat. vol. xili.
11. Kunrz, A.—“ Development of the Sympathetic” in Mam-
mals, 1910; in Birds, 1910; in Fishes, 1911; in
Amphibia, 1911; in Man, 1920. Journal of Compara-
tive Neurology.
12. Kuntz, A.—‘* Bxperimental Studies on the Histogenesis of
the Sympathetic Nervous System.” Journal of Com-
parative Neurology. 1922.
13. Mtiivr, W., & Sven Incyar.—“ Ueber den Ursprung des
Sympathicus bei den Amphibien.” Upsala, Likaref.
Forh, xxvi. 5-6.
14. Tiripaunr, M. V.i—‘ Etude des rapports qui existent entre
les systémes pnewnogastriques et sympathiques chez Jes
viseaux.” Ann. des Sciences Naturelles. 1898.
15. Keren & Mani.—'l'ext-book of Human Hmbryology.
16. Ross1, O.—‘“ On the Afferent’ Paths of the Sympathetic
System, &e.” Journal of Comparative Neurology.
1922.
iF Wi i
I.
1923, ABRAHAM, PI.
Th N30
ee
John Bale,Sons & Danielsson. L'?
Ss.
la Mare Norri
Photos. F. de
Sp-n.
NEST OF LIPHISTIUS MALAYANUS
A NEW SPIDER OF THE GENUS LIPITISTIUS. 769
r
37. A new Spider of the Genus Liphistius from the Malay
Peninsula, and some Observations on its Habits.. By
H. C. Apranam, F.Z.8., F.L.S8.
[Received May 25, 1923: Read October 23, 1923.]
(Plate I.; Text-figure 1.)
I have recently received, through the courtesy of Mr. F. de la
Mare Norris, of the Department of Agriculture, Federated Malay
States, a specimen *-of the female of a large species of Liphistius,
together with the trap-door and portion Of the silk lining of its
ear, photos of the burrow in situ, and notes of a number of,
observations on the habits of the spider.
In the preamble of a former paper t I mentioned Ihaing 6 seen
2 photo of the present species, and, at that time, I was under the
impr ession that it would turn out to be LZ. birmanicus Simon, but
since I have had the opportunity of examining the spider itself 1
have, rather reluctantly, come to the aamvakusiian that it must be
referred to a new species which appears, however, to be very
closely related to the above-mentioned.
My reasons for this are, briefly, as follows :-—
(a) In birmanicus the legs of the Ist, 2nd, and 3rd pairs are
subequal, the difference in length (cox excluded) between legs i
and ui being only | mm., with a difference of 0:25 mm. between
hand u3. whilst in the ‘species described below the difference
between 1 and i ii is 4°8 mm., whilst that between legs i and ii is
0-7 mm.
() In birmanicus the 4th tee is rather less than 3 times as-
long as the cephalothorax, in my specimen it is nearly 3°5 times
as long. Similarly, the Ist leg is nearly 2°25 times as long as the
cephalothorax, instead of slightly over twice, which is the case in
birmanicus.
(c) The palp (@) is nearly twice as : long as the cephalothorax
instead of being a little more than 1:5 times as long.
(d) The superior claws of the legs have 2-3 teeth instead of
3-4; also the inferior claws of the legs and the palpal claws
have 1-2 very small teeth instead of 2-3 small ones.
In cther words the legs and palpi are much longer in pro-
portion to the size of the cephalothorax than is the case in
LL. trmanicus, and furthermore there is a greater diversity in the
lengths of the legs, especially noticeable in those of Ist, gnd, and
3rd pairs.
* Unfortunately in a somewhat damaged condition. eve,
+ “A New Spider of the Genus Liphistius,” Journ. Malayan Binh Royal
Asiatic Soc. 1. p. 18 (1923).
770 MR. If, C. ABRAHAM ON A NEW
Before going on to the detailed description of the spider for
which, as it appears reasonable to suppose that it is the usual
jungle species of the genus in the southern part of the Malay
Peninsula, I propose the name Liphistius malayanus, I wish to
record my indebtedness and express my thanks to Messrs. H. R..
Hogg and F. de la Mare Norris; to the former for much help
and advice, and to the latter for having given me the opportunity
of examining and writing up this extremely interesting spider as
well as for kindly allowing me to make use of the excellent
photos which illustrate the description of the burrow and trap-
door. :
Family Lirwistrip#.
Genus Lipuistrius Schiddte.
Liphistius Schiddte, in Kroyer Naturh. Tijdsk. 2, ii. p. 621
(1849); Simon, Hist. Nat. des Ar. i. p. 64 (1892), ii. p. 875
(1903).
Lirnisrius MALAYANUS, Sp. Nn.
Lemale.—-Colour ; Cephalothorax and mandibles black ; fangs
dark reddish brown ; labium and coxe of palpi brownish ; cox
and sternum dark grey, nearly black ; legs and palpi black above,
dark grey (nearly black) below, with undersides of ‘ joints”
nearly white and sides of femora greenish; abdomen black,
opercula and fringes of spinnerets brown.
Cephalothorax.—About 1} times as long as broad, narrower in
front than behind, with the cephalic region rather more elevated
than is shown in Cambridge Natural History, Arachnida, fig. 201,
p. 386, or Fauna Brit. Ind., Arach., fig. 52, p. 155. There are
signs of a longitudinal series of bristles having existed on the
caput, behind the ocular tubercle. A number of bristles also
project forward from the anterior margin, below the ocular
tubercle, the median ones being by far the longest.
Iyes.—Closely grouped on a circular tubercle which is —
higher in front than behind and situated close to the front
margin of the cephalothorax on a distinct declivity (text-
fig. la & b).
The anterior laterals are considerably larger than the rest,
semicircular in shape, and contiguous, with their lower margins
occupying all the front half of the margin of the tubercle; the
posterior laterals, next in size, touch the hind margins of the
anterior laterals, are also nearly semicircular but rather pointed
behind, and occupy the remainder of each side of the tubercle ;
the posterior medians are oval, rather pointed behind, and about
7 as wide as long, they are situnted above and between the’
posterior laterals with their rear ends level with the middle of
the latter and the front edges a little in advance of the level of
the hind margins of the anterior laterals; the anterior medians
SPIDER OF TIE GENUS LIPHISTIUS. 771
are quite small, between the front margin of the posterior medians
and the upper edge of the anterior laterals and separated by
about their own diameter from each other and from the front of
the posterior medians: they are each placed on the top of a
small tubercle (text-fig. 1 @).
The eyes are brown except the anterior medians which are
black.
The ocular tubercle has a longitudinal median row of 4 or 5
curved bristles, and there are two more bristles near the inner
hind margin of each posterior lateral eye.
Mandibles.—Stout, about 2/5 as long as the cephalothorax.
The falx is strongly arched anteriorly, strongly convex on the
outside but hollowed at the base towards the lower edge, the
inner surface is flat. The anterior and upper surfaces are clothed
with slightly curved bristles which are longest along the inner
front margin.
Text-figure 1.
(ey) Profile. (6)- From above.
Kyes of Liphistius malayanus, sp. n.
The fang-groove has a fringe of reddish hair; the inner
margin is armed with 10 or 11 stout, bluntly-conical teeth, of
which the 2nd, 4th, and 8th are larger than the remainder, which
are irregular in size.
The fang is slightly curved, long, and stout, with slight longi-
tudinal striations along its upper surface.
Labium.—About twice as broad as long, the length being equal
to the breadth of the anterior margin of the sternum. It is
vounded anteriorly and provided with numerous slender curved
bristles. It is sunk below the level of the coxs, of which the
basal ends of the Ist pair close in behind it.
Sternum.—Long and narrow, and sunk much below the level
of the coxee of the legs, of which the basal ends approach one
another over it; about 3} times as long as its greatest breadth,
which occurs between cox ii and iii. In the only specimen
UU MR. H. C. ABRAHAM ON A NEW
available the posterior end of the sternum is somewhat damaged,
but the remaining part appears to have been clothed with
bristles.
Legs.—Compressed so as to make them squarish in section,
this is particularly noticeable in the femora, metatarsi, and tarsi ;
clothed with numerous black bristles arranged in longitudinal
rows. In addition to this there are a number of spines which are
distributed as follows :—-
The femora have two on the upper surface near the apex and
a row along each lower margin.
The patella have a row along the anterior upper margin, those
at the apex being the longest and curved.
The tibize have a series along each lower margin (5 in i and iii,
6 in ii, and 4, much finer, in iv; the basal one in each case much
smaller than the others), as well as an apical spine on each side
and a number of curved ones along the front margins.
The metatarsi of the Ist and 2nd _ pairs have 7 stout spines
along each lower margin, and one at the apex on the inner side ;
on the 3rd and 4th pairs there is, in addition, an apical spine on
the outer side and 3, rather finer, along each upper margin.
The tarsi have a row of stout spines along each lower margin
(7 or 8 along the front, 6 or 7 along the posterior); there are
also 3 ox 4 sane spines in a transverse row on the underside at
the base and about 4, also small, arranged in a V just below the
inferior claw and apparently intended to fulfil, to some degree,
the functions of ungual tufts or scopule.
The superior t tarsal claws are long, well curved, and stout ; ey
are armed on the basal half with 3 teeth, except in the athe one
of the Ist and 2nd legs, where there are only 2 teeth; in the
claws of the 4th pair all the teeth are long, sharp, and subequal,
but in the other cases where 3 teeth occur the basal one is very
small.
The inferior tarsal claws are armed with 1 or 2 very small
teeth.
Palpi.—Very similar to the legs both in general appearance
and armament.
The coxe have a thick fringe of long reddish curved hair along
the inner margins. i
The femora have a series of slender spines along each margin
of the underside, as well as a single spine near the Tease and of or
3 near the apex on the upper surface,
The patellses have a spine on the lower inner margin near the
distal end and also show signs of having had 2 or 3 on the upper
surface near the apex.
The tibiz have 5 5 stoub-spines along each margin of the under-
side, and also have a series of 2 along the inner guiding.
The tarsi have 7 long stout spines along each margin of their
lower surfaces.
The claw is stout, slightly curved, and armed with 2 very
small teeth near the base.
SPIDER OF THE GENUS LIPHISTIUS. . 773
Abdomen.—Apparently oval in shape and about 13 times as
long as wide (the abdomen of the only ayailable specimen is,
however, somewhat shrivelled, so that these proportions may not
be accurate). It is furnished with 9 distinct dorsal terga, and
there is a rudimentary 10th one midway between the 9th and
the anal tubercle. The 4th tergum is the largest and the 9th
the smallest. The sides and lower surface of the abdomen are
-wrinkled and thickly clad with bristly hairs; there are signs also
of the terga having been armed with bristles along their posterior
margins.
The spinnerets form a compact group about the middle of the
lower surface of the abdomen, the base of the anteriors being
nlmost exactly midway between the base of the abdomen and the
anal tubercle. ‘They are typical in form with the basal segment
of the posterior laterals about half the diameter of that of the
anterior laterals, and the anterior medians a little larger than the
posterior medians, The apical segments of tia laterals are
divided into about 12 false articulations, The anterior laterals
are separated at the base by about half their basal diameter; and
their apices reach a point halfway between the anterior margin
of their base and the anal tubercle, The anterior medians are
close together and just in front of the anterior laterals, whilst
the posterior medians, also contiguous, are situated, between the
two pairs of laterals, The bases of the posterior laterals are close
together and their distal extremities are curved inwards similarly
to those of the anterior laterals. The inner margin of the apical
seyments of all the laterals is furnished with a fringe of coarse
‘brownish hair.
The anal tubercle is conical, clothed with black bristles and
situated’ a little way in front of the posterior end of the
abdomen. é
Measurements (mm.).
Length. Breadth.
Cephalothorax’ 40. ocpcsecienn core e-sine 13°2 10°6
IO COMeMT Weta sces cies aacone tate neees 13:2 (8:6) *
DT bre EARN a ie LORI et Raa nis acer a 5:5
Lengths of Coxe. Tr.&fem. Pat. &tib. Met. &tar. Totals.
Megsi Tos. 6-0 10°3 10°6 8-2 35°]
TD 57 10°2 10°6 . 9-0 35°5
NEE eee 6:0 11-6 11-2 lley 39°9
AS eagecrtc 64 14°3 13°4 17°5 51:6
Ballpligeeceraace 4:5 9°3 9-4 6:0 29°2
* Approximate only, abdomen somewhat shrivelled.
1923, No. LL 51
Proc. Zoou. Soc.
7174 A NEW SPIDER OF THE GENUS LIPHISTIUS.
Habits*.—This spider inhabits web-lined burrows fitted with
a trap-door of the wafer type. The burrows, which are about
28 mm. in diameter and 400 to 450 mm. long, are found, from
the examination of about 20 examples, to be horizontal in general
direction, but usually are somewhat bent so that a straight stick
cannot be thrust in very far; they are unbranched and the inner
end is somewhat enlarged. ‘The trap-door is invariably hinged to
the mouth. of the tube at the upper edge, and another very inter-
esting fact is that the lines of web, which in Z. batuensis Abr.
ave employed to support the nest upon the vertical cave-walls f,
are by the present species used to keep the web-lining stretched
_ over the edge of the burrow so as to form a “lip” against which
the lid can close down tightly; this may be seen very clearly in
Plate I. fig. 2, in which the lid is shown in the opened position.
‘All the burrows found so far have had their openings on prac-
tically bare banks with perhaps a few small ferns, ete. growing
near, but, as all bare banks in this country are artificial cuttings,
and those in question have been made within the last 20 years or
so, it is almost certain that the spider must live in the jungle on
the neighbouring hills. The trap-doors of the observed burrows
all have small stones and particles of earth woven into them.
The spider appears to be entirely nocturnal, as pins placed in
front of the entrances of burrows were never disturbed during
the day, but have always been pushed out of position during the
night; the spider also seems quite “lost” in bright light. It
does not appear to be common as, in annual visits to the hill
upon which it was found from 1918 to 1922 inclusive, only com-
paratively few individuals—-not more than about 20 altogether—
have been observed. As is generally the case with burrowing
spiders, LZ. malayanus occupies the same nest for a long time ; in
fact, it probably does not change its quarters unless forced
to do so.
Locality —-Gunong Angsi, Negri Sembilan, F.M.S.; 2500 feet.
December 1922 (coll. Mr. F’. dela Mare Norris).
Specimens examined.—One adult female, which I have sent to
the British Museum, Natural History Department f.
* From notes of his field observations kindly supplied by Mr. F. de la Mare
Norris.
+ Abraham, l. e. supra, p. 19, and plate i. fig. 2.
f£ Together with its trap-door.
-
ASIATIC LIZARDS OF THE GENUS TROPIDOPHORUS. 775
38. A Review of the Lizards of the Genus Yropidophorus
on the Asiatic Mainland. By Maucorm A. Smrrx,
WERICrS Cal a Zias:
[Received May 28, 1923: Read October 28, 1923.]
This article concerns those species that are to be found distri-
buted throughout the Indo-Chinese peninsula—Burma, Siam,
and French Indo-China—and the region immediately north—-
Assam, Yunnan, and southern China. «Incidentally it completes
our knowledge of the whole group, the species of the Indo-
Australian region and of the Philippine Islands having been
recently undertaken by Nelly de Rooij* and Edward H. Taylor f
respectively.
The whole genus now includes some 18 species. It is essen-
tially a highland one, and is largely aquatic in its habits. Its
members are to be found in the vicinity of rocky streams, living
among the damp herbage on the banks, or hiding under stones
and boulders, sometimes almost completely immersed in the
water to which they invariably take to avoid capture. They are
nocturnal in thei habits and are not remarkably agile. One
species, Z', microlepis, 1 have found to be viviparous.
Different characters have at times been used to group the
species, but the one, so far neglected, which appears to me to
present the most natural basis for classification, is the manner in
which the scales are disposed along the sides of the body. It
divides the genus into two broad groups: (1) those with the
lateral scales directed straight backwards, with smooth or keeled
dorsal scales and smooth or feebly rugose upper head-shields, and
(2) those with the lateral scales directed obliquely upwards and
backwards, with strongly keeled, often mucronate dorsal scales
and with head-shields always rugose.
The preanal shields vary in number, but appear to be constant
as regards each species. On the other hand, the number of
scales round the body shows considerable individual variation,
and unless some other differential character can be shown to
exist, does not seem suflicient for specific distinction.
Jn coloration the species are all much alike.
Within the geographical limits discussed inthis paper I
recognize the following forms :—
I: Lateral scales directed straight backwards.
a. Head-shields smooth.
Fronto-nasal entire ............ccccceseeeeeseeeeeceseaveae 2". berdmovei.
Fronto-nasal divided .......0......ecceceeeeeeeeeeeeeeeee 1", lavtus.
b. Wead-shields feebly rugose ...........c..ceeeeseseeeeeeee ZL robinsont.
* © Reptiles of the Indo-Australian Archipelago,’ 1915.
+ ‘The Lizards of the Philippine Islands,’ 1022.
51*
776 MR. M. A. SMITH ON ASIATIC
II. Somo or all of the lateral scale-rows directed obliquely.
a. ‘Two prwanal shields.
Frontal and fronto-nasal entire, 4th supralabial
largest, dorsal scales strongly mucronate ......... T. assamensis.
Frontal and fronto-nasal entire, 4th supralabial
_ largest, dorsal scales not mucronate, median row
BiGATINNtO! se cise cache tenaMecccessenbsamuarecrseeen al a RAInanUs.
Frontal and fronto-nasal entire, 5th supralabial
IEVPRSIRS coocooseagadndoseoonsn, Lesboogccnon see oaN das cco ne T. cocincinensis.
Frontal entire, fronto-nasal divided .................. . sinicus.
Frontal and fronto-nasal divided ............-........ BP. thai.
b. Three prwanal shields .............:.:00seeeeeteeeeeeeee LD. microlepis.
The separation of many of the forms here mentioned is ex~-
tremely slight, and depends sometimes upon a single scale-
character. Most of them have a limited range of distribution,
and, so far as I am aware, no two forms have yet been found in
the same locality.
1. TROPIDOPHORUS BERDMOREI.
Aspris berdmorei Blyth, Journ. Asiat. Soc, Bengal, 1853, xxii.
. 61.
ray 'ropidophorus berdmoret Theob. Journ. Linn. Soe. 1868, p. 24;
Anderson, Zool. Res, Yunnan, 1878, p. 796, pl. Ixxvi. fig. 3;
Boulene. Cat. Liz. Brit. Mus, 1887, iii. p. 362; Annandale, Rec.
Ind. Mus. 1912, viii. p. 59; Mlem. Smith, Journ. N. 11.8. Siam,
1919, iii. p. 225.
Tropidophorus yunnanensis Bouleng. Cat. Liz. Brit. Mus. 1887,
iii. p. 362; Mlcm. Smith, Journ. N. H. 5. Siam, 1919, iii.
p. 224. —
Through the kindness of Dr. Annandale, Director of the
Indian Museum, I have been able to exainine the types—three in
number—of this species. ‘The specimens are somewhat faded
but are otherwise in excellent preservation. One has 32 scales
round the middle of the body, the other two have 34 each. In
the two smaller specimens the dorsal scales are distinctly, but not
strongly keeled, in the largest one I cannot find any keels at all.
Anderson, when comparing these types with his specimens from
Yunnan*, states that he found keels on all three, and, except
for this difference in the carination of the scales, they agreed
entirely, Hé therefore labelled the Yunnan specimens berdmoret.
his view should certainly be maintained, and 7. yunnanensis,
founded later evidently on a misunderstanding, became a
synonym of it.
My own collections of this lizard from Siam, supplemented by
an excellent series from Burma in the Indian Museum, have
enabled me to examine a large number of specimens. In the
number of scales round the body, and in the degree of carination
* The Yunnan specimens are now lost. Dr. Annandale tells me he has never been
able to discover what happened to the types of Anderson’s reptiles and batrachians.
Many of them never seem to have been incorporated in the Museum ‘collection.
LIZARDS OF THE GENUS TROPIDOPHORUS. 177
of the dorsal shields, they show considerable variation, but. J
cannot find any satisfactory combination upon which they can be
racially separated. Some have the scales smooth, some have
them faintly keeled, others more strongly. Hach locality has its
own slight variation. Where one adult has smooth scales, all
the other adults from that place have the same, and vice versa.
All the Burmese examples have two loreal shields, one behind
the other. Most of the Siamese individuals have three, the
anterior one being divided horizontally. The prefrontal shields
may be in contact, or separated, or have a small shield interposed
between them, irrespective of locality.
The following table shows the variation in the specimens I
have examined :—
BURMA. SIAM.
Locality. Scales. Locality. Scales.
Mercure ees Pi or.o4 ee Sa.on Witaradiubs 32--36
Tenasserim .............0. 36-40 Doi Nga Chang, 36-38
N. Siam. .
Takoo Mts., Central 36-38 Me Wang forest, 32-36
Tenasserim. N. Siam.
Pe mere beeiyaaees ace oa 40 5
Bia Po, Karin Hills ... 34
a SUNY eTVO) get ars hy mee ee 32
The allied Bornean TJ. becearit (Peters) appears to have a
variation similar to that of 7’. berdmorei, and J’. mocquardi
Boulenger should, in my opinion, be united with it. The speci-
mens in the British Museum show the range in scale-rows to be
from 28-36 round the body; apart from this I can find no
character upon which they can be separated.
2. 'TROPIDOPHORUS LAOTUS, Sp. n.
Types. Adults $ and 2, author’s Nos. 5410 and 5414, col-
lected at Muang Liep, N. of Pak Lai, Upper Mekong, French
Laos, in January 1920. Presented to the British Museum
(Natural History).
Description of the types.—Upper head-shields smooth; a pair
of fronto-nasals, each one Jonger than broad: prefrontals sepa-
rated by a small azygos shield; frontal as long as the parietal and
fronto-parietal together; parietals in suture behind .the inter-
parietals; 4 supraoculars, Ist largest, 1st and 2nd in contact
with the frontal, 4th entering the supraciliary border ; nostrilin a
single shield; two superposed anterior loreals, succeeded by a larger
posterior one; 6 supralabials, 4th largest and forming subocular ;
5 infralabials, the first longest and in contact with the postmental
and first chin-shield ; temporals small and like the body-scales,
except the superior, which is much larger; a single postmental
succeeded by two pairs of large shields. Tympanum as large as
the eye-opening.
778 : MN. M. A. SMITH ON ASIATIC
» Thirty-three scales round the middle of the body in the Bn
34 in the 9°, all quite smooth; ventrals largest, Jaterals easy
and directed straight backwards): ; a pair of salar ged preanals ;
series of enlarged scales beneath the tail. The higndelinns Pcie:
the wrist in the ¢, and not quite so far in the 2; subdigital
lamellee smooth, 18 to 20 beneath the 4th toes.
Dark brown above, with lighter black-edged V-shaped bars ;
sides of body with small white spots ; below whitish, throat and
tail thickly spotted with black.
Variation.—The azygos shield separating the pr sefrontals varies
in size, and may be absent, when either the preefrontals are in
contact or the frontal touches thé fronto ‘nasal, 82 scales round
the body occurs in nearly half the series. In one example
(No. 5416) there are only 30 scales. Im the young the dorsal
scales are faintly keeled, and the limbs are proportionately longer,
the leg reaching the wrist or the elbow of the ad pressed fore-limb:
7’. lotus is identical with the smooth-scaled form of 7’. berd-
morei except for the divided fronto-nasal. Two examples
(Nos. 5412 and 5425 from Muang Liep and Nong Kai respec-
tively) have the fronto-nasal single as in true berdmorei; but
with so large a series at hand I prefer to regard them as aberrant
examples of 7’. laotus.
Sixty-eight specimens examined, from Muang Liep, Nong Kai,
Pak Maat, ~Pak Men, and Hoi King, all localities on the Mekong
river, to the north onl south of, “ool within 80 kilometres, of,
Pak Tai.
Adult males can be recognized from Fame ke by the broader
hend at the angle of the jaw. Most of the old males also have
the head-shields considerably scratched and scarred, and | gather
from this that a considerable amount of fighting amongst them-
selves takes place.
Measurements of specimens of T. laotus in mm.
Author’s No. ... 6410 5414 6411 6415 65440 6417 6424
Head and body ......... 71 71 74, 75 76 51 40
Daily cwastemhoceonceseececs 95 105 100 96 105 82 62
Length of head .......... 18 17 17 17 19 13 115
Width of head ......... 12 10 11 11 12 8 65
Fore-limb ............... 20 19 20 20 21 14 12
Hind-limb ................ 30 27 | 28 30 31 21 18
SOx iiitaccnersedeecssoy a 2 2 2 rei juv. juv.
No. of scales ............. 33 34, 34 34 32 32 82
3. TROPIDOPHORUS ROBINSONI.
Tropidophorus robinsont Micm. Smith, Journ. N. H.S8. Siam,
1919, iil. p. 223,
Besides the large series from the type-locality there are six
specimens from ‘Cavoy, Tenasserim, in the Indian Museum
(Nos. 12722-12727). They agree with the types in every
respect.
LIZARDS OF TIE GENUS TROPIDOPHORUS. 779
4. TROPIDOPTIORUS ASSAMENSIS.
Tropidophorus assamensis Annandale, Rec. Ind. Mus. 1912,
Vill. p. 58.
Still known only from the type-specimen, which I have ex-
amined. From 7’. cocincinensis it differs in having the 4th
supralabial largest and below the middle of the eye, in “having no
small shields interposed between the loreals and supralabials,
and in the strongly mucronate dorsal scales. It has one large
anterior and one posterior loreal scale. ‘Che ventral keeling is
probably an immature Coaeaet er as the specimen is not yet fully
grown.
5, ‘TROPIDOPMORUS MAINANUS, sp. n.
Type. Author’s number 6997, collected at Ang Mao, alt. 600 m.,
near the Five Finger mountain, island of Hainan, in January 1923.
Presented to the British Museum (Natural Histor y).
Description of the type-—Upper head-shields strongly striated ;
fronto-nasal single, as long as broad, in contact with the frontal ;
frontal once and two-thirds longer than broad, as long as the
fronto-parietal and interparietal together; parietals in contact
behind the interparietal; 4 supraoculars, the anterior two
touching the frontal, the fourth just entering the supraciliary
border; 5 supraciliary shields anterior to the fourth supraocular ;
nostril in a single shield ; two anterior and two posterior loreal
shields ; temporals small, ‘with strong strie like the upper head-
shields ; 6 supralabials, the fourth largest and below the middle
of the eye; 5 infralabials, the first in contact with the azygos
postmental and anterior chin- shield. tympanum as lar ge as the
eye-opening.
Thirty-two scales round the middle of the.body; dorsals and
laterals strongly keeled, not mucronate, most of the scales in the
median dorsal row with two, keels, or where unicarinate smaller
than the others ; laterals smaller than the dorsals, the upper rows
with a feeble obliquity upwards and backwards ; ventrals larger
than the dorsals, quite smooth; gulars feebly keeled ; ; a pair of
enlarged praanals; three rows of smooth scales below the tail,
the median row transversely enlarged, The hind-limb reaches
the wrist.
Dark reddish brown above, with fodlisifinc! light dark-edged
cross bars, the anterior two V-shaped ; flanks with large whitish
dark- edged blotches ; belly white speckled black ; throat with
white longitudinal streaks.
Var iation. —Six specimens examined from the type-locality
(Nos. 6996 to 7001), and one (No. 7002) from the foot of the
Five Finger mountain about 20 miles distant.
The prefrontals are separated in every example; there are
seven supralabials in No. 7002, the 5th being below the middle
of the eye; one specimen hes only 30 sonllas round the body,
another has 34; in some examples only: a few scales of the
780 MR; M. A. SMITIE ON ASIATIO
median dorsal row are bicarinate; in two juveniles the ventral
seales are feebly keeled; the hind-limb may reach the elbow.
''wo examples are light brown in colour, with the usual
markings. _. .
Although Hainati is not the mainland of Asia, the fauna of
the island is so closely related to that of Indo-China, that this
species should be included here.
Measurements of specimens in mm.
, INO Sie aes sasaseeeesseenOOOd 7001
Length of head and body......... 47 49
AUMUN T cppecenascnoooas Sicletehe Soleo acre aoe 60 dD
IO RSENTTND. SbaGancooe BW asraraeve bats delle 13 12
HIG) Se onces0oaa00e0 o0c poboe oe eAD) 18
6. Tnorrpor HORUS COCINCINENSIS.
Tropidophorus cocincinensis Dum, & Bib. 1839, v. p. 556,
pl. lvii. fig. 1.
Tropidophorus éochinohinensis Bouleng. Cat. Liz. Brit. Mus.
1887, iii. p. 363 (in part).
_ ho types, three in number, two adults and one juvenile, are
in the Paris Museum. M. Angel has kindly examined them for
ime, and hns give mé the following particulars.
They have 30 and 32 scales round the body, the laterals a little
smaller than the others. Dorsal, lateral, ventral, and gular scales
keeled in the young example, the ventral and gular smooth in the
adults. ‘Iwo large preanal plates. ‘l'ail longer than the head
and body. Caudal scales strongly keeled except for the three
median .lower rows. The fifth supralabial is largest and below
the middle of the eye, and there are small scales interposed
between the supralabials and the loreals.
The exact type-locality of 7. cocincinensis is unknown; it is
said to have come from Cochin-China, but in view of the known
range of 7’. microlepis it seems probable that it came from
farther north.
7. TROPIDOPIIONUS SINICUS.
Tropidophorus sinicus Woettger, Zool. Anz. 1886, p. 519;
Bouleng. Cat. Liz. Brit. Mus. 1887, iii. p. 362; A. Mell, Archiv
fiir Naturgesch, 1922, 10 Heft, p. 114.
The type-locality is near Canton, and it has also been found in
the Man Son Mts., ‘Tonkin.
I recently obtained two half-grown specimens on the Peak in
Hongkong. ‘They have 30 scales each round the middle of the
body, the laterals and dorsals strongly keeled and feebly mucro-
nate, the ventrals feebly keeled; loreals 1+1; 5 infra-labials,
the first long and in contact with the divided post-mental and
first chin-shicld. Belly in life salmon-pink.
LIZARDS OF THE GENUS TROPIDOPHORUS. 781
8. TrorIpoPHORUS TIIAI.
Tropidophorus that Micm. Smith, Journ. N. H.S. Siam, 1919,
lil. p. 226,
Habitat. N. Siam. Still known only from the original spe-
cimens.
9, TROPIDOPHORUS MICROLEPIS.
Tropidophorus microlepis Giinther, P. Z. 8. 1861, p. 188; id.
Rept. Brit. Ind. 1864, p. 76, pl. x. fig. A.
Tropidophorus cochinchinensis (in part) Bouleng. Cat. Liz, Brit.
Mus. 1887, iii. p. 363; Mlem. Smith, Journ. N. H.S. Siam. 1919,
iii. p. 227.
As already stated, the preanal shields appear to be constant in
number as regards each species, and in 19 examples of 7’. nvicro-
lepis from the type-locality I find 3 preanal shields in every
instance. Another specimen from Dran, on the Langbian
plateau, S. Annam, has also 3 prianal shields, and agrees in all
other respects with the topotypes.
With 'so important a character to guide one it is justifiable to
maintain Giinther’s species, and to keep it distinct from cocinci-
nensis, the three type-specimens of which have only two preanal
shields each.
In 7’. microlepis the dorsal and lateral scales are all strongly
keeled and spinously produced, all of the lateral rows, except the
lowest, being obliquely directed. The first lower labial is very
small, allowing the second labial to make broad contact with the
post-mental.
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LARVAL ANATOMY OF THE GOUT-FLY OF BARLEY, 783
39, On the Larval Anatomy of the Gout-Fly of Barley
(Chlorops tantopus Mcig.) and two Related Acalyp-
trate Muscids, with Notes on their Winter Host-Plants.
sy J. G. IL. Frew, M.Sc. (Birmingham), F.Z.8.,
Ministry of Agriculture Research Scholar.
[from the Entomological Department, Rothamsted Experimental Station,
Harpenden].
[Received June 4, 1923: Read October 23, 1923.)
(‘Text-figures 1-23.)
TABLE OF CONTENTS. Page
Mi troduction cute seccsaneccesisne ces ssecereinenasctsceestoteranscsenctesseassersecay (Od
Chlorops Peniopne Wale,
A. en item e ORMIALV ao issiieeseenssessioessecetanieees seajsesoses LEO
Rady: segments Hed dsdacoTEH cod baGece ne: BeSbooBod Goo ca npCHadedRo cob aoe ace. Ae held/
B. Internal Anatomy of Tea hi ek 788
Cephalo-pharyngeal skeleton .. Berodopnrcbae sobcerecoocnocenm osc: Cite
Tracheal system ...... beaoondertecooad seoepecontod WAU!
Alimentary canal and its appendages .. 706
Central nervous Lelie imaginal discs and related
DIL IS Ta SGC ea PIR asH or Me A RI a EA ue a Marana Rar 7 te)
Other imaginal discs ieseasaniies Saal iacederecdsuneseeeaaes veces eIOOl
The dorsal vessel, nephirocytes, ete. . seebcsicesnasity slesseatenensnocc eee OUD
(nocytes .. scaeeaneesaneadvss cgades ees tedesceversestesasesente, 7 GUO
Fat body ..... BEER E BEE ee HEOCELI CHA ELLO CCU RDO CECE Sechelt,
Integument .. Be eke ecebedissilede dee Sania aaiden naasebvaeacelesea tease BOO
Gonadial rudiments. ......... Mocaauasutanmoucdwessmeenccacen (DOL
C. Other Immature peers of Chtoreps Laiwabad sinatdneeerieacroeen ie, TOOe
The puparium ......... AGiiaiercuaneaeeeacammenesnecheses OOM
The first larval instar . eek Cate da vtheanenc aalanaoe aL Ue hea ave aeereccr, OOS
Tho second larval instar ...cs.csssccscsssssesesseeserceseaecercsseseeee 809
The egg ......... Mawsrissuegedecbsateiaesascesesaareiesoomessteeete, 1 OU
Meromyza nigr jaenerts Mea.
The mature larva .. BE SS RRE SH ca AGA EE RR ARRON EA ees cae REM Ee aitidet oy) KO)
AMINO (VEG UR acnea cba cee coanbona ceca nod ouaackaocioononco cepeddacteabnccememir ton Ul
Oracle SsesmientS yin wceke uae seeeieanecs anon ceases ccnnen perbeciacsse see
Abdominal segments . SER SPOONS tint Pisa RS URGES cin GOS nCE TSO GUEAT ere
Cephalo- pharyngeal skeleton .. SHEARER oA GRAB an Gu poeisac abe aoumm eH
Tracheal eatery piers He Shy, Mecano ise ctaaeh oh eaio GE aaa i Sop totes aoa
Puparium .......... PESADOS RSE eae e oer anata ana remote metas ke}
Balioptera Lomb inate Ty ea test
The mature Jarva.. Be Arata nt) a eee salaccie Ma wircr ane ara Rerum nasal A
Dreshe nde eh eee eee en cae oatecsuan ites couse create oenalece ean OL
Mhoracicssepinentsice ss ren coe eee ee eee cee ecu easeetices OLD
Abdominal segments .....- PEP aera a cere een mena eee eOOL OD
Cephalo- pharyngeal skeleton .. RN Wer ein aati eneit cous iasee TOLO:
Muryaclicallisyis bem: scessece sss ccnhececnes seve tacseeser sesh eae te ites eeen Od:
Puparium ........ BERR OH SoA Groep TED EER Bria acORCo Seater oe Teco ia elue)
’ Chloropisca glabra Meinie RS Ma We GLa Tada oe tae Ost a OLD.
Winter Hosts of Species considered SA RS GERD BEC Oro nen Ciclaa One uoR Senn rele
Summary .. wh Meese tras cnn lsneaue cum neee eas mnainion igi OOO
Reference statin of Text AVR, Seana aat uM Sc tags el ODO)
WEES CLEFT EOVTELYS): Goa ena consegonn csonoobdoneeapousereocecadeon dos sanduseeacenccd (SPU)
784 - MR, J.G. 1. FREW ON TTE LARVAD,
INTRODUCTION.
The following paper deals with the preliminary work done in
the course of an investigation on the life-history, possible means
of control, etc., of Chlorops teniopus, the gout-fly of barley.
The investigation was commenced in October 1921 under the
direction of Dr. A. D. Imms. At the beginning of the work a
search was made through the literature of the subject to find
a recognisable description of the larva of Chlorops, but no such
description could be found. Gout-ry is usually easily recog-
nisable by the type of damage which it causes to the barley (or
other host-plant). ‘This, however, is not always the case and one
has, fairly frequently, to depend on the morphology of the larva
itself for certain diagnosis of the cause of damage to the host-
plant. A description of the morphology of the larva seems there-
fore to be desirable, as relatively little is known concerning the
morphology of Acalyptrate larve... The.internal anatomy of the
larva of Chlorops is also dealt with in this paper.
During the winter 1921-22 an extended series of examinations
of various grasses was made primarily in order to discover the
winter hosts of the gout-fly. During this grass examination
several other Acalyptrates besides Chiorops were bred out, and the
larval forms of two of them are described briefly at the end of
this paper. Although these larye normally pupate within their
host they will, if removed towards. the end of their larval period
when they have finished feeding, pupate and complete their
emergence if placed on iioist filter-paper in glass tubes. In this
way one can be quite certain.which larva is being bred out,
For the investigation of the-internal-anatomy of the larva of
Chlorops both dissections and serial sections were made. A certain
amount of the internal anatomy of the larva can be made
out in a living larva in a drop of salt solution, a method useful
in checking results obtained by other means.
I wish to express my indebtedness to Dr, A. D. Imms for
help and advice in all branches of this investigation, ‘To
Dr. W. E. Brenchley I am indebted for her kindness in
identifying all grasses examined during the difficult winter
period when only the vegetative eharacters were available.
For the identification of the imagines bred out I have to thank
Mr. J. E. Collin. :
Cuxonrors ‘raniopus Meig.
A, Eeternal Anatomy of Larva.
The following description applies to the third instar larva of
either the winter or summer generation ; the points in which the
first and second instars differ anatomically froi the third will be
indicated later.
The full-erown larva (text-fig. 1) is approximately cylindrical,
about 63mm. Jong and 1:2mm. in diameter in the posterior
ANATOMY OF THE GOUT-FLY OF BARLEY. 785
region of the third ‘thoracic segment, At the latter region the
diameter tapers off rather rapidly anteriorly tu the minute head
and very slightly and gradually posteriorly to the rounded anal
segment. The larva is apodous and of the usual Muscid type;
T consider that it consists of a head, three thoracic and nine
. Text-figure 1.
C. teniopus. Vateral view of mature larva.
abdominal segments. The larva is ainphipneustic, the posterior
spiracles being at the apices of two very small papillw carried
postero-dorsally by the last abdominal segment. The anterior
spiracles lie one on each side, projecting from the first body-segment
near its posterior border, slightly dorsal to the mid-lateral line.
The Head (text-fig. 2).
Tn 1904 Henneeuy proposed the term “ pseudo-cephalon” for
the anterior segment of Museid larvee, and Hewitt (1914) follows
him in this nomenclature. Becker (1910), however, has shown
that the head of a Muscid larva is homologous with the head of
such a eucephalous type as the larva of Stmuliwm, and there is
therefore no valid objection to the use of the term “head” in
describing Muscid larve.
The head is « small rounded body capable of almost complete
retraction within the first body-segment. Ventrally it bears
several paired sense organs and also the mouth-opening through
which project the apices of the mouth hooks; its surface is quite
smooth, with none of the chitinous ridges so commonly found on
the ventral surface of the head of eyclorrhaphous larve.:
The most prominent cephalic sense organs are the mawillary
palps (m.p.), each of which consists of a small rounded group of
sense papillz surrounded by a dark brown chitinous ring. Hach
sense papilla appears in surface view asa small shining ring with
a dark central region which is apparently a shallow depression at
the apex of the papilla. .The two maxillary palps lie close to the
anterior border of the mouth with their median borders almost
touching. From between: the two. maxillary palps a median
depression runs forwards along the ventral surface of the head
dividing it into symmetrical halves; unless the head is almost
completely exserted very. little of this groove can be seen.
Close to the antero-lateral borders of the mouth there is on
each side a pre-oral sense organ (p.o.). In the majority of
eyclorrhaphous larve this organ lies anterior to the imouth
between the latter and the maxillary palps; its lateral position
786 MR. J. G. I. FREW'ON THE LARVAL °:
in’ Chlorops is probably due to the unusually large size of the
maxillary palps and their close approximation to the anterior
edge of the motth-opening. Hach pre-oral sense organ is a
slightly raised papilla bearing several small rounded bodies similar
to those on the maxillary palps. «, According to Keilin (9) the pre-
oral sense organ is innervated by a branch of the nerve which
supplies the maxillary palps, of which, according to him, it is
probably a part.
The position of the antenne (a.) is peculiar in being not
anterior, but lateral, on a level with the anterior border of the
mouth and actually, therefore, posterior to the maxillary palps.
Kach is two-jointed, the basal joint being little more than a sinall
papilla with a flattened apex. ‘The second joint, which fits into
the apex of the first, is a cylindrical body of firm yellow chitin
Text-figure 2.
C. teniopus. Veutrel view of larval head.
A. Thoracic sense organ and neighbouring chitinous denticles.
B. Abdominal chitinous denticles.
with a rounded apex. Internally, at about one-third of its height,
the second joint is produced into a circular ledge whose inner
edge is bent at right angles towards the base of the antenna,
ISeilin (9) describes, in connection with the antenna of Pollenia
rudis Fabr., a ‘petit batonnet chitineux,” one end of which ig
attached to the outside of the base of the second joint of the
antenna while its other end is in connection with a “bulbe
nerveux.” In a later paper (10) he describes a similar organ in
the larval antenna of Graphomyia maculata Scop. with the
difference, however, that it arises from the inside of the second
antennal joint, and he seems to imply that such an appendage to
the antenna is universal in Cyclorrhaphous larve. J have failed
to find this appendage in the larva of Chlorops or any other
Acalyptrate larva, and although the number of species examined
ANATOMY OF THE GOUT-FLY OF BARLEY. 2 TEE
does not at present justify any generalization, it is .perhaps
possible that the absence of this appendage may be a reliable
distinguishing character of Acalyptrate larvee. h:
At the posterior border of the mouth: is a small lightly
chitinized plate (/.) bearing two small rounded sense organs (I.p.)
close to the median line and near the posterior end of the plate.
I think there can. be no doubt that these sensory organs are the
labial palps said by Keilin (9) to be present in all cyclorrhaphous
Jarvee which he has examined, though these are in many cases,
notably in certain carnivorous Anthomyid larvee (10), much more
prominent structures than they are in Chlorops. If this ter-
minology is correct the chitinized plate must be considered as a
Labium—the presence of which as a distinct sclerite in a cyclor-
rhaphous larva is of considerable interest.
There are several smal] sense organs on the surface of the lead,
and their position is shown in text-fig. 2 (¢.s.0.). Each is slightly
raised above the surface of the head and consists of a minute
rounded pit with an encircling chitinous rim.
Body-Segments.
There are twelve body-segments, three thoracic and nine
abdominal, the first thoracic and the last abdominal bearing
respectively the anterior and posterior spiracles. It is usual in
describing Cyclorrhaphous larve to distinguish a segment,
‘“‘Newport’s Segment,” between the head and the first body-
segment, which “is usually invaginated within the first thoracic
segment, so that it cannot be seen except when the larva is:
forcibly extended” (Lowne, p. 34). Lowne (p. 748) failed to find
this segment represented in the embryo, and I am unable to find
any morphological reason for considering it as other than the
invaginable anterior end of the first thoracic segment.
The lines of junction of the body-segments are marked by
bands of minute chitinous denticles which completely encircle the
body. The bands are not continuous but are formed of short
separate series of denticles. The thoracic teeth are long and
narrow, pointed posteriorly, and attached by their broad base
anteriorly. Very rarely their apices are bifurented (text-fig. 2 A).
The abdominal teeth (text-fig. 2 B) are considerably smaller and
approximately semicircular in shape with the flat side attached ~
anteriorly and the rounded apex projecting backwards.
The dorsal surface of the first thoracic segment is completely
covered by the rows of chitinous teeth. Ventrally it has a broad
anterior band .and a narrow posterior band leaving a narrow
transverse area between the two which is free from teeth. The
second thoracic seginent is very similar to the first in the arrange-
ment of the chitinous teeth. The third thoracic segment has a
broad band round its anterior border only.
Keilin (9) drew attention to the presence on the thoracic
segments of the great majority of Cyclorrhaphous larvie, of two
types of sense organs, both of which occur in the larva of
788 MN, J.G. Il. FREW ON THE LARVAL
Chlorops. Near the anterior ventral border of each thoracic
segment is 2 transverse series of about six small rounded sense
organs (ts.0.). ‘these are similar to, but rather larger than,
those occurring.on the ventral surface of the head, from which
they also appear to differ in being sunk in extremely shallow pits,
instead of being slightly raised above the surface. ‘These sense
organs have not been found on the dorsal surface of the body,
Each thoracic segment also bears two little sense organs at about
the middle of the ventral surface, one on each side of the median
line (text-fig. 2,v.J.). Hach consists of three tiny papille situated
very close together and each bearing a minute stiff bristle.
Keilin (9) has established the fact that the peduncles of the
imaginal leg rudiments are attached to the hypoderm immediately
below these organs which he regards as vestigial legs. I have
been able to verify this relationship in the eucephalous larva of
Forcipomyia piceus Winn, where there is no peduncle to the
imaginal dises, but I have failed to do so in Chlorops, where the
peduncles are extremely long and fine.
ach of the abdominal segments, except the ninth, has a
broad band of chitinous teeth round its anterior border. What
I regard as the ninth abdominal segment is considerably smaller
than any of the others. Ventrally it bears a longitudinal slit,
the anus, bordered on each side by a slightly raised rounded lip ;
postero-dorsally it bears the two posterior stigmatic papille.
When the larva is fully extended the division between the eighth
and ninth abdominal segments is indistinguishable; the ninth
‘segment is, however, capable of being partially withdrawn within
the eighth and should, I think, be regarded as a separate segment.
This conclusion is strengthened by a consideration of the distri-
bution of the abdominal sense organs in the larva of Merumyza
nigriventris Meq., which will be dealt with later. No abdominal
sonse organs were found in the larva of Chlorops.
The form of the spiracles is described later in connection with
the tracheal system.
ea B. Internal Anatomy of Larva.
The Cephalo-Pharyngeal Skeleton (text-figs. 3, 4 & 5, A-H)
consists of two mouth hooks, an intermediate sclerite (i.s.) which
bears anteriorly an articular surface for each mouth hock, and a
large pharyngeal sclerite (p.s.). The two latter sclerites are
fused, but the line of junction is clearly visiole owing to the
heavier chitinization of the intermediate sclerite. For the sake
of convenience the mouth hooks are, throughout this paper,
referred to as “mandibles,” though it must be understood that
no attempt is thereby made to bomologize them with the man-
dibles of Orthorrhaphous larve. According to Weismann’s
embryological work (16) they are entirely new structures, Holm-
gren (5) considers them to be homologous with the mandibles
of Orthorrhaphous larve but brings forward no embryological
ANATOMY OF THE GOUT-FLY OF BARLEY. 789
support for his theory, and his comparisons of certain larval
types by which he attempts to establish his view are adversely
criticised by Becker (1).
Hach mandible (m.) is a stout, heavily chitinized hook with a
pointed claw-shaped anterior end and two posterior processes,
a smaller heavily chitinized dorsal one with a rounded posterior
surface which articulates with the corresponding articular surface
of the intermediate sclerite, and a Jarger ventral one flattened
from side to side, into which are inserted the depressor muscles
Text-figure 3.
@ ~Pp.s. ;
o :
‘2 mm.
C. teniopus. Dorso-lateral view of cephalo-pharyngeal sclerites,
of the mandible. At the point of junction of the claw and the
anterior end of the ventral process there is a small, less heavily
chitinized, but sharply pointed, accessory tooth. The dorsal
process is broad from side to side and somewhat flattened dorso-
ventrally. It is held in close apposition to the articular surface
of the intermediate sclerite by the tension of the depressor and
levator mandibular muscles, and when these are cut, or even when
the larva is killed, the mandibles very easily come apart from the
Y ‘Text-figure 4,
tt
On may
Lateral view of cephalo-pharyngeal sclerites.
intermediate sclerite. The two mandibles are fused together
medianly over a small area but their articular surfaces are
separate.
The relations of the intermediate and pharyngeal sclerites can
best be understood by a reference to text-figs. 3 & 4 and text-
fis. 5, A-K, which represents a series of transverse sections.
through the sclerites at the points marked by arrows on text-.
fig. 3. As mentioned above, the intermediate and pharyngeal
sclerites are fused. The form of the whole compound sclerite is.
Proc. Zoou. Soc.—1923, No. LI. 52
790 MR. J. G. HH. FREW ON THE LARVAL
that of a trough with a double ventral wall, the space between
the two ventral walls being the cavity of the pharynx. The
trough is surrounded by hypodermis and the cavity of the trough,
above the pharynx, is also lined by hypodermis, the two layers
being continuous round the posterior end of the trough. (This
hypodermis is only shown in text-fig. 5, D & H.) The dorsal
edges of the trough are joined together by a sheet of hypodermis
(which is really double though its double nature is never very
distinctly visible in sections) which appears to secrete a thin
layer of cuitin in places. I consider that this double layer of
Text-figure 5.
Ol mm.
‘Transverse sections tirough the points marked A-Fi in text-fig. 3.
Chitin shown black or dotted according to the density of the chitin. Hypoderm
and dorsal membrane cross-hatched. The complete hypoderm is shown only
in text-fig. 5B. It is partially shown in 5D; not at all in the other
figures.
hypodermis is homologous with the hypodermal fold in the larva
of Chironomus, which is derived from the dorsal region of junction
of head and neck and extends backwards into the thorax and
from which arise the cephalic imaginal dises (14). In Chlorops
the cephalic imaginal discs arise from the posterior end of this
double hypodermal sheet. In the region represented by text-
fig. 5, A & B, the dorsal membrane is present, but I am very
doubtful as to its nature,as I failed to find any hypoderm in con-
nection with it.. It may possibly be chitinous, which is certainly
ANATOMY OF TILE GOUT-FLY OF BARLEY. 791
its appearance, and may be the result of the complete transfor-
mation of the original bypoderm into chitin. Further back
(text-fig. 5C) the dorsal membrane consists only of hypoderm
In text-fig. 5D hypoderm is present (not shown in the figure)
cand secretes a thin layer of chitin. In text-fig. 5 HE, only hypo-
derm is present.
The dilator muscles of the pharnyx (d.m.) arise from the upper
parts of the walls of the trough and are inserted into the dorsal
wall of the pharynx, which they raise by their contraction, thus
enlarging the capacity of the pharynx and sucking up food
through the mouth. In the extreme anterior region (text-
fig. 5 A) there are no dilator muscles, their space being occupied
by a blood-sinus. ‘The posterior end of the pharyngeal sclerite
shows two lateral incisions extending far forwards and dividing
it into three arms, a ventral one containing the pharyngeal cavity
‘and two solid dorsal ones.. Owing to their depth the dorsal arms
appear, when viewed dorsally, to be more heavily chitinized than
the ventral one; that this is not so is shown in text-fig. 5 KH.
The hypoderm surrounding the ventral arm is directly continuous
posteriorly with the epithelium of the oesophagus.
The appearance shown in text-fig. 3 of a hole through the
posterior region of the intermediate sclerite is due to the thin
ventral walls of the trough at this point (text-fig. 5C), and the
light chitinization of the lower of the two. It is in this region
that the salivary duct opens into the pharynx.
The Tracheal System (text-figs. 6 to 10) is of the usual Muscid
type, the larva being amphipneustic.
The Anterior Spiracles (text-fig. 6) are of the usual radiating
type. The anterior end of the main tracheal trunk expands
slightly, forming a small bulbous region (0.), the wall of which is
strengthened by a ridge (7.) formed apparently as a hollow out-
pushing of the wall, which goes round the bulb, forming
alternately anterior and posterior loops. Anterior to the bulb is
the short Stigmatie Trunk or Felted Chamber (fe.), the Filz-
kammer of De Meijere (6), ‘Chis is of somewhat greater diameter
than the bulb, and sections show that its lumen is broken up by
numerous fine chitinous hairs and ridges arising from its walls.
The distal end of the felted chamber expands slightly and,
bending upwards at right angles to its proximal portion, projects
from the surface of the body in the dorso-lateral region of the
first body-segment near its posterior border. The expanded part
‘of the felted chamber which is antero-posteriorly flattened bears
a number of radiating arms. Stigmata bearing 6, 8, 9, and 11
arms have been found, 8 being the usual number, In all cases
examined, the two anterior stigmata of a single larva had the
same number of arms. Each arm bears on its posterior face near
its apex a minute longitudinal slit (s.). The lateral edges of the
slit are curved so that the slit is narrowest in the middle, widens
out slightly at its proximal end and considerably at its distal end,
‘The proximal end of the slit is produced upwards into a minute
52*
792 MR. J. G. 1. FREW ON THE LARVAL
triangular flap, and its distal end is produced downwards into
a similar but more prominent flap, the two flaps largely filling
up the widened ends of the slit. The whole felted chamber
with its branches is enclosed in a delicate, chitinous sheathing-
membrane (s.m.).. ‘Chis membrane appears to he absent from the
extreme apices of the branches and does not, presumably, cover
over the openings. These two points could not, however, be
settled definitely.
The Posterior Spiracles (text-figs. 7 & 8) are situated at the
apices of two very small stigmatic papillae carried postero-dorsally
by the last body-segment. Each papilla bears at its apex a
rounded stigmatic plate (s.p.) in which are the three stigmatic
openings. The posterior stigmata are of the same general type
as the anterior ones. A bulb, which is, however, of no greater
Text-figure 6,
Posterior view of anterior spiracle.
diameter than the tracheal trunk, js interposed between the felted
chamber and the tracheal trunk. Its wall is strengthened by two
sinuous ridges which may be separate from each other, or the
proximal one may be incomplete with its ends fused with the
distal ridge as shown in text-fig. 7. I have not as yet seen a
case where the distal ridge was incomplete. The felted chamber
is relatively narrow and long, with its walls produced internally
into delicate ridges and hairs; distally it divides into three very
short branches (not, however, all in one plane as are those of the
anterior stigmata), each of which opens on the stigmatic plate
by one of the openings mentioned above. In an apical view of
the stiginatic plate (text-fig. 8) a central dark area is seen caused
by looking through the plate into the conical space between the
ANATOMY OF THE GOUT-FLY OF BARLEY. 793
three short arms of the felted chamber. In neither anterior nor
posterior stigmata have I been able to distinguish any Stigmatic
Scar. According to De Meijere, the visibility of this structure
varies greatly in difterent species. It is, however, probably
present in all third instar Cyclorrhaphous larvee.
On each side of tire body there is a dorso-lateral main tracheal
trunk uniting the anterior and posterior stigmata of its side
(text-fig. 9, l.¢.). Uniting the two dorsal trunks are a series of
ten dorsal commissures, the second of which is rather strongly
looped forwards, the remainder being practically transverse. The
first and tenth commissures, particularly the latter, ave thicker
than the intervening ones; the first gives off from each end an
anteriorly directed branch which goes to the pharyngeal mass ;
Text-figure 7.
?.
Dorsal view of posterior spiracle. (The stigmatic plate is seen by
transparence.)
the remainder are quite devoid of branches. ‘The middle point.
of each commissure shows a small rounded thickening which
sections show to be attached to the dorsal body-wall by a slender
suspensory ligament. The distribution of the dorsal commissures
is as follows :—No. 1, anterior region of body-segment 3. No. 2
middle of segment 3, but looping strongly forwards. No. 3,
anterior region of segment 4 (7. ¢. tirst abdominal). No. 4 arises
at about the junction of segments 4 and 5, and nos. 5-10 arise
similarly at the points of junction of the other body-segments.
There is no transverse commissure at the point of junction of
segments 1] and 12.
Each dorso-lateral trunk gives off eleven lateral trunks, the
anterior three of which are small and are mainly distributed to
794 ’ MR. J.G. WH. FREW ON THE LARVAL
the pharyngeal mass and its associated muscles, forming in this
region a complicated plexus, The remaining eight lateral trunks
pass down the lateral body-wall and bifurcate into anterior and
posterior branches, by the joining up of which branches (text-
fig. 9) acontinuous but undulating ventro-lateral trunk is formed,
from which branches ave given off to the ventral body-wall
and a few small ones to the lateral body-wall. ‘The ventral body-
wall branches of one side do not appear to anastomose with those
of the other side.
Text-figure 8.
0-03 mm.
Apical view of posterior spiracle.
Each of the eight posterior lateral trunks except the second
and the last gives off two large branches, an anteriorly distributed
and a posteriorly distributed; the latter may arise from the
anterior: side of the trunk; the second and last lateral trunks
have no anterior branch, and the last has three posterior branches.
The anterior branches of the lateral trunks are deep lying; they
Text-figure 9.
——Wadun
Dorso-lateral view of larva to show tracheal system.
supply the viscera and then run forwards, approaching the median
ventral line, and supply a large ventral tracheal plexus in the
region of the ganglionic complex. ‘The posterior branches divide
into two parts, one supplying the lateral body-wall, the other
passing dorsally superficial to the dovso-lateral trunk to supply
the dorsal body-wall. Zhe tracheation of the dorsal body-wall is
supplied exclusively by branches from the lateral trunks, the dorsal
commissures supplying no branches (text-fig. 10 A). oe
The first three (small) anterior tracheal trunks ave joined to
the corresponding trunks of the opposite. side by three very
ANATOMY OF THE GOUT-FLY OF BARLEY. 795
delicate ventral commissures which do not unite the trunks them-
selves but corresponding branches. No other ventral commissures
could be discovered. ‘Che second tracheal trunk on each side
gives off a backwardly-directed branch; these two branches con-
verge to the dorsal surface of the brain, where they are joined by
a short transvere commissure. From here they continue back-
wards, one on each side of the dorsal lobes, and pass downwards
one on each side of the cesophagus, probably ending in the ventral
tracheal plexus.
Text-figure 10.
OS mm.
Dorsal commissures and their tracheal branches in A. Chlorops teniopus,
B. Balioptera combinata, and C. Meromyza nigriventris.
All the lateral trunks except the first two bear a small
appendage (v.f.) appearing as a very short tracheal branch. An
attempt to investigate the nature of these by means of sections
was not completely successful, but indicated that they were
solid strands directed towards the lateral body-wall and in all
probability attached to the latter. They are probably vestigial
stigmatic trunks such as have been recorded by Imms (8) in the
larva of Anopheles, and by Carpenter and Pollard (2) in the larva
of Hypoderma.
Peristigmatic Glands.
Lying at the base of each posterior stigmatic papilla just ventral
to the tracheal trunk and close to the hypoderm, is a group of two
or three somewhat flask-shaped cells. A duct Jeaves each cell, and
the three ducts join forming a common duct which passes back-
wards towards the stigmatic plate, upon which it probably opens,
796 MR. J.G. H. FREW ON THE LARVAL
though it could not be traced to any opening. The gland cells
are probably modified hypoderm colls. Keilin has described the
peristigmatic glands in several Cyclorrhaphous Jarvee.
The anterior stigmatic trunks are surrounded as they traverse
the hypoderm by modified hypoderm cells which are probably of
the nature of peristigmatic glands, although they are not nearly
such prominent structures as the posterior peristigmatic glands
and are apparently unprovided with ducts.
The Alimentary Canal and its appendages.
The structure of the pharynx has been described in connection
with the pharyngeal sclerites. he hypoderm surrounding the
ventral posterior arm of the pharyngeal sclevite is directly con-
tinuous posteriorly with the epithelium of the esophagus, which
is a straight, narrow tube with a very small lumen. In the first
abdominal segment it traverses the ganglionic mass, slightly pos-
terior to which it enters the cardiac valve (proventriculus). ‘This
is of the type commonly found in Dipterous larvee. It is a pear-
shaped body, its broad anterior end being continuous with the
cesophagus, its narrow posterior end with the cardiac division of
the midgut. ‘ne structure of the proventriculus is essentially
simllar to the house-fly larva (see Hewitt, p. 187) and will not
therefore be described, The cardiac ring of imaginal cells occupies
the same position as it does in the larva of the house-fly and other
Dipterous larve.
The proventriculus lies in the anterior region of segment 5
(2nd abdominal), From here the midgut runs backwards to the
anterior region of segment 10; turns dorsally and runs forwards
to the posterior region of segment 4; backwards to seginent 6,
where its diameter decreases considerably ; forwards to the pos-
terior region of segment 5; backwards, its diameter increasing
somewhat, to the middle of segment 10; forwards asa wide, thin-
walled tube with a very poorly developed epithelium to the
posterior region of segment 7, where it passes into the hindgut.
"This last section of the gut is liable to become enormously
distended, when its epithelium becomes extraordinarily thin and
ill defined ; its contents are usually of a dark brown colour.
No attempt has yet been made to work out the detailed
‘histology of the different regions of the midgut. As mentioned
above, its last section was, in all examples studied, considerably
distended; other regions of the midgut are also liable to dis-
tension, particularly the regions of the bends, and the appearance
of sections of the gut-wall varies very greatly according to the
degree of distension, great distension being frequently accompanied
by partial degeneration of the epithelium of the affected region.
‘Lhe possession of a striated border is a constant characteristic of
the cells of all regions of the midgut epithelium, and this border
can usually be made out by careful examination even where the
gut is very greatly distended. ‘I'here is no peritrophic membrane
ANATOMY OF THE GOUT-FLY OF BARLEY. 197
in the midgut. From its point of junction with the midgut, the
hindgut passes forwards for a short distance near to the dorsal
body-wall and then turns backwards and runs as a practically
straight tube to the anus. At its commencement the hindgut
is slightly swollen and bulbous; on this follows the narrow
anteriorly-directed region. ‘Lhe posteriorly-directed region is
dilated until a short distance in front of the anus, where it again
becomes a narrow tube. The whole of the hindgut is lined with
a well-marked chitinous intima, which is thrown into numerous
longitudinal folds, some of which are very prominent. There
is a well-defined ring of imaginal cells at the junction of the mid-
gut and hindgut.
Salivary Glands.
The salivary glands are two simple tubular bodies with a con-
tinuous central intercellular Jumen. ‘he glandular portion (as
distinct from the duct) begins at the level of the proventriculus
and passes backwards to about the posterior limitof the ninth body-
segment, the glands lying one on either side of the gut. In about
the sixth or seventh body-segment each gland turns dorsally,
then forwards for a short distance, then again dorsally, and then
backwards and ventrally to its termination near the ventral body-
wall. These bends are always present, but their position is not
absolutely constant. In transverse section the gland is circular,
with a central star-shaped lumen surrounded by four to six gland
cells. At the anterior end of each gland there is a narrow ring
of small imaginal cells surrounding a small central lumen which
is continuous posteriorly with the lumen of the gland and an-
teviorly with that of the salivary duct.
The duct is a narrow tube lined by a chitinous intima, its
walls being formed of flattened cells, of which three are usually
seen in any transverse section except anteriorly, where there are
only two. Hach duct runs forwards, becoming very closely applied
to the ganglionic mass, and lying in the groove dividing the sub-
esophageal from the supracesophageal ganglion. Anterior to the
ganglionic mass the two ducts lie ventro-lateral to the esophagus
and directly ventral to the lateral edges of the antennal imaginal
discs. As they pass forwards they approach the median line
finally joining, slightly posterior to the pharyngeal mass, to form
a single median duct which opens through the ventral wall of the
pharynx at the point marked X in text-fig. 3. The duct narrows
considerably just before entering the pharynx.
Malpighian Tubes.
There are two pairs of malpighian tubes, an anterior and a
posterior. ‘The anterior pair unite to form a short common tube
which narrows considerably as it passes mesially to open into the
left side of the gut at the point of junction of midgut and hind-
gut. ‘Che posterior pair of tubes forms a similar short common
tube opening in a corresponding position on the right side of the
798 _ MR. J. G. H. FREW ON THE LARVAL
gut. From their point of junction the posterior tubes pass back-
wards in the dorsal region of the body, having a somewhat wavy
course but no convolutions or sharp bends. The anterior tubes
are almost twice the length of the posterior ones; from their
point of junction they pass forwards in the dorsal region of the
body to about the level of the proventriculus. They then turn,
abruptly posteriorly. The distal limb of each tube lies lateral to.
the proximal one, and has a somewhat greatef diaineter, ‘The
distal limbs extend backwards to a point slightly posterior to the
point of origin of the malpighian tubes from the gut.
The proximal limbs of the anterior tubes are transparent and
appear dark by reflected light; their lumen is very narrow.
The distal limbs are opaque, glistening white by reflected light,
and have a relatively wide lumen which contains a frothy mass.
At present no more precise details can be given concerning this.
difference in structure, and presumably function, of the two-
regions of the anterior malpighian tubes, but it is hoped shortly
to investigate the matter more fully. All that can be said at
present is that the frothy mass in the distal limbs of the anterior
tubes does not appear to consist of CaCO, which does cccur in the
malpighian tubes of certain Cyclorrhbaphous lJarve (11). The
posterior tubes resemble in structure the proximal limbs of the:
anterior tubes, The Jumen of the malpighian tubes is in all places.
intracellular.
Central Nervous System—Cephalic Imaginal Discs and
related parts (text-fig. 11).
The cerebral ganglia are rounded lobes lying in the first ab-.
dominal segment. ‘They are separated from each other by a
longitudinal dorsal fissure, except in their antero-ventral regions,
where they are united by a short, and moderately thick, trans-
verse commissure, Hach is united by a broad csophageal
commissure to the ganglionic mass lying below the csophagus.
This ganglionic mass (called hereafter the subcesophageal ganglion)
is a more or less cone-shaped body; it hes mainly in the first
abdominal segment, but its posterior end extends a short distance
into the second abdominal segment. It represents the fusion
of the subewsophageal, thoracic, and abdominal ganglia of less
specialised insects.
The wsophagus passes backwards from the pharynx along the
dorsal surface of the subeesophageal ganglion. For part of its
course it lies at the base of the fissure separating the dorsal lobes
of the brain, and the transverse commissure (¢.c.) uniting the
two lobes lies immediately dorsal to it. The dorsal vessel (d.v.)
dips down from the dorsal body-wall and passes through the
dorsal region of the fissure separating the dorsal lobes of the
brain. Between the posterior ends of the dorsal lobes it is
surrounded by Weismann’s Ring (w.7.), an oblique cellular collar,
of which the dorsal part is anterior, the ventral part, posterior,
The ring is relatively thick at the sides but less so dorsally, where
ANATOMY OF. 'THE GOUT-FLY Of BARLEY. 799:
is pvoiluced laterally on each side into a fairly stout suspensory
process (s.p7.) whose other end is attached to the surface of the
dorsal lobe of its side. hese suspensory processes do not appear
to contain any nervous elements. ‘The dorsal and lateral regions
of the ring are formed of fairly large cells, each with a single
large nucleus containing a big, densely-staining nucleolus and a
few scattered chromatin granules. The ventral and posterior part
of the ring is composed of very small cells which have exactly the
appearance of the cortical ganglionic cells. This region is un-
doubtedly a ganglion; it becomes more prominent in mature
larvee, in which it is marked off from the lateral regions of the
ving by a deep fissure on each side. <A strand of tissue
(presuinably nervous) (p.7.) leaves this ganglion and passes back-
wards and downwards to the dorsal surface of the proventriculus
where it forms a small ganglion (p.g.). Another strand (m.n.)
goes forward and downward from the ganglion and passes under
the transverse commissure uniting the dorsal lobes of the brain,
in front of which it breaks up into four nerves which pass
forwards to supply the esophagus and pharyngeal mass. In the
blow-fly larva, according to Lowne (18), the corresponding nerve
swells out to form a small median ganglion just posterior, to
o
the transverse commissure, and from this ganglicn nerves pass.
forward along the csophagus, round which they form a plexus.
The median ganglion could not be distinguished in Chlorops.
From the posterior region of the transverse commissure a stout
vertical strand of a fibrous nature (v.s.) passes upwards to be
inserted into the ventral wall of the dorsal vesssel. ‘There is no:
ganglion at its point of junction with the dorsal vessel and it
gives off’ no nerves; it is probably, therefore, purely suspensory in
function. The dorsal vessel passes forwards and, slightly anterior
to the transverse commissure, it fuses with the posterior region
of the cephalo-pharyngeal band in the manner described below.
The dorsal cornua of the pharyngeal sclerite ave each surrounded
by hypoderm, the hypodermal sheaths being united by a narrow
arched band of hypoderm, which is continuous anteriorly with
the hypodermal sheet which. unites the dorsal margins of the
pharyngeal sclevite and which, as mentioned in the description of
the pharyngeal skeleton, is really a double sheet. ‘The hypo-
dermal sheaths of the dorsal cornua are continued posteriorly
beyond the termination of the pharyngeal sclerite as hollow tubes
of hypoderm (i.s.p.) united by the arched dorsal hypodermal
plate (c.p.0.). The hollow tubes are the peduncles of the cephalic
imaginal sacs and the arehed plate is the cephalo-pharyngeal
band. The relationships of the parts are most easily made out in
fairly young third instar Jarvee: in mature larve the enormous
growth of the cephalic imaginal discs makes the interpretation of
sections somewhat more difficult. In such:a young larva the
peduncles of the sacs decrease rapidly in diameter as they pass.
backwards and their cavity becomes practically obliterated. In
the third thoracic segment they increase in size to form the
800 MR. J. G. H. FREW ON THE LARVAL
cephalic imaginal sacs which contain the imaginal dises and a
well-marked though narrow. cavity. The cephalo- -pharyngeal
band extends outwards under the median region of their ventral
surface, and its lateral margins are then produced downwards to
form a lateral flap (1) on each side. Slightly more posteriorly
the imaginal sacs separate completely from the cephalo-pharyngeal
band, their posterior ends being attached to the dorsal lobes of
the brain by nervous tracts.
The lateral flaps are not deep anteriorly, but as they pass pos-
teriorly their depth increases so that they hang down close against
the sides of the cesophagus, their ventral edges lying slightly
below the level of the ventral surface of the cesophagus, and
being attached to two small longitudinal traches which will be
mentioned later. There is thus a well-defined sinus whose floor
is formed by the esophagus, and whose roof and sides ave formed
by the cephalo-pharyngeal band and its lateral flaps respectively.
At about the anterior limit of the brain the dorsal vessel passes
into this supracesophageal sinus, its dorsal and lateral walls fusing
with the cephalo-pharyngeal band: The vessel ends by opening
into the sinus at the point marked Y in text-fig. 11. Slightly
posterior to the opening of the vessel into the sinus the dorsal
regions of the lateral flaps approach one another below the vessel
and fuse, forming a vertical median plate (m.pl.). ‘The vertical
plate is short in the antero-posterior direction, and posterior to
its termination the ventral parts of the flaps form an arched mem-
brane over the cesophagus. As it passes backwards this membrane
becomes less and less extensive and more and more closely
applied to the oesophagus, and where the latter passes beneath
the transverse commissure its dorsal and lateral regions are closely
surrounded by the membrane, the ventral edees of which are
united to the membrane covering the dorsal surface of the sub-
cesophageal ganglion and to the two small longitudinal trachez
already mentioned. The ventral parts of the lateral flaps do not
extend posteriorly beyond the transverse commissure.
The two small longitudinal trachez: mentioned above arise in
the head region and pass backwards, one on each side of the
«esophagus, until they come into relation with the ventral edges
of the lateral flaps along which they pass to the dorsal sur face of
the subeesophageal ganglion.
As the larva approaches maturity the cephalic imaginal sacs
and their contained discs increase enormously in size. ‘The two
sacs extend upwards above the cephalo-pharyngeal band until
they are in contact along the median line; probably in a very
late stage the two sacs communicate with each other as is the
case in AMelophagus ovinus (15), but this stage has not been
observed, Hach sac when almost fully developed consists of an
antero-vential region containing the antennal disc and a postero-
dorsal region which contains the eye rudiment and also the rudi-
ments of certain regions of the head capsule. ‘he eye rudiment
is innervated from the posterior region of the corresponding
ANATOMY OF THE GOUT-FLY OF BARLEY. 801
dorsal lobe; the antennal rudiment is, I believe, innervated
from the anterior region of the dorsal lobe, but I have not
been able to establish this quite definitely. The ocular half of
the imaginal sac overlies the anterior half of the corresponding
dorsal lobe.
The process of evagination of the cephalic discs is believed to
be as follows:—The right and left sacs fuse in the median
line above the cephalo-pharyngeal band so that their cavities
communicate, This fusion gradually extends forwards to include
the peduncles whose central cavities come to form merely the
lateral regions of the peripodal cavity of the conjoined sacs.
When this fusion extends as far forwards as the point of origin
of the peduncles from the pharyngeal part of the cephalo-
pharyngeal band, which as mentioned above appears to be a
Text-figure | Ie
10mm.
Diagram to show relations of cephalo-pharyngeal band, dorsal vessel, and related
structures. The right dorsal lobe of the brain, and the tight cephalic
imaginal sac are represented as having been removed ; left cephalic imaginal
sac and its peduncle not shown. Part of the cephalo-pharyngeal band
between the points E and E! is represented as having been removed. (Recon-.
structed from serial sections and dissections.)
double layered structure in this region, the two layers of the
band become more definite and split apart, the space between
the two layers being continuous posteriorly with the cavity of the
united imaginal sacs, and it is through this space that the
imaginal discs evaginate. I have not seen this process but, as is
mentioned below, the commencement of such a fusion of the sacs
and their peduncles has been seen in connection with the pro-
thoracic leg dises.
Thoracic Imaginal Discs.
On each side of the body there are three leg dises—prothor-
acic, mesothoracic, and metathoracic. From the antero-ventral
region of the sub-cesophageal ganglion there arise on each
802 MR, J. @. WW. FREW ON THE LARVAL
side three stout nervous tracts one supplying each leg disc.
Each imaginal leg sac is pear-shaped with its broad posterior
end receiving the nerve from the ganglion, its narrow anterior
end being produced forwards into the peduncle of the sac.
The prothoracic leg sacs and their contained imaginal discs lie
in the median region of the third thoracie segment. In all
specimens examined the peripodal cavities of the sacs communi-
cated with each other, but this union would probably not be
found if sufficiently young specimens were examined; it is quite
certain that the prothoracie leg discs reach a given stage of
development sooner than do the meso- and metathoracic leg
discs. ‘This point is referred to again helow. The peduncle of
each sac passes forwards close to the ventral body-wall to end in
the ventral hypoderm of the prothoracic segment slightly to one
side of the median line. At the point where the peduncle
reaches the hypoderm the chitin of the body-wall is traversed by
a minute fissure elongated transversely to the longitudinal axis of
the body. ‘The fissure in the chitin is, of course, due to the dis-
continuity of the hypoderm over « small area brought about by
its invagination to form the peduncle of the sac. The fissure
passes obliquely through the chitin, so that im a longitudinal
section through the peduncle and fissure the two form a practi-
cally continuous line. I am unable to offer any explanation of
the obliquity of the fissure. Similar fissures occur opposite the
peduneles of the other leg discs and opposite those of the wing
and haltere discs,
One specimen which, when killed, was approaching its time of
pupation, showed the method of evagination of the prothoracic
lees. ‘'he united imaginal sacs had grown forwards between
the peduncles of the sacs so that the inner two-thirds of the
peduncles were united by two closely approximated horizontal
hypodermal sheets, a dorsal and a ventral, the space between the
two sheets being continuous posteriorly with the cavity of the
united imaginal sacs. The central cavities of the inner two-
thirds of the peduncles had not remained separate, but were
probably represented by the extreme lateral regions of the space
between the dorsal and ventral hypodermal sheets. There can
be little doubt that eventually the whole length of the peduncles
becomes united by the dorsal and ventral hypodermal sheets;
thus, eventually, there is only one imaginal sac with a single
‘broad, dorso-ventrally flattened, peduncle, and containing the two
prothoracic leg dises. Probably. when evagination is to take place,
the dorsal and ventral walls of this flattened peduncle separate,
and the sac evaginates through the space between them. In the
specimen from which the above particulars were obtained the
meso- and metathoracic imaginal leg sacs showed no sign of union,
though they almost certainly undergo the same process at; a later
stage. In all larvee examined the degree of development was
greatest for the prothoracie leg discs, less for the mesothoracic,
and least for the metathoracie. .
ANATOMY OF 'THE GOUT-FLY OF BARLEY. 803
The mesothoracie leg discs lie slightly posterior and ventral’ to
the prothoracie discs; their peduncles pass forwards to the
ventral hypoderm of the mesothoracic segment. In the speci-
mens studied the mesothoracic imaginal leg sacs did not
communicate with one another, though they probably do so at a
later stage.
The metathoracic leg dises are posterior to the mesothoracic
and rather more lateral and dorsal. They are united to both
the wing and haltere dises by strands of tissue accompanied by
small trachese derived from the third lateral tracheal trunk.
The figure given by Lowne (p. 84) of the wing, haltere, and meta-
thoracic leg discs of the larva of Calliphora approximates very
closely to the condition found in Chlorops.
In a fairly mature larva a small imaginal dise can be found
arising just posterior to the anterior stigmatic trunk on each side
at the point where this traverses the nypoderm. In the most
advanced stage so far examined these discs are only shallow
invaginations of imaginal cells and cannot be said to consist of
peripodal cavity, peripodal membrane, and imaginal disc. They
may, however, and probably do, increase in complexity towards
the end of larval life.
The wing rudiments are the largest of all the imaginal dises;
they lie mainly in the mesothoracic segment, but their posterior
rounded ends extend into the anterior region of the metathorax.
Each lies mainly dorsal and internal to the dorso-lateral tracheal
trunk of its side, but anteriorly it narrows and passes dorsally
over the trunk to the external side of the latter. Its narrow
anterior end is prolaced into a short peduncle which passes to
the dorso-lateral hypoderm in the posterior region of the meso-
thorax.
The haltere disc lies ventral and external to the dorso-lateral
tracheal trunk, the bulb of the disc overlying the base of the
third lateral tracheal branch. Its peduncle joins the hypoderm
in the mid-lateral region of the metathorax.
The innervation of the spiracular, wing and haltere discs has
not been followed out.
A minute trachea accompanies the peduncles of all tlie thoracic
imaginal discs.
Imaginal Discs of the Mouth Parts.
There are two pairs of imaginal mouth rudiments. The larger
pair, the maxillary dises, lie one on either side of the anterior
region of the pharyngeal mass. ‘They are broad posteriorly, but
taper off anteriorly into peduncles which are attached to the head
capsule just latero-dorsal to the mouth-opening. Posteriorly
they appear to be united dorsally to the hypoderm overlying the
pharynx, which is continuous posteriorly with the cephalo-
pharyngeal band. They are united ventrally by a hypodermal
sheet passing below the pharynx;
804 MR, J. G. U. FREW ON 'THE LARVAL
Below the anterior region of the pharynx there are two
extremely small discs which appear to consist of solid longitudinal
rods of embryonic cells lying close together near the median line.
They are attached posteriorly to the hypodermal sheet uniting
the maxillary discs below the pharynx: they are the labial discs.
Their peduneles are attached anteriorly to the median ventral
hypoderm of the head immediately posterior to the mouth-
opening and some little distance anterior to the opening of the
salivary duct in the pharynx.
I have found no mandibular imaginal dises. These are also.
absent in the Blow-Fly (13) and the House-Fly (4). According
to Lowne (p. 83) Kunckel d’Herculais described and figured
mandibulsr discs in the resting larva of Volucella, but I have
been unable to find either the figure or the statement in Kunckel
d’ Herenlais’ work (12).
Imaginal Discs of the External Genitalia.
Immediately in front of the anus there are two imaginal sacs:
lying against the ventral hypoderm in the median line, each
sac containing twe imaginal] discs, one on each side in its anterior
region, The posterior sac is the larger; it has a very short,
transversely broad peduncle, which opens out anteriorly into a
small imaginal sac, in whose antero-lateral regions lie the two
dises. ‘lhe anterior imaginal sac is similar to, but rather smaller
than, the posterior. Both sacs lie in the mid-ventral region of
the eleventh body-segment (8th abdominal). The posterior dise
develops before the anterior one, being well formed when the
latter is a mere hypodermal thickening. It is possible that
differences exist in the formation of these disesin g and 9 larva,
which would enable one to detect the sex of a larva, but such
differences have not yet been. observed.
From the examination of two specimens in which the anterior
dise was in the form of a hypodermal thickening, I am inclined
to believe that the paired discs are from the commencement of
their formation enclosed in a single common imaginal sac, and
that this single sac with its single peduncle is not formed by the
fusion during later stages of larval life of two originally separate
dises. The evidence on this point, however, is by no means
conclusive.
Imaginal Discs of the Abdominal Segments.
In the most mature larva examined the abdominal discs were
in the form of two small lenticular thickenings in the dorso-
lateral hypodermis on each side about the middle of each seg-
ment. ‘The cells of these discs originate by the amitotic division
of an ordinary hypoderm cell; the nuclear membrane dis-
appears, and the nacleoplasm splits up into numerous little masses.
each of which becomes the nucleus of an imaginal cell. It is
possible that the same process goes on in other regions of the
ANATOMY OF THE GOUT-FLY OF BARLEY. 805
hypodermis at a later stage, but it has only been actually
observed in the dorso-lateral regions. The abdominal imaginal
dises, of course, never become invaginated, and therefore have
no imaginal sac and no peduncle,
The Dorsal Vessel.
The dorsal vessel is a tubular organ lying close beneath the
dorsal hypoderm in the median line, its posterior end lying at
about the junction of segments 10 and 11. From here it runs
forwards as far as the anterior region of segment 6, where it dips
ventrally to the brain to take the course already described in
connection with the central nervous system, etc. The part of
the tube lying in segment 10 is relatively wide, but it narrows
considerably as it passes forwards into ‘segment 9. The wide
posterior region may be defined as the heart; the portion
anterior to segment 6 may be termed the aorta. The ostia are
confined to the cardiac region, but I have been unable to make
certain as to their number and position. The posterior end of
the heart is slightly anterior to the posterior transverse com-
missure. The latter is suspended from the dorsal integument by
a delicate double transverse membrane, the space between the
two layers being wide dorsally but narrowing ventrally where
the membrane fuses with the commissure. From the middle of
the anterior membrane a delicate ligament passes forwards to
become attached to the ventral wall of the posterior end of the
dorsal vessel. At about the level of each pair of pericardial cells
(see below) in segments 10--6 inclusive the dorsal vessel is
attached to the dorsal hypoderm by a delicate ligamentous
strand. :
Pericardial Nephrocytes, and Alar Muscles.
On each side of the dorsal vessel there are 26 pericardial cells
(nephrocytes) situated as follows :—
In each of segments 10, 9, and 8, four cells on each side.
In segments 7 and 6, five cells on ench side.
In segment 5, four cells on each side.
The cells lie in pairs, one on each side of the vessel, exactly
opposite each other. Seen in surface view in the living larva
the cells are perfectly circular; in transverse or longitudinal
section they are oval. They are all uninuclear. In the majority
of cyclorrhaphous Jarve the pericardial nephrocytes are of two
kinds; posteriorly there are a few large cells somewhat separated
from each other, while anteriorly the cells, though similar to the
posterior ones in structure, are smaller, more numerous, and
closely crowded (see Keilin, 1917, for a review of the subject),
in Chlorops, however, there is no such distinction, the cells being:
all alike.
In connection with each pericardial cell (with the exception of
the four anterior pairs which lie alongside the ventrally-directed
portion of the vessel) there is an alary muscle. The alary muscles.
Proc. Zoon. Soc.—1923, No. LIII. D3
806 MR. J. G. If, FREW ON THE LARVA
of any segment arise close together from the mid-lateral region
of that segment, slightly ventro-lateral to the dorso-lateral
tracheal trunk. Each arises by a short dorsal and a short
ventral root, which unite to form a very thin strap-like muscle
which passes inwards and dorsally below the dorso-lateral
tracheal trunk to the pericardial cell to which it is related. Here
it: splits into dorsal and ventral layers which are inserted
respectively into the dorsal and ventral walls of the dorsal vessel,
and between which lies the pericardial cell. The dorsal and
ventral layers do not retain their narrow strap-like form but
spread out fanwise in the antero-posterior direction, so that the
anterior ends of their insertions into the heart extend as far
forwaris as the posterior ends of the insertions of the alar muscles
next in front of them. The figures given by Imms (8, figs. 9
& 10, PI. ix.) show the arrangement of the alar muscles in rela-
tion to the dorsal vessel in Anopheles, and the arrangement in
Chloraps is almost identical.
Ventral Nephrocytes.
In Muscid larvee the ventral nephrocytes usually form a trans-
verge loop of cells passing below the gut, each end of the loop
being attached to one of the salivary glands at the junction of
the glandular region and duct. In Chlorops, however, the ventral
nephrocytes consist of two separate masses each of seven or eight
cells, the groups lying one at the anterior end of each salivary
gland between the gland and the proventriculus and attached to
the gland at the region of junction of the duct with the glandular
portion. These cells are very much smaller than the dorsal
nephrocytes and are binucleate; they also differ from the dorsal
nephrocytes in several cytological details which will not, how-
ever, be considered here.
(Hnocytes occur in each of abdominal segments 1 to 7 inclusive,
as two gioups of about five or six cells, the groups lying one on each
side of the segment close against the hypoderm in the mid-lateral
region. The cells are all uninucleate with rather dense, finely
granular, and nonvacuolated protoplasin.
Small Genocytes (8) appear to be absent.
The Fat Body consists of lobes and chains of large cells lying
between the various organs of the body. ‘There is no fat body
in the head and very little in the thoracic segments.
Integument,
The chitin consists of two layers, an outer thin and dark
coloured, and an inner thick and light coloured. The surfaces
of the two layers where they come in contact are very minutely
serrated, the serrations engaging with each other. The hori-
zontal split by which the puparium will open is marked out in
the third instar larva by what is apparently a closed fissure,
_ appearing in sections as a thin dark line through the chitin,
which passes horizontally round the anterior end of the larva
ANATOMY OF THE GOUT-FLY OF BARLEY, 807
anil back on each side to the posterior region of the metathoracic
segment. ‘The peduneles of the wing discs reach the hypoderm
ata point immediately opposite this fissure. ‘The vertical splits
which pass up and down the lateral regions of the puparium
from the posterior ends of the lateral splits do not appear to be
similarly marked out in the third instar larva.
Gonadial Rudiments.
__ The rudiments of the ovaries or testes lie one on either side of
the fifth abdominal segment in its posterior region. They are
solid oval masses of cells with the long axis parallel to the long
axis of the body, and are attached to the neighbouring fat body.
As I have not so far been able to distinguish a Javval ovary from
a larval testis, it would be quite useless to give any account of
the structure of the gonadial rudiments,
Muscular Systen.
The details of the muscular system have not been worked out.
The arrangement of the muscles probably approximates very
closely to their arrangement in Jfusca domestica which is de-
scribed by Hewitt (4). ;
C. Other Immature Stages of Chlorops.
The Puparium (text-fig. 12).
Length 5:4 mm.; maximum breadth 1:5 mm. Colour ex-
tremely variable; all shades between pale yellow to very dark
brown. ‘Thoracic segments slightly dorso-ventrally compressed,
much wrinkled transversely and obliquely, always darker coloured
than the abdominal segments; apices‘of wrinkles usually appear-
ing dark brown or black, the spaces in between being lighter.
The last two abdominal segments are also considerably wrinkled
and darker coloured than the others. The other abdominal
segments are also transversely wrinkled, but the wrinkles are not
pronounced or close together and are not darkened.. The larval
cephalo-pharyngeal skeleton lies pressed against the ventral wall
in the anterior end of the puparium. ‘he larval head is com-
pletely withdrawn within’ the prothorax and is quite invisible.
The larval stigmata stand out clearly. There are no pupal dorsal
respiratory trumpets projecting through the dorsal wall of the
puparium (cf. Keilin, 1917).
Hor the exit of the imagines the anterior end of the puparinn
splits into dorsal and ventral halves by means of a horizontal
‘split running round its anterior end and backwards along each
side as far as the anterior region of the first abdominal segment.
From each posterior end of this split a dorsal and a ventral split
extends round the anterior end of the first abdominal segment.
The extent of these transverse splits is very variable; sometimes
they are very short, sometimes the dorsal ones meet in the mid-
53*
808 . MR. J. G. U. FREW ON THE LARVAL
dorsal line and the ventral ones in the mid-ventral line, so that
when the imago emerges the whole of the thoracic region of the
puparium falls off in two halves. Usually the ventral splits are
longer than the dorsal. When the pupariuin splits the larval
spiracles are situated on the dorsal half of the split thorax, very
slightly dorsal to the split.
The First Larval Instar is extremely similar to the third imstav.
The head and cephalo-pharyngeal skeleton ave identical in form
to those of the third instar; the arrangement of the tracheal
trunks is the same, but the smaller branches of the tracheal
system ramify less extensively, many being either not yet formed
or not filled with gases so as to be visible. On emergence from
the ege the larva is completely apneustic. The dorso-lateral
tracheal trunks extend from the anterior region of the second
body-segment to the posterior region of the eleventh body-
segment. There is no indication whatever of anterior spiracles.
Posteriorly, in the position occupied in the second and third
instars by the posterior stigmatic papillw, there are two stout
and prominent papille with a truncated apex. These show no.
Text-figure 12.
‘ni,
C. Eeniopus. Lateral view of empty puparium.
trace of stigmatic openings, and the cdorso-lateral tracheal trunks.
do not extend into them; four rather stout pointed sete radiate
outwards from the dorgo-lateral border of the apex of each of
these papillee.
In the first larval instar there is no difference in shape between
the thoracic and abdominal denticles, all being of the approxi-
mately semicircular shape found on the abdominal segments of
the third instar. Their distribution is slightly different from
that in the third instar in that they do not appear to extend
into the dorsum of the thoracic segments. Such characters as
these, however, require very careful examination with an oil
immersion. J have not yet succeeded in finding the vestigial
leg-sense organs in the first instar; if they are present, however,
they will naturally be extremely minute, and Tam not, therefore,
prepared to say definitely that they are absent.
Maximum length of cephalo-pharyngeal sclerites
~ Maximum clorso-ventral depth of sclevites
third larval instar and about 43 in the first larval instar.
= 34
~
in the
ANATOMY OF THE GOUT-FLY OF BARLEY. 809
With the exception of the above points the first instar is a.
miniature replica of the third.
The Second Larval Instar is identical in external anatomy,
except for size, to the third. A mature second instar is only
recognisable from a young third instar by the fact that in the
former there can be seen the developing spiracles and cephalo-
pharyngeal skeleton of the third instar alongside those of the
second instar itself. .
The Egg (text-fig. 13) is approximately cylindrical with its
ventral surface flattened and its dorsal surface strongly convex
transversely and slightly so longitudinally. The egg narrows
slightly at both ends. There is a definite polarity of the eggs
which are always laid on the leaf of the host-plant in a certain
way. The anterior end, which points towards the apex of the
leaf, is slightly dorso-ventrally flattened and very slightly truncate
at the apex. The posterior end is evenly rounded and not dorso-
Text-figure 13.
e
'
. ’
Anterior RE 5; Posterior
C. teniopus. A. Outline drawing of egg viewed dorsally.
B. Outline of pits on dorsal surface seen at a higher magnification.
ventrally flattened. The upper surface and sides of the egg are
ornamented with small polygonal depressions set in longitudinal
rows, which may or may not extend the whole length of the egg.
The ridges separating two adjacent depressions in a row are not
quite so high as the longitudinal ridges which separate adjacent
rows. The ventral surface of the egg bears a few longitudinal
rounded ridges. The egg is firmly attached to the plant upon
which it is laid bya transparent cementing material. No micro-
pylar apparatus has been observed.
Host- Plants.
As the question of the host-plants of Chlorops will: be con-
sidered in detail in a later paper, it will be sufficient to state
here that Agropyrwm (= Triticum) repens is the only wild grass
in which the larva has been found up to the present.
810 MR, J. G. Hf. FREW ON THE LARVAL
MernroMyYZA NIGRIVENTRIS Meq.
The full-grown larva (text-fig. 14) is about the same length as
that of Chlorops, but is rather move slender. It is almost always
a bright green colour, but occasionally almost colourless speci-
mens are found, Other prominent characteristics are the sessile
Text-figure 14.
P-Sp- aie MAT NGL aie | a.5p.
i | H i | ee
« eis Ss
—— =
liam.
M. nigriventris. Lateral view of mature larva.
posterior stigmata, the presence in the cephalo-pharyngeal skele-
ton of a median ventral piece, and the peculiar shape of the last
abdominal segment when seen laterally.
The Head (text-fig. 15) differs in several points from that of
Chlorops. The antenne are anterior instead of lateral. The
Text-figure 15.
0:2mm.
ML. nigriventris. Ventral view of larval head.
maxillary palps are situated on two prominent swellings separated
by a rather deep median fissure: the chitinous ring surrounding
the maxillary palp sense organs is incomplete antero-laterally.
The pre-oral sense organs are situated on the postero-ventral
regions of the swellings bearing the maxillary palps; each is a
ANATOMY OF THE GOUT-FLY OF BARLEY. 811
roughly triangular plate bearing four small circular sense organs,
a larger central one and an anterior and two posterior smaller
ones. The labial plate bearing the two labial sense organs
occupies the same position as in Chlorops, but differs somewhat
in shape. The position of the small circular cephalic sense
organs is shown in text-fig. 15. At each side of the oval aper-
ture, running more or Jess parallel to its lateral margin, there
are three chitinous ridges which extend from the antero-lateral
region of the mouth-opening to its postero-lateral region.
Thoracic Segments.
Instead of rows of chitinous teeth such as occur on Chlorops,
Meromyza has low transverse ridges of varying lengths, the
apices of the ridges being minutely serrated. Almost the whole
of the dorsal surface of the prothoracic segment bears these ridges,
only a narrow posterior region being free from them. Ventrally
the ridges extend across the whole breadth of the anterior
margin, but passing posteriorly they become less and less exten-
sive, so that they form a roughly triangular area, the base of the
triangle lying across the anterior margin of the segment, the
apex of the triangle lying in the median region of the segment a
little anterior to its posterior margin.
The meso- and metathoracic segments have ridges all round
their anterior borders, the ridges being numerous, closely placed,
and covering broad areas in the middle of dorsal and ventral
surfaces, but becoming much less numerous, wider apart, and
confined to a narrower area laterally.
In the antero-ventral region of each thoracic segment is a
transverse series of circular sense organs, five on each side of the
median line. ‘The vestigial rudiments of the legs are similar to
those of Chlorops.
Abdominal Segments.
There are nine abdominal segments, the ninth being rather
small with its posterior region capable of partial withdrawal into
its anterior. The anus, which is situated ventrally on the
anterior region of the ninth abdominal segment, is a narrow,
Jongitudinal slit, bounded on each side by a fairly prominent oval
swelling; there are no adjacent chitinous teeth or special sense
organs. ‘he last abdominal segment is divided posteriorly into
two lobes by a deep and broad vertical fissure; the sessile
posterior stigmata lie facing oné another, one on each side of the
fissure.
Chitinous ridges, similar to those on the thoracic segments,
surround the anterior regions of each abdominal segment except.
the last. No ridges occur round the posterior border of any
segment, ‘There are no ambulatory swellings.
Circular sense organs, similar to those on the thoracic segments,
occur in transverse rows of about ten on the ventral surface in
812 MR. J. G. H, FREW ON THE LARVAL
the posterior half of each abdominal segment. ‘There is a row
behind the anus, and their presence makes it practically certain
that what in this and the other larvee described heve (see Chlorops
above) I have called the ninth abdominal segment really is a
separate segment,
Cephalo-Pharyngeal Skeleton. (Text-fig. 16.)
This is of the same general type as that of Chlorops, and the
differences of detail can aye made out from the figures. ‘Lhe most
striking difference is perhaps the presence of a median ventral
Text-figure 16.
M. nigriventris. ateral view of cephalo-pharyngeal sclerites.
piece. This plate also occurs in Balhoptera combinata, where its
relations with the neighbouring parts are more easily made out
(see below).
Tracheal System.
The anterior stigmata are of the usual branching type and are
situated near the. posterior margin of the first ‘body- segment,
slightly dorsal to the mid- lateral line. ‘The branches are ex-
tremely short and project very little from the body-surface
(text-fig. 17).
The posterior stigmata (text-fig. 18) are, as already mentioned,
sessile and not at the apex of a stigmatic papilla; there is no
stigmatic plate and each of the three openings projects very
slightly above the body-surface. Four pairs of branching hairs
arise, as shown in the figure.
The arrangement of the tracheal trunks and branches is very
similar to that found in Chlorops. There are many differences of
detail which do not, however, merit the extended description
necessary to make them clear: one difference only is noteworthy.
In Chlorops the dorsal commissures give off no branches to the
dorsal region of the body, which is supplied by branches from the
lateral trunks. In Meromyza {text-fig. 10C) each dorsal com-
missure, except the second and the last, gives off two anteriorly-
directed branches, which ramify extensively to supply the dorsal
ANATOMY OF THE GOUT-FLY O! BARLEY. 813
tracheation of the segment in whose posterior region arises the
dorsal commissure from which they take origin. Very little of
the dorsal tracheation is supplied by branches from the lateral
trunks.
Text-figure 17.
-- ESN
BN
Sa
M. nigriventris. Anterior spiracle. ‘The dotted line indicates the
surface of the body.
Text-figure 18.
0°03 mm.
Posterior spiracie of I. nigriventris.
Pupariwum.
‘Length 5:7 mm. Maximum breadth 1:2 mm. Practically
colourless, but appears green owing to colour of pupa within.
Wrinkling as in Chlorops; wrinkles have a light golden-brown
colour. Splitting of puparium as in Chlorops except that the
vertical splits always meet dorsally and ventrally, so that the
thoracie segments always separate off completely in dorsal and
ventral halves (except that they may remain attached to the
nyraphal integument).
814 MR, J. G. W. FREW ON TILE LARVAL
BaALIoPrERA COMBINATA L.
The full-grown larva (text-fig..19) is shorter and relatively
thicker than that of Chlorops. Its cephalo-pharyngeal skeleton
is proportionately larger than in any other larva considered here.
Text-figure 19.
ss eli Wl STE °
[en ee wit
LO nim. mM.
Balioptera combinata. Juateral view of mature larva.
Its most prominent characteristics are its large anterior stigmata
and the heavily chitinized median ventral piece of its cephalo-
pharyngeal skeleton.
The Head (text-fig. 20) is chiefly remarkable for the arrange-
ment of the numerous chitinous ridges on its ventral surface,
Text-figure 20.
0-2 mmm.
B. combinata. Ventral view of larval head. (The rows of chitinous denticles
on the prothorax have been omitted.)
which is divided by them into small polygonal areas whose distri-
bution is shown in the figure. Certain of the ridges in the
region antero-lateral to the moutli-opening bear small chitinous
teeth (c.7.¢.) pointing posteriorly.
ANAVOMY OF THE GOUT-FLY OF BARLEY. 815.
The maxillary palps are borne by two fairly prominent
swellings; their circular sense organs are surrounded by chitinous
rings which are incomplete antero-laterally, and two slightly
larger circular sense organs lie just outside the gaps in the rings.
The antenne are only remarkable in that the apical joint is
almost as broad as long, instead of being markedly longer than
broad as in the other lave considered here. ‘The pre-oral sense
organs are inconspieuous, each consisting of two small circular
sense organs situated close together but not carried on any special
swelling; they are quite separate from the swellings which bear
the maxillary palps. The labial plate is again characteristic in
shape, so far as the larve considered in this paper are concerned,
and is reinarkable in bearing two pairs of Jabial sense organs, the
anterior pair being the larger. The distribution of the small
cephalic sense organs is shown in the figure, except for one pair
which are not visible in a ventral view, as ‘they are situated one on
each side of the head slightly dorsal to the mid-ventral line and
on a level with the anterior margin of the mouth-opening.
Thoracic Segments.
The chitinous denticles on the thoracic segments are consider-
ably larger than those on Chlorops. ‘Yhey occur all round the
srahenion margins of the first and second thoracic segments as
moderately broad bands. The third thoracic segment has no
anterior teeth ventrally and only.a few laterally and dorsally ;
a little in front of its posterior margin, however, it bears a single
row of teeth extending across its ernie region and up most Wee
its lateral regions. There are six thoracic sense organs on each
segment forming a transverse row behind the region bearing the
chitinous teeth. ‘he vestigial lee rudiments are similar to those
in Chlorops.
Abdominal Segments.
There are nine abdominal segments, the last bearing the anus.
ventrally and the posterior stigmatic papillae posteriorly. The
morphology of the jast two body-segments is rather obscure, and
this point is dealt with below. The arrangement of the chitinous
teeth on abdominal segments 1 to 7 is practically identical.
Hach bears near its anterior and posterior margin a transverse
band of chitinous teeth, which are from two to four rows broad
ventrally, but become less numerous as they pass up the lateral
regions of the segment. In the first four abdominal segments
the teeth are continued over the dorsum as a single anterior and
posterior row on each segment, but on the other segments they
are not continued above the mid-lateral line. The posterior
teeth on the seventh abdominal segment form only a single row
ventrally. Ventrally there is a narrow space between the
posterior rows of teeth of one segment and the anterior rows of
the next segment behind it, and this space, which corresponds
816 MR. J. G, WH, FREW ON THE LARVAL
largely to the thin intersegmental membrane, is capable of a
slight degree of protrusion to form a ventral pseudopod.
Behind the seventh abdominal segment there appears to be one
large segment bearing the longitudinal slit-like anus in its mid-
ventral region, and a very small postero-dorsal segment, cou-
sisting mainly of the two stigmatic papille and capable of partial
withdrawal into the apparent segment in front of it. ‘This, how-
ever, is not the true nature of this region. A study of the
musculature shows that the large segment consists of two seg-
ments, abdominal segments 8 and 9, and the small posterior
stigmata-bearing region is almost certainly to be regarded merely
as 2 specialized region of segment 9. Some indication of the com-
posite nature of the large segment is available without a study
of the musculature. In front of the slit-like anus is a transverse
row of about 17 sharp, posteriorly-directed, chitinous teeth, and
behind it is a small patch of about seven similar teeth pointing
forwards. It seems probable that the anterior row represents
the posterior band of teeth of segment 8, and the posterior
patch the anterior band of segment 9, the anus lying therefore
between the two segments. ‘The anus is not bounded by lateral
swellings,
Cephalo-Pharyngeal Skeleton. |
The form of this is sufficiently shown in text-fig. 21, and only
two points require special mention, the ventral wall of the pharynx
Text-figure 21.
ml.
B. combinata, Yateral view of cephalo-pharyngeal sclerites.
and the median ventral piece. The ventral wall of the pharynx in
the region of the pharyngeal sclerite has distinctly the appearance
of being longitudinally ribbed when seen in surface view. As I
have not examined sections of the larva, 1 can give no indication
of the true nature of these ribs. The median ventral sclerite is
a somewhat shield-shaped chitinous plate with its lateral margins
bent strongly upwards. Its anterior margin shows a central]
curved emargination bounded on each side by a rather prominent
process from which a cord, apparently of a tendinous nature, goes
forward to the ventral process of the corresponding mandible.
Posteriorly the plate becomes less heavily chitinized and appears
ANATOMY OF TUE GOUT-FLY OF BARLEY. 817
to be serrate; into it are inserted the depressor muscles of the
mandibles. ‘Che median ventral piece bears two vather prominent
circular sense organs, one on each side of its anterior region.
Tracheal System.
The anterior spiracles (text-fig. 22) are borne dorso-laterally
by the first thoracic segment in its posterior third and project
forwards close to the surface of the segment. They are very
prominent structures, consisting of a rather broad dorso-ventrally
flattened central stem, bearing about 13 lateral branches. Along
the median region of the central stem are a series of small
fissures (72), which apparently pass right through the stem, heing
only covered by the delicate membrane by which the whole struc-
ture, except the apices of the branches, is covered.
Text-figure 22.
B. combinata. Anterior spiracle.
The posterior spiracles (text-fig. 23) are carried at the apices of
two prominent posterior stigmatic papille. There is no definite
chitinized stigmatic plate. The form of the stigmata and the
branching hairs may be seen from the figure. The stigmatic
sear (s.c.), which I have failed to find in Chlorops. and Meromyza,
is fairly prominent in Balioptera, but it does not occupy the
position in which it is found (as the “ Button”) in most Muscid
Jarvee. 1t lies on the median side of the stigmatic papillz some
distance below the apex. »
The distribution of the traches is essentially similar to that in
Chlorops, the chief difference being the branches given off from
818 MR. J. G. H. FREW ON THE LARVAL
the ten dorsal commissures. the first and last of which are, as
usual, thicker than the others.. The tenth commissure gives off
no branches: the first. gives off at each end a moderately large
anteviorly-directed branch; the second loops strongly forwards
over the first and gives off a few branches anteriorly; the third
is also strongly looped forwards and gives off a few branches
anteriorly. Commissures 4 to 9 inclusive are similar and are
represented in text-fig. 10B. They are roughly transverse and
give off branches both anteriorly and posteriorly, each commissure
supplying the dorsal tracheation of the posterior half of one
segment and the anterior half of another. ‘The dorsal commissures
themselves occupy approximately the same positions with regard
to the body-segments as they do in Chlorops.
Text-figure 23.
B. combinata. Posterior spiracle. Tho dotted line represents the edye
of the apex of the stigmatic papilla.
Puparium. Length 3°2 mm. Maximum breadth -9 mm.
Colour golden brown; anterior and posterior ends much
wrinkled and blackish. ‘The thoracic segments are much flattened
dorsally, so that their dorsal surface shows a large shallow con-
cavity, from the antero-lateral corners of which the prominent
anterior spiracles of the larva projects forwards.
For the emergence of the imago the puparium splits in the
usual manner along the lateral regions of the thoracic segments,
which in this case correspond to the lateral margins of the dorsal
concavity. ‘Lhe posterior ends of the lateral splits are joined by
1. dorsal transverse split, and the whole of the dorsal region of the
thoracic segments splits off as a roughly quadrangular plate,
which may or may not become completely detached from the
puparium. ‘The anterior spiracles are attached to the dorsal
plate.
ANATOMY OF THE GOUT-FLY OF BARLEY. 819
Chloropisca glabra Meig.
This fly was bred out in considerable numbers from various
grasses and has also been found infesting Winter Wheat and
Barley. It was originally intended to describe here the larval
form. The larva is, however, so extremely similar to that of
Oscinis frit that it has been decided to defer the description of the
Chloropisca until a detailed comparison of these two larve can be
made. ‘The only differences so far observed are the somewhat
larger size of the Chloropisca larva and its rather more yellow
colour; such differences are obviously unsatisfactory.
The chief winter host of Chloropisca is Lolium perenne.
Winer Hosr-Piants or CuLorors T-4Nn10Pcs, Mreromyza
NIGRIVENTRIS, AND BALIOPTERA COMBINATA.
After a considerable amount of purely qualitative grass
examination had been done, an attempt was made to reduce the
work toa quantitative basis by bringing in samples of the various
grasses, examining each shoot in the laboratory, and finding a
percentage figure of infestation. The figures were, however,
found to vary very greatly in samples of the grasses taken from
different localities and are not, I think, worth giving. I consider
that to obtain a reliable infestation figure for a grass in this way,
at least 20,000 shoots would have to be counted, taken in samples
of about 1000 shoots from different localities. ‘This is obviously
impossible when one is examining a considerable number of
(different grasses.
Chlorops teniopus has only been found in Agropyrum repens
among the wild grasses examined. Jleromyza nigriventris occurs
in A. repens, Festuca ovina, and Alopecurus pratensis; Balioptera
combinata occurs in A. repens, Mestuca elatior, Loliwm perenne,
Holeus lanatus, and Agrostis alba. ‘The following grasses have
also been examined but do not appear to function as winter hosts
for any of the above three species of flies:—Loliwm italicum,
Poa pratensis, P. trivialis, P. annua, Agrostis vulgaris, Alopecurus
agrestis, Arrhenatherum avenaceum, Anthowanthum odoratum,
alvena pubescens, Cynosurus cristatus, and Dactylis glomerata.
Winter Barley and Wheat are also attacked by all three
species; the conditions under which they are attacked by Chlorops
will be dealt with in detail ina later paper. The effect on the
host of either J/eromyza ov. Balioptera is very similar to that of
Frit Fly. The larve live in the base of the shoot and destroy
the growing point, causing also. the yellowing and death of the
central leaves by destroying their basal regions. Winter is passed
in the larval state. In spring the Jarva turns round in the grass
shoot until its head is directed upwards, ascends a short distance
up the shoot, and then pupates.
820 MR. J. G. H. FREW ON THE LARVAL
SUMMARY.
The external and internal anatomy of the mature larva of
Chlorops teniopus ave described in detail.
The external anatomy of the first and second larval instars is
compared with that of the third instar and points of difference
noted. ‘The puparium and the egg are also described.
The external anatomy of the third larval instar and the
puparium of JMJeromyza nigriventris and of Balioptera combinata
ave described and compared with the corresponding stages of
Chlorops taniopus.
The winter host-plants of the three species are given, together
with a list of eleven grasses which have been examined but do
not serve as winter hosts for any of the above species.
ReEererence LErrerine
or Trexr-Fieurns.
a. antenna, m.pl. mediante plate of cephalo-pharyn-
a.sp. anterior spiracle. geal band.
b. bulb of stigmatic trunk. n.v.l. median ventral sclerite.
bh. branching hair of posterior p.g. proventricular ganglion.
spiracle. ph. pharynx.
e.p.b. cephalo-pharyngeal band. p.w. proventricular nerve.
c,p.s. eephalo-pharyngeal sclerites. ‘p.o. pre-oral sense organ.
e.. cephalic chitinous ridges. pr. proventriculus.
ert. “teeth” on cephalic chitinous p.s. pharyngeal sclerite.
ridges. p.sp. posterior spiracle.
c.s.o. cephalic sense organ. r. ridge on bulb of stigmatic trunk.
d.m, dilator muscles of pharynx. s. spiracular opening.
d.v. dorsal vessel. s.c. stigmatic scar.
f. fissures in stem of anterior S.m. sheathing membrane of anterior
spiracle of Balioptera. spiracle.
fe. felted chamber of spiracle. s.p. stigmatic plate.
hy. hypodermis. s.pr. suspensory process of Weis-
i.s. intermediate sclerite. mann’s ring.
i.3.p. peduncle of cephalic imaginal sac. t.c. transverse commissure.
1. labinn. t.s.o. thoracic sense organ.
1.6. lateral tracheal branch. v1, vestigial lex sense organ.
Lf. lateral flap of cephalo-pharyngeal v.s. vertical suspensor of dorsal
band. vessel.
lp. labial palp sense organ. v.t. vestigial stigmatic trunk.
1.¢. dorso-lateral tracheal trunk. w.r. Weismaun’s ring.
m. mandible. XX. point: of junction of salivary duct
. median nerve.
with pharynx.
m.niw, muscle nucleus. Y. opening of dorsal vessel inte
my. maxillary palp. supra-cesophageal sinus.
Lisr or Lrrerarurn.
(Only those titles are given which are referred to in the text.)
1. Becrer, R. 1910. “ Zur Kenntniss der Mundteile und des Kopfes der Dipteren-
8. Hennecuy, L, F. 1904.
4, Hewitt, C. G. 1914,
Larve.”
B. No. 4.
ser.
Les Insectes.
The House-F ly.
Zool. Jahrb. Abt. fiir Anat. u. Ont. der Thiere, Bd. xxix.
2. Carpenter, G. H., and Potzagp, F. J. 38. 1918.
Spiracles in the Larva of Hypoderma.”
“The Presence of Lateral
Roy. Ir. Acad. Proce. vol. xxxiy.
Paris.
Cambridge.
ANATOMY OF THE GOUT-FLY OF BARLEY. 821
5. HWormennn, M. 1904. “ Zar Morphologie des Insektenkopfes: II, Einiges tber
die Reduktion des Kopfes der Dipteren-Larven.” Zool. Anz. Bd. xxvii.
6. Dr MrtsErn, J.C. H. 1895. - “ Ueber Zusammengesetzte Stigmen bei Dipteren-
larven, nebst cinem Beitrag zur Metamorphose von Hydromyza livens.”
Tijd. v. Entomol, vol. xxxviil.
7. Imus, A.D. 1907. “On the Larval and Pupal Stages of Anopheles maculi-
pennis Meigen.” ‘The Journal of Hygiene, vol. vii. No. 2.
8. Inns, A. D. 1903. Idem. Parasitology, vol. i. No. 2.
9. Kerntn, D. 1915. “ Recherches sur les Larves de Diptéres Cyclorhaphes.’’
Bull. Se. de la Fr. et de la Bel. 7¢ sér. t. xlix. fase. 1-2.
10. Ket, D. 1917. “Recherches sur Jes Anthomyides & Larves Carnivore.’”
Parasitology, vol. ix. No. 3.
11. Kerri, D. 1921. “On the Calcium Carbonate and the Calcospherites in the
Malpighian Tubes and the fat body of Dipterous Larvee and the Eedysial
Elimination of these products of excretion.”’ Q. J. M.S. vol. Ixv. No. 260,
n. s.
12. Kuxexern p Hrrcunars. 1875-1881. Recherches sur Vorganisation ct le déve-
loppement des Volucelles, Insectes diptéres de la famille des Syrphides-
Paris. :
13. Lowng, B. T. 1890-95. Anatomy, Physiology, Morphology, and Development
of the Blow-Fly. Tendon.
14. Mrass, L, C., and Hammonn, A. R. 1892. “The Development of the Head of
the Imago of Chironomas.” 'Try. Linn. Soc. Lond. vol. v. pt. 8, 2nd ser. Zool.
15. Prarz, If. S. 1900. “The Embryonic History of the Imaginal Dises in Aelo-
hagus ovinus L.., together with an account of the earlier stages in the
evelopment of the Insect.” Proc. Boston Soc. Nat. Hist. vol. xxix.
No. 13.
16. Weismann, A. 1865. “ Die Entwicklung der Dipteren in Ti, nach Beobacht-
ungen an Chironomus sp., Musca vomitoria, und Pulea canis.” Zeitschr.
f. Wiss. Zool. Band xili.
Proc. Zoou. Soc.—1923, No. LIV. o4
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ON TIE ISOPOD GENUS LIGIDIUM. 823
40. A Revision of the Isopod Genus Ligidium (Brandt) *
—Crustacea. By Harotp Gorpon Jackson, F.Z.S.,
Birkbeck College, University of London.
{Received July 13, 1923: Read November 6, 1923.]
(Text-figures 1-10.)
ConrEnrTs.
Page
(1) Introduction .. ws stPho's cheraeec nas adh craves ene 2S.
(2) Generic and Spnatite Olbmnevaias, Coane eos UrnboBEd RooaaGoe | ert
(3) Key to the genus Ligidiwm w....0....cccseseeceeessceeeeeeee 828
Gy eiherenusmiigedeum) nae ee ee S28
L. Ligidiwm hypnorwm 0.0... cccsveveecseeeresteeseecesene 829
2. Digidirm fragile oo... ..c.cccseccnessssecceceseseoseeee SOL
Sy TENG AUCTTD GAREXOIG can suocos daonsanbnnboonobandeoboncacee (hi)
Ae Teiigiiai Lat... cccccc ses cceetcccesonsicsesssecee veees D4
5. Ligidium germanicwm .....ccccceccccseeseeccsceeess es. 885
6. Ligidium japonicutr......ccccccceccceereerecees ves 836
7. Ligidiwm bosniense .........scc cee ceecse tec cce eee eeeess 837
8. Ligidiam nodilosum... 10... ccccce cen cececeeee eee scess. 837
9. Ligidium pee aei STN ox ai TREES OOS
10. Ligidiun (lyphiatgidunyie COXCHED — sactereoncon ter 838
(1) Introduction.
The genus Ligidiwm was separated from Ligia by Brandt in
1833 with this dingnosis ; “ Articulus appendicis caudalis apicalis
exterior articuli basalis apici interius Bera processul proprio
ex articuli basalis apice prodeunti insertus.” It is well defined
in structure and habitat from Ligia, although nearer to that
genus than to any other Terrestrial Isopod and linked to it by
the newly-described genus Ligidioides Wahrberg (1922); and
there is but slight range of structure within it. Budde-Lund
(1885) gives five species which he describes in Latin with the
utmost brevity and without figures. The most important and
indeed often the only character given is the form of the
uropod, but as the appendage is brittle and often lost, it is
sometimes impossible to make a certain identification from his
description. I only retain two of these species, but several
good species have been added since Budde-Lund’s work, and his
collection contains specimens, mostly unidentified by him, which
have enabled me to describe all but four species. The revision
of this genus has been greatly aided by a recent paper by Verhoeff
(1918) dealing with the European species, in which much-needed
new characters are proposed, and a new subgenus, Z'yphloligidium,
is set up, on, to my mind, amply sufficient grounds, to contain
Carl's cave-dwelling Ligidium caecum.
* This paper is the second of a series. :
Se OA
824 MR. H. G. JACKSON ON THE
(2) Generic and Specific Characters.
T have not found any marked sexual dimorphism to occur in
Ligidium, the difference between the sexes being confined to the
modified pleopods of the male. In some species of Ligiw the
sutures separating coxal plates from tergite are differently
marked in the sexes. It is interesting to note that no such
difference is to be found in this genus, and that the sutures are
marked, if at all, in an entirely different manner by fine semi-
circular grooves on the last four thoracic somites of both sexes
(text-fig. 7, a).
The proportions of the body and the size vary remarkably
little.
The colour of preserved specimens is of little value, but the
distribution of the pigment is sometimes. characteristic and
always worth noting.
The general surface of the body is remarkably smooth and
polished in all but two species, which have a roughened appear-
ance owing to the presence of scales or knobs.
Cephalon. The line of the epistome is continuous. Above and
between the insertion of the antenne it forms a downwardly-
directed V, which differs in length and sharpness in different
species. When prolonged and sharp it commonly projects
forward to form a slight triangular rostrum. ‘This condition is
termed ‘ produced ” in the following descriptions (text-fig. 7, b).
The eyes are moderately large in all species of the subgenus
Ligidium, and occupy the lateral corners of the head. In front.
of the hind margin of the cephalon is a more or less deep trench
(‘‘ transverse groove ”), opening to the cheek more or less behind
the eye on each side. The two pear-shaped pits on the forehead
of Liyia are represented in this genus by grooves (‘frontal
grooves”) which originate behind the eyes in the transverse
groove, run forwards on the inner side of the eyes, and tum
inwards on the top of the head torun towards each other parallel
to the transverse groove. They never meet in the mid-line, but
end abruptly. ‘The demarcation of these grooves has some
systematic value. (Text-figs. 3,6; 7, 6.)
The thoracic somites vary little in general form. The hind
margins of the first three are more or less straight, the fourth is
slightly concave, and the remainder more deeply so owing to the
backward production of the lateral corners. These ave never,
however, much produced. The first somite usually differs from
the remainder, as has been pointed out by Verhoeff (1901). In
many species the tergite is dinted on the postero-lateral corner
of each side, and the dint may take the form of a wide shallow
pit or of a shallow groove forming a “‘re-entrant” fiom the hind
border of the tergite (text-fig. 3, c&d). In either case it super-
ficially appears as if the tergite had been carelessly nipped by a
fine pair of forceps. This structure is referred to hereafter as
the ‘lateral depression.” When the depression is extended to
ISOPOD GENUS LIGIDIUM, 825
the hind border of the tergite, there is often present on its hind
edge a patch of stiff bristles arranged in about three transverse
rows, forming, as it were, a thick fence of stakes at the entrance
to the little valley. Verhoeff (1918) has suggested that. this
‘pparatus may be used as a comb for cleaning the antennee, but
confirmatory observations have yet to be made.
The edges of the tergites are beset with “ Schuppenborsten ”
similar to those of Ligia, as described by Wahrberg—that is to
say, the bristle usually projects slightly, but never conspicuously
from the scale; itis sometimes not easy to detect.
The abdomen (metasome) is always abruptly contracted. The
first two somites are covered by the last thoracic somite, and are.
without drawn-out pleural plates, but the remaining somites
have these moderately drawn out. The terminal somite is very
similar in form in all species. The hind border ig arcuate or
very bluntly angled in the median line; it is notched above the
insertion of the uropods to a greater or less extent in different
Species, but no distinctive angles or spines are developed.
Appendages.
The antenne (antennules) are of three segments, the distal: of
which is almost vestigial. 'They always project beyond the front
of the head in this genus. Sensory .bristles are found on each
segment and a terminal bunch on the last of » few short stiff
rods. (Text-fig. 2, a.)
2nd antenne. Kive segments and an obscurely segmented
flagellum of not more than 15 or less than 9 segments, ending in
a dense brush of sete. In the subgenus Z'yphloligidium the
flagellum has 19-23 segments. The number of segments on
the flagellum appears to increase in number till fully adult.
(Text-fig. 2, b.) .
hight mandible. Three strong biting teeth; large lacinia
mobilis beset with many strong bristles, but not chitinized ;
3 hairy sete between biting teeth and molar tubercle on base
of lacinia mobilis. Molar tubercle not high-crowned, and bearing
a dense row of setose bristles on its free posterior edge. A
bunch of long non-setose bristles between lacinia mobilis and
molar tubercle. (Text-fig. 1, a &c.)
Left mandible. Three or four strong biting teeth; lacinia
mobilis with 3 chitinized teeth; 3 or 4 hairy bristles between
biting teeth and molar tubercle on base of lacinia mobilis.
Molar tubercle high-crowned ; its surface strongly ridged.
(Lext-fig. 1, b & d.)
Ist maailla. Lacinia exterior with 4 large teeth and 3 or 4
small ones. Lacinia interior with three very stout hairy bristles,
the two inferior of which are equal and ‘larger than the upper-
most one. (Text-fig. 1, e& f.) ie :
2nd maxilla. Obscurely divided into two lappets, the smaller
‘on the outer side; on the inner side are seen two richly setose
826 MR. UW. G. JACKSON ON THE
bristles, the upper of which is small and rounded, the lower
larger than the upper and blade-shaped, (‘Text-fig. 1, y.)
Mavillipedes. Mndopodite distinctly divided into 3 segments,
the distal one ending in a blunt cone; on inner side of all but
proximal one a prominent bunch of about 7-9 bristles. The
Text-figure 1.
Ligidium hypnorum. a & oc, right mandible; b & d, left mandible ;
e & f; st maxilla; g, 2nd maxilla; 1, maxillipede.
inner blade densely covered with plain and setose bristles on the
apex. (Text-fig. 1, /.)
Percopods. 'V'ypical in form and without distinctive features..
The bristles are of simple form, ending in two spikes with a
median small “hair.” A single bristle projects beyond the
ISOPOD GENUS LIGIDIUM. 827
unguis in all species but ZL. japonicum, in which, according to
Verhoef, the 7th foot has four ciliated bristles projecting beyond
the unguis.
Pleopods. In the male the Ist and 2nd pairs are modified.
The 1st endopod is drawn out on the inside into a process upon
which is set a variable but small number of sete. The Ist
exopod has also sette on the postero-median corner, The 2nd
endopod is of two segments and greatly elongated. The inner
edge is thickened, and the outer appears to be folded over in some
forms. At the end of the distal segment this fold sometimes
becomes a wide lJappet, which shows considerable individual
variation in size and shape. Verhoeff bases a key on the
characteristics of the male pleopods, but although my material is
not conclusive, I do not feel disposed to allow much importance
Text-figure 2.
Ligidium hypnorum. a, antenna ; b, antennule,
to them. I suspect that they appear different in forms collected
during and out of the breeding-season. (Text-figs. 3,e; 4,a&6;
6,d,e&f; 8,ckd; 10, c&d.)
Uropods. The striking inequality of the two rami and the
characteristic inner process of the base make these appendages
systematically valuable, a fact which causes their brittleness to
be all the more regrettable. Budde-Lund divides the species
dealt with by him into two groups—the first in which the exopod
is longer than the endopod, and the second in which the reverse
is the case. Further knowledge has shown that the first group
contains but two species—the common L. hypnorum and Z'yphlo-
ligidium cecum,—and that all the other known forms fall into:
the second group. The grouping does not seem to be of primary
importance. (Text-figs. 4,¢; 6,9; 8,e; 9,¢; 10,e.)
828 MQ. Il. @. JACKSON ON THE
(3) Mey to the genus Ligidium.
The diagnoses of the genera T'yphloligidiwm and Ligidium s. 8.8.
are taken from Verhoef (1918) with one omission.
A. Eyes and body-pigment absent. Antennal flagellum 19-23 segments.
Mandibles with large interval between the biting teeth and triturating part ;
in intervening space 10-11 hairy bristles. Uropods widely separated.
Subgenus Typhloligidium Verh.
One species, L. ccecwm (Crimea—caves).
B. Eyes large, body-pigment well developed. Antennal flagelluin 10-15 segments.
Mandibles with 3-5 He setse, their biting teeth aud triturating part not
widely separated.............02.4. levees, Subgenus Ligidium Verh.
With bristle group on hind lateral eee of Ist
1 SOMITE WE, Re ah RRL Rar eM onieral a Metetee 2.
Without bristle group.. Yan nhSh Eatat eee SAO KG Heo 6.
9. ¢ With lateral decusione on inna Worder of 1st somite 3. Europe).
*¢ Without lateral depression .. betetatrteseseeseseese germanicum (Central
» § Surface covered with little ole beetesesserseseeseeee 2odulosum (Caucasus).
2b) Surface smooth and shining Bee aAEE 4,
A { Endupod of uropod longer than aorta cate hin hypnorum (Europe).
Hxopod longer than endopod wo... cece eee cee eee 3.
;_ § Mrontal grooves deep and ileal bcitttrssecseeeee bosiionse (S. Bosnia).
© U Frontal grooves absent or very shallow............ Jragile (C ‘aucasus).
6 ee rough and scaly ......cccceeeseceeeeeseeeeeeeeeee Cade (California).
* (Surface smooth and shining ...........:.cecce eee eee eee Up
Inner process of propodite of uropod shorter than
te breadthiof baseieaaraacuccscecetoesccerssr es athe japonicum (Japan).
Tnner process longer than breadth of base ......... 8.
8 { Endopod less than one-sixth longer than exopod... gracile (California).
*¢ Endopod about half as long again as exopod ...... longicaudatum (New
York).
(4) The genus Ligidium,
The diagnosis given below is modified from Sars (1899).
Body oblong ‘oval, moderately convex, attenuated behind.
Cephalon without lateral lobes. Mesosome with coxal plates
marked off from tergite by faint semicircular grooves on last four
somites. Metasome moderately small and abruptly contracted,
without produced postero-lateral angles on last somite. Eyes
large and compound. Antennula of 3 segments, small, distal
seement rudimentary, but distinctly projecting in front of head.
Antenna with multi-articulate flagellum of 10-15 (Ligidiwm) or
19-23 (Typhloligidium) segments. Mandible with Jacinia mobilis
and 3-5 (Ligidium) or 10-11 (Typhloligidium) setose bristles
between it and the molar tubercle. Ist maxilla with 3 stout
setose bristles on interior lacinia. 2nd maxilla membranous,
with 2 stout setose bristles on inner side. Maxillipede with
endopod of 5 distinct segments; inner blade linguiform and
setose. Legs very slender and greatly increasing in length
posteriorly. Opercular plates of pleopoda very thin, without any
obvious branchial structure. Uropoda moderately large, base
produced on inner side into a process upon which the endopod is
set; endopod usually longer and more slender than exopod and
provided with 2 long and slender apical bristles.
ISOPOD GENUS LIGIDIUM. 829
Text-figure 3.
fine -
¥
(c
Nu:
7 O00 SEEN eg ee ars ens
Ligidium hypnorum. a, head from front; 6, head from above; c, 1st tergite, .
lateral edge; d, Ist and 2nd tergites from side; e, Ist pleopod, g-.
1. Ligtp1um nypnorum (Cuvier). (Text-figs. 1, 2, 3, 4.)
Oniscus hyprorum Cuvier, 1792, p..19. Ligidium personat
Brandt (1833), p. 173. Ligia melanocephala Koch (1847), H. 22
(162), 18. Zia melanocephala Koch (1847), Zusammenstellung,
p. 212. Z. paludicola Koch (1847), H. 34 (180), 21. Ligidium
amethystium Schébi (1861), p.. 311. L. hypnorum Stuxberg
(1875); Budde-Lund (1885), p. 254 (q. v. for other references).
LL. cursorium Budde-Lund (1885), p. 256. L. hypnorwm
Michaelsen (1897), p. 132; Sars (1899), p. 157; Verhoeff (1901),
p. 40. Z. melanocephala Koch (1901), p. 69. £. hypnorum
Koch (1901), p. 69; Richardson (1905), p. 686; Verhoeff (1918),
p- 114; Dahl (1916 & 1917). ialiabies
Length: 6 6mm., 9 75m, Breadth: $ 25mm., 2 3° mm.
Surface smooth and shining. Head. Frontal margin slightly
830 Ml. H. G. JACKSON ON THE
sinuate ; median V very obtuse and almost linear, not produced.
Transverse groove deep and passing behind eyes. Frontal
grooves well marked, joining transverse groove at obtuse angle
behind eyes. Eyes large and pear-shaped. Vhoraa. 1st somite
with deep lateral depression on each side, extending to hind
border, On hind border of depression a dense bristle group.
Coxal plates: sutures well marked on last four somites in both
sexes; moderately drawn backwards on last three somites.
Antennal flagellum reaching back as far as hind margin of 2nd
thoracic somite; with 11 segments. Uropods. Inner process of
base long, curved, and nearly as long as base. Endopod by itself
not more than half exopod; combined with inner process reaching
nearly as far, or as far as, but not beyond, tip of exopod.
Exopod about three times as long as inner process. Zelson
arcuate ; not notched or only faintly notched over uropods.
Pleopods of 3. 1st exopod with 2 or 3 large sete; 1st endopod
drawn out internally into long narrow process with 3 large
Text-figure 4.
Ligidium hypnorum, a, 2nd pleopod, 8; b, endopod of same from another
specimen; c, uropod from above,
bristles; terminal segment of 2nd endopod with rounded or
almost triangular lappet.
Colour. Brown and yellow mottled. Dark more or less con-
tinuous band over junction of coxal plates and tergites.
Distribution. Kurope; California; Niagara (Canada), This
species has not been recorded from N. America since Stuxberg
(1875), on which Budde-Lund (1885) comments, “quid mihi
minus verisimile videtur.”
Figured in full by Sars (1899).
I have included Z. cursoriwm Budde-Land and L. melano-
cephala Koch as synonyms of this species after a minute exami-
nation of the original specimens. Verhoeff rightly conjectures
that LZ. cursorium is a large variety of L. hypnorwn, and as such
Thave distinguished it. ZL. melanocephala is a colour variety,
with slight structural features to separate it from L. hypnorwm.
J agree with Verhoeff that Z. amethystiwm is almost certainly a.
synonym.
ISOPOD GENUS LIGIDIUM. 831
Varieties.
LL. hypnorum, var. cursorium (Budde-Lund). Length 9 mm.
Breadih 4 min. Frontal margin sinuate, median V_ blunt
and slightly produced. Frontal grooves shallow. Colouring as
in L. hyprorum. Inner process of uropod base perhaps «a little
shorter than in ZL, hypnorwn.
LL. hypnorum vay. atromaculatum Verb. The dark markings
make 2 longitudinal bands on the tail in front of the telson.
L. hypnorwm var, melanocephala (Koch). Body very convex
above; antennal flagellum 9-12 segments; fourth segment of
antenna scarcely longer than the third; colour predominantly
dark brown.
2, LicibiuM rraGiLE Budde-Land. (‘Text-fig. 5.)
Ligidium fragile Budde-Land (1885), p. 257.
Jagidium ewxium Verhoeff (1918), p. 114.
Length 75 mm. Breadth 3°5 mm.
Surface smooth and shining. Head. Krontal margin sinuate ;
median V moderately sharp; not produced. Transverse groove
deep and narrow, and passing behind eyes; frontal groove very
shallow and joining transverse groove abruptly. Hyes large and
pear-shaped. Thorax. 1st somite with lateral depression on each.
Text-figure 5.
a
Ligidium fragile. a, antenna; b, head from front.
side extending to hind border. On hind border of depression a
bristle group. Coxal plates: sutures faintly marked on last
four somites; slightly drawn backwards on last three somites.
alndennad flagelluni reaching back as far as halfway across 3rd
thoracic somite; with 12-15 segments. Uropods. Inner process
of base nearly as long as base; endopods reaching further back
than exopod ; exopod about twice as Jong as inner process. (I
have not seen a complete uropod.) J'elson. Sides obtuse-angled
and with slight notch over uropods.
832 MR, If. G. JACKSON ON ‘TUE
Colow. Mottled brown and yellow; median yellow stripe from
head to tail; legs yellow.
Disobution: ‘Caucasia, Crimea.
Verhoett’s suspicion that his Z. ewainwm can be brought into
relation with Budde-Lund’s Z. fragile proves to be well founded
on further examination of Budde-Lund’s material. I have no
hesitation in placing it as a synonym of that species.
Variely.—L, fragile cauwcasium Verhoeft (1918). Epimera of
thorax predominantly coloured with irregular brown marks.
Hind angele of tergite of 2nd somite with similar depressions to
Ist terwite, Antennal flagellum with 11 segments.
LigmruM Gracie (Dana). (‘lext-fig. 6.)
Styloniscus gracilis Dana (1856), p. 176.
Ligidium tenue Budde-Lund (1885), p. 258.
Ligidium gracilis Wlolmes (1904), p. 318; Richardson (1905),
p. 690 (q.v. for other references).
Ligidiwm gracile Verhoeff (1918), p. 114.
Length: 67mm, 299mm. Lreadth: ¢ 2mm., 9 3 mn.
Surface smooth and shining. /ead. Ta marein sinuate 5
median V sharply drawn out and produced to slight rostrum.
Transverse groove deep and short, reaching to inner edge of
eyes. I[rontal grooves obsolete or very faintly indicated. Hyes
rather small and somewhat pear-shaped. Vhoraa. 1st somite
finished with bristles at regular intervals on lateral edges, none
on posterior marein, A deep lateral depression on each side
reaching hind border. No bristle group. Coxal-plete sutures
distinetly marked on last four somites in both sexes; drawn
backwards on last three somites only, the 5th somite little if at
all drawn back. Antennal flagellum reaching back to hind
margin of 2nd somites with 12 segments, Uvropods. Inner
process of base stout, slightly curved oad about half as lone as
base. Hndopod by itself slightly longer than exopod ; combined
with inner process exceeds exopod by about one-sixth of the
latter. Mxopod about four times as long as inner process,
Yelson with blunt postero-lateral angles” moderately deeply
notched over uropods.
Pleopods of 3. 1st exopod with one long bristle; 1st endopod
with process moderately drawn-out and blunt, with 3 bristles ;
2nd endopod with small rounded lappet.
Colour. Brown ground; brown median stripe; mottled on
each side with yellow. Yellow stripe along and above sutures of
coxil plates, which ave brown on each side. Legs yellow. ITead
mottled.
Distribution. California, St. Clara; Massett (British Columbia).
Sitka Island. San Francisco.
The specimens in Budde-Lund’s collection that he has identified
with Z. gracile Dana are from St. Clara (California). They
ISOPOD GENUS LIGIDIUM. 833
differ in slight particulars from Holmes’s (1904) description of
Dana’s material ; the antenne (judging by Holmes’s imperfect
figure) are shorter, but as Dana’s specimens are larger than these,
the point is not of much systematic value; the eyes of Dana’s
specimens are described as “rather large”—a verbal difference
depending on an unknown standard. These specimens have
smaller eyes than the average in Ligidiwm. There cannot be
wuch doubt that they are the same, however.
Text-figure 6.
Ligidium gracile. a, antenna; 5, head from front; ¢, Ist tergite, lateral edge ;
d, e & f, Ist and 2nd pleopods of g; g, uropod from below.
The specimens from British Columbia are labelled with an
unpublished name by Budde-Lund. They are, however, speci-
fically identical with L. gracile, but have interesting differences
that entitle them to rank as a variety. Melanin pigment is
entirely absent, and the eyes are slightly smaller. I have no.
834 MR. H. G. JACKSON ON THE
history to these specimens, but one may surmise that they are
cave varieties of LZ. gracile.
Budde-Lund (1885) described from Sitka Island a species,
L, tenue, but the oniy important points in his description are the
detail of the uropods and the size. These agree closely enough
with 4. gracile to justify a provisional inclusion of the species in
the synonymy of Z. gracile, but I have seen no specimens.
Figures in Richardson (1905).
Variety.—L. gracile var. flacwm, n. var. Pigment entirely
absent. Hyes rather small, round, and not reaching to lateral
margin of head. |
4. Licipium LArum, sp. nu. (‘Lext-figs. 7 & 8.)
Length: 6 6mm., 9 85mm. Breadth: § 3am. 2 41min.
Surface rough and covered with small scales, ZZead. Mrontal
margin sinuate; median V very sharp and produced. ‘Transverse
groove deep and passing behind eyes; frontal grooves very deep
and curving back to join transverse groove abruptly. Myes large
and pear-shaped. Zhoraz. 1st somite without lateral depressions
or bristle groups. Sete at intervals on lateral border, but absent
on posterior border. Coxal-plate sutures well marked on last
four somites in the female, only lightly marked on last two in
male; weil drawn backwards on last three somites and slightly
on fourth somite. Antennal flagellum long, reaching back as
+
a
Text-figure 7,
RS eae
;
Cee
~ ——— as
ea
a 6
~~
Ligidium latum. a, last four thoracic somites from left side; 2, head from above.
far as hind margin of 4th somite; with 12 segments. Uvropods.
Tnner process of base shorter than base by about half its length,
stout and conical. (None of my specimens have undamaged
uropods.) ‘Lhe single seta arising from outer side of base, set
on sharp and pointed process. J'elson deeply notched over
uropods ; blunt, rounded postero-lateral angles.
Pleopods of 3. 1st exopod with one (?) bristle ; 1st endopod
with moderate process and one(?) bristle; 2nd endopod ending
in pointed process.
Colour. Brown and yellow mottled; slightly lighter streak
down middle of back. Coxal plates light and sharply defined
ISOPOD GENUS LIGIDIUM. 835
from brown of tergite at the suture. Legs banded with yellow
and brown.
Distribution. San Francisco.
Text-figure 8.
ope
Ligidium latum. a, head from front; 5, Ist tergite, lateral edge ; ¢ & d, endojiod ‘
of Ist and 2nd pleopods, %; e, uropod from below.
Cc
t
m
his species is distinguished from all but LZ. nodulosum by its
very distinctive rough surface. It differs from that species in
the character of the granulations and the form of the lst somite.
5. Ligiprum cermanicum Verhoeff. (Text-fig. 9.)
Ligidium germanicum Verhoeft (1901), p. 41; Verhoef (1918),
p- 114.
Ligidiuwm herzegowinense Verhoeff (1918), p. 114; Dahl (1916
& 1917).
Length 7mm. Breadth 2-5 mm.
Surface smooth and shining. Head. Frontal margin slightly
sinuate, median V almost obsolete, not produced. ‘Transverse
groove deep and passing behind eyes; frontal grooves shallow
and curved, passing into transverse groove at less than a right
angle. Eyes large and pear-shaped. Thorax. Ist somite without
lateral depressions or bristle groups. Coxal plates: sutures
836 MR. . G. JACKSON ON THE
faintly visible on last three somites of male, very faint or obsolete
on female; hind corners of last three but little drawn out.
Antennal flagellum veaching back as far as halfway across
3rd somite ; stout and somewhat setose; with 9-10 segments.
Uropods. Inner process of base curved and longer than base.
Endopod by itself about as long as exopod ; combined with inner
proeess exceeding exopod by «about one-third of the latter.
Exopod about twice as long as inner process; setie of endopod
Text-figure 9. '
SS
mo
SS
aa X\
sf : 6 4
(os
a WA
Pa
SS
Ligidium germanicum. a, head from front ; b, antenma; ¢, vropod from below.
about three-quarters as long as that branch. Zelson arcuate,
inclining to a bluntly angulate condition iu some; no notch
above uropods.
Pleopods of 3. Similar to L. hypnorum.
Colour. Predominantly brown, mottled with yellow; legs
yellow.
Distribution, Central Kurope.
6. Liciprum saronicum Verhoeff. (Text-fig. 10.)
Ligidium japonicum Verhoeft (1918), p. 114.
Length 6mm. Breadth 2°5 mm.
Surface smooth and shining. J/ead. Frontal margin sinuate,
forming moderately sharp V; not produced, ‘Transverse groove
‘narrow and deep, and passing behind eyes. Frontal grooves
shallow, joining transverse grooves abruptly. NMyes large and
pear-shaped. Zhoraxz. Ist somite with deep lateral depressions ;
no bristle groups, but a group of little processes lies in each
depression. Coxal plates: sutures not visible; hind corners of
last three slightly drawn out. Antennal flagellum veaching as
far back as hind margin of 3rd thoracic somite; with 14 segments ;
long and setose. Uvropods. Inner process of base short and
blunt, hardly one-third base and shorter than width of base.
Endopod by itself about half as long again as exopod, combined
with inner process nearly two-thirds as long again. Exopod
about six times as long as inner process. T'elson. Only slightly
notched above uropod; no postero-lateral production,
ISOPOD GENUS LIGIDIUM. 837
Pleopods of 3. 1st exopod with 5-6 large sete; Ist endopod
drawn out internally into sharp triangular process with 5-6 large
sete ; 2nd exopod ‘divided very distinctly by a suture in two
parts” (Verhoeff) ; 2nd endopod with small rounded projecting
terminal lappet and three small spines on inner edge at distal
end.
Text-figure 10.
Ligidium japonicnm, a, head from front; b, antenna; ¢ & d, endopod of
Ist and 2nd pleopods, g; e, uropod from below.
Colowr. Light brown with whitish speckling; longitudinal
band of large spots over coxal plates; median brown band without
mottling ; legs whitish.
Distribution. Japan ; “ Moheri” (Japan).
Figured by Verhoeff (1916).
I have not seen specimens of the following species :—
7. LicrpiuM BosnieNsE Verhoeff.
Ligidium bosniense V erhoeff (1901), p. 40; (1918), p. 114.
Distribution. Sarajevo, Bosnia.
Briefly described but not figured.
8. Licrp1um NopuLosum Verhoeff.
Ligidium nodulosum Verhoef (1918), p. 114,
Distribution. Caucasus.
Described with figures by Verhoeff.
Proc. Zoon. Soc.—1923, No. LY. 5D
838 MR. W. G. JACKSON ON THE
9, LigrIpIUM LONGICAUDATUM Stoller.
Ligidium longicaudatum Stoller (1902), p. 208; Richardson
(1905), p. 688; Verhoeff (1918), p. 114.
Distri nos New York,
Verhoeff remarks of this species that the figure given by
Stoller can scarcely be accepted. The descriptions ave also in-
adequate, and the specimens should be redeseribed in the light of
recent work.
10. Licrprum (TypHLoLierpium) caz:cum (Carl).
Ligidium cacum Carl (1905), p. 327.
Lyphlotigidium cecum Verhoeff (1918), p. 114.
Distribution. Crimea, in caves.
Verhoeff supplements Carl’s description, but no new figures
have been given. Carl figures the telson and uropods and the
second pleopod of the inale.
I wish to acknowledge my debt to Dr. W. T. Calman, F.R.S.,
for allowing me the use of the British Museum inaterial, and
for his readiness in advising on questions of Crustacean
morphology and literature,
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Hamburgs.” Mitt. Naturhist. Mus. Hamburg, xiv.
pp. 121-134.
ISOPOD GENUS LIGIDIUM. 839
Ricnarpson, Harrger.—l905. “ A| Monograph on the Tsopods
of North America.” Bull. U.S. Nat. Mus. no. 54.
Sars, G. O.—1899. Crustacea of Norway. 1. — Isopoda.
Bergen.
SCHOBL, J. 1861. Korysi stejnonozi (Crustacea Isopoda) .
iva. Casopis Pitrodnicky. Prague, pp. 307-821.
SrouLer, J. LP.—1902. “Two new Tanne Tsopods.” Rept. N.Y.
State Museum, 54, vol. i. (for 1900), pp. 1238-1213.
2 text-figs.
SruxperG, A.—1875. “Om Nord-Amerikas Oniscider.” Ofver-
sigt K. Vetenskaps-Akad. Férhandl. xxxii. 2, pp. 43-63.
Verworrr, K. W.—1901. “Uber paliiarktische Isopoden.” Zool.
Ane. xxiv. pp. 38-41. Leipzig.
Veruorrr, K. W.—1918. “Zur Kentniss der Ligidium ete.”
Archiv f. Naturges. 82. Jahrg. (for 1916), Abt. A, 10 Heft,
pp. 108-169, 2 pls., 3 text-figs.
Waurverc, R.—-1922. “Terrestre Isopolen aus Australien.”
-Arvkiv f. Zool. xv. 1, Stockholm. ‘Text-figs.
co
Ct
*
LIZARDS OF BRITISH 'PERRITORIES IN EAST AFRICA. 841
41. A List of the Lizards of British Territories in Bast
Africa (Uganda, Kenya Colony, Tanganyika Territory,
and Zanzibar), with Keys for the diagnosis of the
Species. By Arrnur Loveriper, F.E.S., C.M.Z.S.
| Received October 3, 1923: Read October 23, 1923. ]
As no work on the Lizards of the British territories in East
Africa, as a whole or in part, has ever appeared in the English
language, the writer hopes that the accompanying keys and check-
list may prove of use to his fellow herpetologists and perhaps
help to stimulate an interest in the Lizards of this region.
Both Tornier* and Niedent have published works dealing
with the reptile fauna of German Hast Africa (now ‘Tanganyika
Territory) and have included in these many species from British
Kast Africa (now Kenya Colony) and Uganda likely to be found
in G.H. A. Owing to the many species ‘descr ibed by Boulenger,
Sternfeld and other rs, however, both these works may be con-
sidered out of date.
Like all herpetologists, I must confess my indebtedness to tie
writings of Mr. G. A. Boulenger. In the case of the Lizards
proper (Lacertidz) I have adopted, or extracted, from his keys +
to suit my purpose.
Out of the 112 species and races considered valid in this paper,
no fewer than 64 have heen described since the publication of his
‘Catalogue of Lizards’§. A still more significant fact as to the
amount of work done in recent years is that, of the 48 species which
appear in the Catalogue, only 15 are recorded from, or credited to,
the area dealt with in the following pages, which have perforce been
compiled from a mass of papers that have appeared since 1887.
Wn . 1d Chaprae 10e
Number of Species characterised. Preseutiyiper,
By Bler. 1885-87. ‘Torn. 1897. Nied. 1913. 1923.
Geckonide............ 1 11 16 25
INGENIOUS Goo sosseoses I 8 6 13
Zonunidceweee 0 2 4. 4
Varanidee ............ 0 3 3 3
Amphisbeenide ...... ] 0 4 5
Lhacertidie ............ 3 6 13 20
Gerrhosauride ...... I 3 2 4
DCUMC) es eee 8 17 24. oi
Anelytropidee ...... 0 1 1 1
“NOtEAIS. so snoxconos 15 51 73 112
* "Tor nier, ‘ Die Kriechtiere Denfech- Ost- Afr ikas, i 1897, and Zool. Jahrb. Syst. xiii.
Band 6, 1900.
+ Nieden, “Neues Verzeichnis der Kricchtiere . . . von Deutsch-Ost-Afrika,’ 1913.
ft Boulenger, ‘ Monograph of the Lacertida,’ i. & ii. 1920-21.
§ Boulenger, ‘Catalogue of the Lizards in the British Museum,’ i.-iii. 1885-1887.
842 Mh. A. LOVERIDGE: LIST OF TILE LIZARDS
As brevity is so essential in a key I have omitted lists of
bibliographical veferences, giving in the case of those 50 species
appearing in the ‘Catalogue of Lizards’ the volume and page
reference of that work only, in the case of those subsequently
described only the reference to the publication in which its
description first appeared.
‘The synonyms, which are also numerous, have been omitted for
the same reason, and only a few given of ‘recent date which may
not be generally ‘know n.
With the exception of the generic keys for the Geckonidw and
Lacertidee, t have been aisle: to arrange my keys according to
characters of taxonomic value, so that the genera or species
fall into their natural order. In the two generic keys men-
tioned this was not possible without making thera unduly
cumbersome.
Synopsis of the Families.
1, Upper surface of head with scales or granular tubercles.
A. Tongue short with villose papille.
No movable eyelids; dentition plenrodont ............ GECKONID™
Movable eyelids; dentition acrodont..................... AGAMID&.
B. Tongue long, smooth, deeply bifid, sheathed at base... VARANID.%.
IJ. Upper surface of head with large or symmetrical shields.
A. poueue with villose papillee ; femoral pores ............ ZONURID .
B. Tongue with scale-like imbricate papilla or oblique plicie.
Body vermiforn ; scales soft, squarish, not imbricate. AMPiise.uNnipa:.
No bony plates under the ceales feinoral pores
usually present ............5. eee ANGI aay
Bony plates under ‘li enleae it iidieen fold femoral
pores present. . adheres j veceeeeeeee GERRIOSAURLD A.
Bony plates nition! Alo etloals ; no eel fol; no
femoral pores ; eyes distinct ...... ogpttnee san abo an cada ScIncipa,
Bony plates under the scales ; body vermiform; eyes
concealed beneath skin... 0... ccc eee ANELYTROPID.U.
Family GeEcy ONID®.
Ten genera.
I. Digits not, or but slightly dilated.
A. Punil vertical
IN@ WOES coates sed nuuodedeb dou ossvanena .eodnneenG0bOGH As Boonosdon bee SLADMEMICLIN Us.
Rrganalipores eeveeec. eeicceesteee eee eee ene eBumocnenis:
io TEND ROMINA coo.c6u noc eus'bbo dogo noe den obaRcSonaorodeusneoouncaren ona. . Cranmutiatas.
If. Digits dilated.
A. Pupil vertical.
1. Digits with a strong claw.
a, Sub-digital plates divided.
Digits dilated at apex only, not at base .................. Diplodacty/us.
Distal phalanges compressed............... 00. 0ceceesee cesses Hemidactylus.
b, Sub-digital plates undivided,
Thumb and inner toe clawless ............0..0. ccc ~Platypholis.
OF BRITISH TERRITORIES IN BAST AFRICA. 343
2. Digits clawless or with a minute claw.
Digits moderately dilated ; nostril pierced between the
rostral and three small scales... .........c00ccce0eeee 0s Pachydactylus.
Digits stronely dilated; nostril pierced between the
rostral, Ist labial, and three small scales ............ Elasmo:lactylus.
B. Pupil cireular.
Sub-digital lamellie in two rows 0.0.0... cccccececcee cece Lygodactylus.
Sub-digital lamella undivided .. ..................0..0..... Phelsuma.
1. STENODACTYLUS. —
Fitzing. N. Classif. Rept. p. 13; Blgr. Cat. Liz. i. p. 16.
1. SrenopacryLus eurratus Cuy,
Blgr. Cat. Liz. i. p. 17, 1885.
Kenya Colony.
2. GONALODES.
Fitzing. Syst. Rept. p. 91; Blgr. Cat. Liz. i. p. 56.
Synopsis of the Species,
5 or 6 upper, and 6 or 6 lower labials ; 7 to 8 preanal
POLES Mh. CAE As cade eer ne cat cosade ack, evn eee
6 or 7 upper, and 7 or 8 lower labials; 8 to 12 pre-
anal pores ee Se le,
quattuorseriatus.
africanus.
1. GoNATODES QUATTUORSERIATUS Stern.
Stern. Ergebn. Deut.-Zentral-Afrika Exp. 1907-08, iv.
(Z. 2), 'p. 202, 1913.
Kenya Colony.
2. GONATODES AFRICANUS Wern.
Gymnodactylus africanus Wern. Verhlg. k. k. zool.-bot. Ges.
Wien, 1895, p. 190.
Kenya Colony; Tanganyika Territory. .
3. DrrLopactryLus.
Gray, Proc. Zool. Soc. 1852, p. 40; Blgr. Cat. Liz. i. p. 97.
1. DIPLODACTYLUS WOLTERSTORFFI Torn.
Torn. Zool. Jahrb. Syst. xiii. p. 584, f. A, 1900.
Tanganyika Territory.
4, Hemipacrytus. 4
Hemadactyles Cuv. R. A, ii. p. 47; Blgr. Cat. Liz. i. p. 113.
844
MR. A. LOVERIDGE?: LIST OF TILE LIZARDS
Synopsis of the Species.
T. Dorsal tubercles, if present, small, smooth, conical, or very feebly keeled,
No tubercles; male with six praeanal pores.. skedetonsudsoudes § COOUG BIS.
Small tubercles on occiput and neck only; ele “ah seven
pruanal pores ........ tropidolepis.
Fecbly keeled scales on heey sarvondell bya a ring Nice Healers
scales ; male with 16 prwanal pores ........ vecseeteceerseee SqQuamulatus.
Sinall tubercles ; male with 15-80 femoral pores ............... mabouia.
If. Dorsal tubercles present and strongly keeled.
bo
oO.
‘4 or 6 lamellae under the inner, and 6 or 7 under the median
toe; 8 to 10 upper and 5 or 7 lower labials .........c00c0s000. ei€ernii.
4 or 5 lamellae under the inner, and 6 to 8 under the tee
toe; 8 or 9 upper and 7 or 8 lower labials, Head covered
with plates more or Jess equal in size and generally keeled. vwspolié.
4 to 6 lamella under the inner, and 7 or 8 under the median
toe; 7 to 10 upper and 6 to 9 lower labials. Head covered
with granules and scattered oval, keeled shields much
larger ‘than the granules ............. brookii.
5 or 6 lamellw under the inner, and 8 tol 10 Hider the sredian
toe; 9 to 10 upper and 8 or 9 lower labials ............... wereri.
7 or 8 Jamellie under the inner, and 10 to 12 under the median
toe; 9 upper and 8 lower labials 2.0.00... eee e eee eee se maeropholis.
HeEMIpACTYLUS IsOLEPIS Bley.
Blgr. Proc. Zool. Soc. 1895, p. 531, pl. xxix. fig. 1.
Kenya Colony.
HEMIDACTYLUS TROPIDOLEFIS Mocq.
Mocq. Mém. Cent. Soc. Philom. Paris, 1888, p. 113.
Kenya Colony; Tanganyika Territory.
HEMIDACryLUS SQUAMULATUS Torn.
Torn. Die Kriecht. Deut.-Ost-Afr. p. 10, 1897.
Doubtfully distinc’ from //. tropidolepis Mocq.
Kenya Colony; Tanganyika ‘Territory.
Hemipactyius MABouta Mor.
Blgr. Cat. Liz. i. p. 122, 1885.
Kenya Colony; ‘Tanganyika ‘Territory; Zanzibar.
Hemipacryius crrerni Bier.
Blgv. Ann. Mus. Civ. st. nat. Genova, ser. 3a, vol. v. (xlv.),
IM
Kenya Colony; Tanganyika Territory.
. Hemipacryius ruspout Bler.
Blgr. Ann, Mus. Civ. st. nat. Genova, ser. 2, xvii. p. 6
1896.
Kenya Colony ; Tanganyika Territory.
OF BRITISH TERRITORIES IN EAST AFRICA. 845
7. WLemipactytus Brookt Gray.
Blev. Cat. Liz. 1. p. 128, 1885.
Uganda; Kenya Colony; Tanganyika Territory; Zanzibar.
8. Hemrmacrynus wernert Torn.
Torn. Arch. Naturg. vi. p. 63, 1897.
Hemidactylus tornieri Mocq. Bull. Mus. Paris, 1902, p. 404.
Kenya Colony; Tanganyika Territory.
9, HemmacryLus MACROPHOLIS Bley.
Blgr. Ann. Mus. Civ. st. nat. Genova, ser. 2, xvii. p. 7, 1896.
Kenya Colony.
5. BUNOCNEMIS.
Giinther, Proc. Zool. Soc. 1894, p. 85.
1. Buxocnemis Moprsrus Gthr.
Gthr. Proc. Zool. Soc. 1894, p. 85.
Kenys Colony.
§. LYGODACTYLUS.
Guay, Proc. Zool, Soc. 1864, p. 59; Blgr. Cat. Liz. i. p. 158.
Synopsis of the Species.
I. Subcaudal scales small... .......00cccccc cee ceeeee cee ceseeeceeeesseesse eee CUPENSiS.
II. Subeaudals enlarged, in two longitudinal rows.
9 upper and 6 lower labials ; 10 prwanal pores in male....., fischeri.
5 or 6 upper and 5 or 6 lower Jabials; 6 preanal pores in [fleri.
THD Oie Rims He et Sette SuRANA Eigte th othe saat daa. ) uafoa MELT sn SCILEJi=
IIL. Subcaudals transversely enlarged in a single median series. (DL. grotet may
combine subcandal arrangement of I, IL, & III.)
6 to 10 upper and 5 to 8 lower labials ; 4 to 6 preanal pores
in male. Throat pure white ........ ......0.6ce ee
5 or 6 upper and 6 or 7 Jower labials; 8 to 9 preanal pores
inmale. Throat usually black in imale.....................6+ picturatus.
5 to 7 upper and 5 to 7 lower labials; 8 to 9 praeanal pores
in male. ‘Throat usually with chevron-shaped markings. p. gutturalis.
grotet.
1. LyGopAcryLus CAPENSIS Sunith.
Blgr. Cat. Liz. i. p. 160.
if a] g G Keowee f
Tanganyika ‘Territory.
2. LyGopacryLus FriscHERI Blgr.
Blgr. Proc. Zool. Soc. 1890, p. 80.
Kenya Colony ; Tanganyika Territory.
3. LYGODACTYLUS FISCHERI, var, SCHEFFLERI Sternf.
Sternf. Ergebn. Deut.-Zentral-Afrika Exp. 1907-08, vol. iv.
(Z. 2) p. 206, 1913.
Kenya Colony; ‘Tanganyika Territory.
846 Mk. A, LOVERIDGEH: LIST OF TILE LIZARDS
4, LyGopAcryLus Grover Sternf.
Sternf. 8.B. Ges. naturf. Berlin, nr. 4, p. 245, 1911.
Tanganyika Territory.
. Lygopacryius picrurarus Peters.
Blgv. Cat. Liz. i. p. 161.
Kenya Colony; Tanganyika Territory; Zanzibar.
6. LyGoDACryLUS PICLURATUS GULTURALIS Bocage,
Blige. Cat. Liz. i. p. 161.
Uganda.
7. PLATYPHOLIS.
Blgv. Proc. Zool. Soe. 1890, p. 80.
1. Pravyrnouis rasciara Blgr.
Bler. Proc. Zool. Soe. 1890, p. 81, pl. vill. fig. 2.
Kenya Colony; Tanganyika Territory.
8. PACHYDACTYLUS.
Pachydactyles, part., Cuv. Régne Anim. ii. p. 45; Blgr. Cat.
Liz. i. p. 200.
Synopsis of the Species.
Dorsal tubercles strongly keeled. Tfead covered with convex,
Keeledbseal es) capscaeas arsine cease atea te ce erateremaceetaoe saci tactert ... bibronii.
Dorsal tubercles feebly keeled. Ilead covered with smooth
cone-shaped [email protected] tee tee tsteeeeseeeeenerseees boulengeri.
1. PacHyDACYYLUS BIBRONII Smith.
Blev. Cat. Liz. i. p. 201.
Tanganyika Territory.
2, PACHYDACTYLUS BOULENGERI Torn.
Torn. Kriechtiere D. O. A. p. 26, 1897.
Tanganyika Territory.
9, PHELSUMA.
Gray, Ann. Phil. (2) x. p. 199, 1825; Blgr. Cat. Liz. i. p. 209.
1. PaeLsuma Laticaupa Bttgr.
Blgr. Cat. Liz. i. p. 215.
P. dubius (Bttgr.), Blgr, Cat. Liz, i. p. 215.
Tanganyika Territory ; Zanzibar.
OF BRITISH TERRITORIES IN EAST AFRICA.
10. ELASMODACTYLUS.
Bler. Proc. Zool. Soc. 1894, p. 727.
1. ELAsmopacryius trigpRus Blgr.
Blgr. Rev. Zool. Afr. iii. p. 104, fig., 1913.
Tanganyika Territory.
Family AGAMID &.
‘wo genera
Hl, JieReeyes SARAAML, GAMA cosccecno css coe soe osdlabe tien oo pee coogooouE cba wenecDeDd
Il. Incisors united into one or two large cutting teeth; tail short,
TTA OW NOTA OLE SPDINES 556 c09 occ osdnae dad Soc anos 00 co noH ceaqno nos peeoDenoD oer
Ile EES:
847
Agama.
Aporoscelis.
Agama, part., Daud. Hist. Rept. iii. p. 8383; Blgr. Cat. Liz.
i. p. 334.
Synopsis of the Species.
I. Occipital scale enlarged.
A. Dorsal scales unequal.
Ventral scales keeled.. eA i
Ventral scales not or but sieallke healed
3. Dorsal scales equal.
1. A slight denticulation or crest on the vertebral line.
Smooth or keeled ventrals much smaller than the dorsals.
Smooth ventrals scarcely smaller than the dorsals .........
2. No dorsal deuticulation or crest.
a. A small nuchal crest.
Longest spines about car and on sides of neck about half
the diameter of the ear-opening ; dorsal scales strongly
keeled . Acie NMA dod aatnici uae dampens aurea aren
Longest spines about ear aun on sides of neck one-third the
diameter of the ear- -opening; dorsal scales keeled, not
or but very slichtly mucronate de Waa ek ERR Re
Longest spilies about ear and on sides of neck upiisidlere
ably shorter than the diameter of the ear-opening ;
dorsal scales strongly keeled, mucronate ............
Longest spines about ear and on sides of neck hardly
more than half the diameter of the ear-opening. 80 or
90 scales round mid-body. Head longer than broad.
Dorsal scales ee smooth or ets: feel Keele not
mucronate .............
Longest spines anand ¢ ear iia on ie ties "Of. acals ‘nearly
equal the diameter of the ear-opening (in type speci-
men); dorsal scales small, very feebly and obbusely,
keeled; throat scarlet in g & ng
Longest spines: about ear and on sides of neck net ate
equal to, but about one-fifth the diameter of the car-
opening; dorsal. scales small, obtusely but strongly
keeled :
(i.) Throat scarlet with bright blue border in gS.
hispida.
h. distanti.
mossambica.
kirkii.
doria.
planiceps.
colonorum.
elgonis.
lionotus.
lionotus, var. dodome.
(ii.) Snout to mid-body including throat bright
metallic pink in OG ......ce eee. Udonotus, var. mwanze.
848 MR. A. LOVERIDGE: LIST OF THE LIZARDS
Longest spines about ear and on sides of neck three-
quarters the diameter of the ear-opening ; dorsal scales
small, very feebly and obtusely keeled, ventrals
smooth. aint nuchal crest ; caudal scales very large
FONG! SUROWENY SPOWOSO — cooaobccocccens oon onancoooe ves oodane ona nod flavicauda.
6. No nuchal crest. Longest of the long spines about
ear and on sides of neck equal the diameter of the
car-opening; dorsal scales very large, mucronate,
imbricate with strong keels ..........cccccceceeeeeeeeeeeeeee Uaillanti.
11. Occipital scale not enlarged ; caudal scales forming more or less distinct annuli;
ventral scales smooth; digits compressed.
Shight nuchal crest; caudal scales large, strongly keeled, aéricollis.
1, AGAMA wisprpa Linn.
Bler. Cat. Liz. i, p. 349.
Tanganyika Territory,
2, AGAMA HISPIDA, Var. DISTANTI Bley.
Bler. Ann. & Mag. N. H. (7) ix. p. 339, 1902.
Ugamla; Tanganyika Territory.
3. AGAMA MOSSAMBICA Peters,
Bler. Cat. Liz. 1. p. 353.
Tanganyika Territory.
(3a AGAMA KInkIT Bler.
Blgv. Cat. Liz. i. p. 384.
Possibly oceurs in Tanganyika Territory.
y Sy, )
4, AGAMA DOoRLE Bler,
Blgr. Ann. Mus, Civ. st. nat. Genova, ser. 2, 11. p. 127, 1885.
o ? ?
Kenya Colony; Tanganyika erritory.
5. AGAMA PLANICEPS Peters.
Blgr. Cat. Liz. i. p. 358.
Tanganyika Territory.
6. AGAMA COLONORUM Daud.
Blev. Cat. Liz, i. p. 356.
Uganda; Kenya Colony; Tanganyika Territory.
7. AGAMA ELGONIS Linnb.
Lonnb. Arkiv for Zoologi, Band 14, no. 12, 1922.
Kenya Colony.
8. AGAMA Lionorus Blev.
Blgr. Proce. Zool. Soc. 1896, p. 214, pl. viii.
Kenya Colony; ‘Tanganyika Territory.
OF BRITISH TERRILORIES IN BAST AFRICA. 849
9. AGAMA LIoNorus, var. Dopoma# Loveridge.
Love. Proc. Zool. Soc. 1923, p. 944.
Tanganyika Territory.
10. AGAMA Lionotus, var. MWANZa: Loveridge.
Love. Proc. Zool. Soc. 1923, p. 945.
Tanganyika Territory.
11. AGAma vaiLiaAnti Blgr.
Blgr. Ann. Mus. Civ. st. nat. Genova, ser. 2, xv. p. 12, 1895.
Kenya Colony; Tanganyika Territory.
12, AGAMA FLAvIcAUDA Wern.
Wern. Zool. Anz. xx. p. 264, 1897.
A. cuudospina Meek, Wield Mus. Nat. Hist. Pub. 147, p. 407,
1910. :
Kenya Colony.
15. AGAMA ATRICOLLIS Sinith.,
Blgr. Cat. Liz. i. p. 358.
Uganda; Kenya Colony; Tanganyika Territory.
2. APOROSCELIS.
Blgr. Cat. Liz. 1. p. 410,
1. AroroscELis Princers O’Shaughn.
Blgr. Cat. Liz. i, p, 410.
Zanzibar.
Family ZonuRID &.
Two genera,
I. Limbs well developed ; caudal whorls very spinous..... ......... Zonuras.
I]. Limbs rudimentary ; body serpentiform ..........0.0.....c00.... Chamesaura.
1. Zonunvs.
Merr, Tent. Syst. Amph. p. 57; Blgr. Cat. Liz. ii. p- 252.
1. Zonunus TRopiwosrerNuM Cope.
Blgrv. Cat. Liz. ii. p. 254,
Tanganyika Territory.
2. CHAMASAURA.
Chamesaura, part, Schneid. Hist. Amph. p. 204; Blgr. Cat.
Liz. it. p. 263.
850 Mk. A. LOVERIDGE: LIST OF THE LIZARDS
Synopsis of the Species.
I. Both pairs of limbs very distinct ; 24 scales round the body.
Hind limb didactyle ..........0:cc0 cee eee cesses neces ees seessseeeesteeseeeees @Mmectans.
Hind limb monodactyle.............cceeseeceeeeeceetee eee eeseeresteeenteees CEnUION.
II. Fore limbs reduced to minute clawed vestiges ; 26 scales round the
[NetN | poyene sedaucrcibdencacnsstie ork sSubs6nt0 n5. desea ccan0 "AG nosesa na jooo cas esa\ a CA DU
1. CuAM#sAURA ANNECYANS Bler.
Bler. Proc. Zool. Soc. 1899, p. 97.
Uganda; Kenya Colony.
2, CHAMESAURA TENUIOR Gthr.
Gthr, Ann. & Mag. N. H. ser. 6, xv. p. 524, 1895.
Uganda ; ‘Tanganyika Territory.
3. CuaMasAURA MIoPRoPUs Bler.
Blgr. Proc, Zool. Soc. 1894, p. 732.
Tanganyika ‘Territory.
Family VARANIDA,
One genus.
1, VARANUS.
Merrem, Tent. Syst. Amph. p. 58; Blgr. Cat. Liz. ii. p. 304.
Synopsis of the Species.
I. Nostril an oblique slit, three times more distant from the end of the snout
than from the orbit.
Scales small; nuchals not larger than the occipitals ......... albigularis.
Scales large; nuchals larger than the occipitals ............... ocellatus.
II. Nostril round, a little nearer the orbit than the end of the snout . xtloticus.
1, VARANUS ALBIGULARIS Daud.
Blgr. Cat. Liz. ii. p. 307.
Kenya Colony ; Tanganyika Territory; Zanzibar.
2. VARANUS OCELLATUS Riipp.
Blgr. Cat. Liz, 11. p. 308.
Kenya Colony; Tanganyika Territory.
3. Vanranus NiLoticus Linn.
Blgr, Cat. Liz. ii. p. 317.
Uganda; Kenya Colony ; ‘Tanganyika Territory ; Zanzibar,
OF BRITISIL TERRITORIES IN EAST AFRICA. 851
Family AMPHISBHZNID&.
Four genera.
I. Segments of the pectoral region not differentiated.
4 Jabials, third largest
2 Jabials, first largest ..
4 shields bordering mouth oc (for med ip fusion of Tabial
and nasal, and prefrontal and ocular) largest ............ Chirindia.
Amphisbena.
Amphisbenula.
II. Segments of the pectoral region slightly enlarged, forming an angular scries.
3 labials, first very small ............... ce cee eeeeee cee eeseesrneees Gteocalamus.
1. AMPHISBENA.
Linn. Syst. Nat. i. p. 392, et omn. auct.; Blgr. Cat. Liz
p. 435.
5 ile
1, AMPHISBENA PHYLOFINIENS Torn.
Torn. Zool. Anz. 1899, p. 260.
ry ee rr ae a
Tanganyika Territory.
2. AMPHISBENULA.
2 Sternf. S.B. Ges. naturf. Berlin, 1911, p. 246. °
1. AMPHISBENULA ORIENTALIS Sternf.
Sternf. S.B. Ges. naturf. Berlin,.1911, p. 246.
Tanganyika Territory.
3. CHIRINDIA.
Blgr. Ann. Mag. N. HH. (7) xx. p. 48, 1907.
1. CHIRINDIA EWERBECKI Wern.
Wern,. Hamb. Jahrb. wiss. Anst. xxvii. p. 37, 1910.
4, GEOCALAMUS.,
Giinth. Ann. & Mag. N. H. (5) vi. p. 234, 1880; Blgr. Cat.
Liz. 11, p. 453.
Synopsis of the Species.
2 lower labials ; 2 temporals: 38 to 42 segments round body
o99.000 acutus.
3 lower labials; 3 temporals; 34 to 38 segments round body
Listas modestus.
1. GEOGALAMUS AcUYUS Sternf.
Sternf. pegs Deut.-Zentral-Afrika Exp. 1907-08, iv.(Z. 2),
p. 209, 1912
Kenya Goi ; Tanganyika Territory.
i)
. GEOCALAMUS MopbEStUS Gthr.
Blgv. Cat. Liz. i. p. 403..
Tanganyika Territory.
852 MR, A. LOVERIDGE: LIST OF THE LIZARDS
Family LACERTID&.
I. Ventral plates smooth.
A. Sub-digital lamellaa smooth or tubercular.
1. Dorsal scales small, juxtaposed or subimbricate.
Nostril pierced between 2 or 3 nasals, separated from
first upper labial; cheeks swollen
Nostril pierced between 3 or 4 nasals, well separated
from the first upper labial ; cheeks not swollen ......
Nostril vata between 2 or 3 nasal and the first upper
labial... roe i ea
2. Dorsal santas ih ze, ie fete “tnouiaey poorer
3. Two series of large smooth plate-like scales allows fie back
and tail Birt ees
B. Subdigital lamellee estas
Nostril pierced between 3 to 5 nasals and the upper
labial, or separated from the latter by a narrow rim .
Nostril pierced between 2 or 3 nasals ..........6.00. ccc ee
IT., Ventral iplates keeled’ ooo Miccccssocansceececcss sobedeceetcvns sdedun coeses
1. Nucras.
Nucras.
Hremias.
Lacerta.
Algiroides.
Holaspis.
Tatastia.
Ichnotropis.
Gastropholis.
Gray, Ann, & Mag. N. H. 1.1838, p. 280; Blgr. Cat. Liz. iii.
p. 52.
Synopsis of the Species.
I. Dorsal scales keeled 10... ..se.0eseeee0e
IT. Dorsal scales smooth. ;
kilose.
Head 4 to 5 times in length to vent; foot as long as or a little
longer than the head; parietal foramen present; 40 to 51
scales across the middle of body; ventrals in 28 to 33 trans-
verse series ; transversely enlarged plates under the fore-arin ...
Head 43 to 6 times in length to vent; foot shorter than the
head; 45 to 63 scales across middle of body; ventrals in 27 to
SAtVANSVELSC!SOLLES Secyocal Wecsattes Mee eran a sigseeoeinct eon ee on aac eee eee
1. Nucnras kinosm Loveridge.
Love. Proc, Zool. Soc. 1922, p. 314.
Tanganyika Territory.
2. Nucras EMINI Bler.
Bier. Ann. & Mag, N. H. (7) xix. p. 593, 1907.
Kenya Colony; Tanganyika Territory.
3. Nucras BOULENGERI O. Neum,
Neum. Ann. & Mag. N. H. (7) v. p. 56, 1900
Kenya Colony ; ‘'anganyika Territory.
2. Lacerra.
Lacerta, part., Linn. 8. N. i. p. 359.
Lacerta Bigy. Cat, liz. iii. p, 8.
emini.
boulengeri.
OF BRITISH TERRITORIES IN EAST AFRICA. 853
Synopsis of the Species.
I. Transverse series of ventral plates with notches between the
plates ; collar serrated ; a single postnasal (exceptionally 2) ;
femoralsporesi@tonl Umea enn eee ene ee RON a arena Tis
If. Transverse series of ventral plates with rectilinear or nearl
ae see : : y
rectilinear border, longitudinal series 6 or 8 in number;
femoral pores 12 to 31; 22 to 26 lamella under fourth toe . Jjacksoni.
Parietals usually not in contact (or very narrowly) with:
upper postocular; temple covered with small scales, as
large as or smaller than dorsals ..2............600ccc000 wees J. Jacksoni.
Parietals more extensively in contact with upper post-
ocular; temple scales larger than dorsals.................. Jj. kibonotensis.
1. Lacerra VAUERESELLI Torn.
Torn. Zool. Anz. xxv. p. 701, 1902.
Tanganyika Territory.
2, Lacerta JAcKsoNI Bler.
Blgr. Proc. Zool. Soc. 1899, p. 96, pl. x.
Uganda; Kenya Colony.
2A. LACERTA JACKSONI, var. KIBONOTENSIS Linnb.
Linnb. Sjostedt, Kilim.-Meru Exped., Rept. Batr. Ded, UO
Tanganyika Territory.
3. ALGIROIDES.
Bibron, in Bory de St. Vine. Expéd. Sc. Morée, Rept. je O08
Blgr. Cat. Liz. iii. p. 48.
Synopsis of the Species.
I. A series of granules between the supraoculars and the supraciliaries ; dorsal
scales strongly keeled.
18 to 24 scales across middle of body; 12 to 17 femoral pores
on each side; 17 to 19 lamellar scales under the fourth toe;
hind limb reaching collar or ear-opening ............c00ccesee ese africanus.
33 scales across middle of body ; 9 femoral pores on each side;
24 lamellar scales under the fourth toe; hind limb barely
reaches shoulder ...............008+ Mrawenke Biisubepacnanacantineecae boulengeri
II. Supraoculars in contact with the supraciliaries; dorsal scales
feebly keeled; the laterals and anterior dorsals entirely smooth. aJleni.
1. ALGIROIDES AFRICANUS Blgr.
Blgr. Proc. Zool. Soc. 1906, ii. p. 570.
Uganda. :
2. ALGIROIDES BOULENGERI Peracca.
Perac. Atti Acc. Torin, lii. p. 531, 1917.
Uganda, — : eta ot
Proc. Zoot. Soc.—1923, No. LVI. 56
854 MR, A, LOVERIDGE; LIST OF TILE LIZARDS
3. ALGIROIDES ALLENI Barbour.
Barb, Proc. New Engl. Zool. Club, iv. p. 97, 1914,
Kenya Colony.
4, LATASTIA,.
Bedriaga, Ann. Mus. Civ. st. nat. Genova, xx. p. 307, 1884;
Blgr. Cat. Liz, ii, p. 54,
Synopsis of the Species.
TI. 89 to 52 scales across the middle of the body; no group of small plates in the
middle of the pectoral region. Dorsal scales sharply keeled, sometimes
doubtfully so in L. johnstoni.
No gular fold; edge of collar serrated; 13 to 16 femoral
pores on each Side ......cscccecese .. Johnston.
A gular fold ; edge of point even ; “10 cal 14 foionnle pores
on each side per opecEniee .. siebenrocki.
II. 52 to 80 scales across the middle of ane bore ee 56 t to 65,
more or less strongly keeled ; a group of small irregular plates
usually present in the middle of the pectoral region ; 5 to 16
femoral pores on each side, Dorsal scales not sharply keeled . longicaudata.
1, Lavasria Jounsront Blgr.
Bligr. Ann. & Mag. N. H, (7) xix. p. 392, 1907.
Tanganyika Territory,
2. LATASTIA SIEBENROCKI Torn.
Hremias siebenrockt Torn. Zool. Jahrb. Syst. xxii. p. 386, 1905.
Kenya Colony ; Tanganyika Territory.
3. LavasrrA LONGICAUDATA Reuss.
Blgr. Cat. Liz. iii. p. 55.
Kenya Colony; Tanganyika Territory.
(Lhe East African longicawdata may al] prove to be of the
var. revoili Vaill., vide Blgr. Monogr. Lacert. 11. p. 30,
1921.)
5, GASTROPHOLIS.
. Fischer, Abh. Naturw. Ver. Hamb. (ix.) i. p. 1, 1886; Blgr.
Cat. Liz. iii. p. 7.
. Gasrroruonis virrata J. G, Fisch.
Blgr, Cat. Liz. iti. p. 7.
JG “luteei Torn. Zool. Jahrb. Syst. xiii. p. 591, fig., 1900.
G. prasina Wern. Zool. Anz. xxvii. p. 462, 1904,
Tanganyika Territory; Zanzibar,
5 ?
6. IcHNorRoris.
Peters, Mon. Berl. Ac. 1854, p. 617; Blgr. Cat. Liz. iii. p. 78.
OF BRITISH TERRITORIES IN EAST AFRICA. 855
Synopsis of the Species.
I, Frontonasal single; subocular bordering mouth .......:............. tanganicana.
JI. Frontonasal pe enn) divided ; subocular not bordering the
MAXON soo da3con o0ec Me panies cela seneiet tans cue eoetene anscMntS QULAIULOSAs
]. IcuNoTROPIs TANGANICANA Blgr,
Blgr. Ann. & Mag. N. H. (8) xix. p. 278, 1917.
Tanganyika Territory.
to
ICHNOTROPIS SQUAMULOSA Peters.
Blgr. Cat. Liz. 111. p. 79,
Tanganyika Territory.
7. Kremias.
(Fitzing.) Wiegin. Herp. Mex. p. 9, 18384; Blgr. Cat. Liz. iii.
p. 80.
Synopsis of the Species.
J. Toes not or but feebly compressed, with bi- or tricarinate lamelle infer‘orly ;
GO to 77 keeled scales across middle of body.
18 to 20 femoral pores on each side; upper head-shields striated ... sperii.
II. oes strongly compressed, with unicarinate lamellie inferiorly.
68 to 82 smooth scales across middle of body; 17 to 23 femoral
pores on euch side; upper caudal scales ey keeled ; upper
head-shields rugose .......... dubabecooreod due. gocoed | CUNdAp Ot
53 to 68 keeled scales across radia ‘of body + 13 to 18 femoral
pores on each side; upper head-shielids coarsely striated ; sub-
ocular bordering the mouth.. Sain .. striata.
53 to 68 keeled seales across smneatilis of bad: ; 20" tes od. temigeal
pores on each side: upper head-shields finely striated; sub-
ocular not reaching the mouth ..........ccccccteeeeeeeeeeeeeeeee brenneri.
1. Kremias spexit Gthr.
Bler. Cat. Liz. i. p. 84.
E. sewteniata Stejneg. Proc. U.S. Nat. Mus. xvi. p. 718, 1894.
Uganda; Kenya Colony; ‘'langanyika Territory.
2. Krentas smrriir Bigr.
Blgv. Proce. Zool. Soc. 1895, p. 534, pl. xxix. fig. 4.
Kenya Colony.
3. HreMiAs srriava Peters.
Bler. Cat. Liz. 11. p. 86 (as 2. brenneri, var. striaia).
Kenya Colony.
4, JEREMIAS PRENNERI Peters. -
>) S {, o ° eee ‘i
Blgr. Cat. Liz. iii. p. 86.
Kenya Colony.
856 MR, A. LOVERIDGE: LIST OF THE LIZARDS
8. Houasris.
Holaspis (Smith) Gray, Proc. Zool, Soc. 1863, p. 152; Blgr.
Cat. Liz. iii. p. 118.
1. Honaspis gurntient Gray.
Blgr. Cat. Liz. 1. p. 118.
Uganda; Kenya Colony ; Tanganyika Territory.
Family GERRHOSAURID &.
One genus. ;
1. GERRHOSAURUS. .
Ptychopleura, part., Wiegm. Herp. Mex. 1834: Blgr. Cat. Liz.
Pills Fy JIS).
Synopsis of the Species.
TI, Tympanic shield small; ventrals in 10 rows ............... major
II. Tympanic shield narrow; ventrals in 8 rows; laterals
i” . .
NUN, dap soneadoadeasuanbes euooodondasedes cod cébaccanoboucoduon pea: 7 MANDLG UCHR
IIL. An intermediate form with keeled laterals based on single
Individual wee ee nen eee een a LOLNaiete nm eded.
IV. Tympanic shield narrow; ventrals in 8 rows; laterals
P > ’ z i
Kecledl pe sh bi ie oan ree area tin inna muntat cceannaines | UEGMOLEMEC LIS.
1. Grernuosaurus MAJorn A.-Dum.
Bler. Cat. Liz. ii. p. 121.
Kenya Colony; Tanganyika Territory ; Zanzibar.
2, GuRRMOSAURUS FLAVIGULARIS FLAVIGULARIS Wiegm.
Blgr. Cat. Liz. iil. p. 122.
Kenya Colony; Tanganyika ‘Territory ; Zanzibar.
3. GERRHOSAURUS FLAVIGULARIS, forma INTERMEDIA Lonnb.
Linnb. Sjéstedt, Kilimanjaro-Meru Exp., Rept. Batr. p. 7,
1907,
Tanganyika Territory.
4, GERRHOSAURUS NIGROLINEATUS Hallow.
Bler. Cat. Liz. iii. p. 122.
Kenya Colony ; Tanganyika Territory.
Family Scrncipa”,
Seven genera.
]. Nostril pierced in the nasal, or between nasal and supra- or postnasal or first
upper labial, not touching the rostral.
A. Pterygoid bones separated on the median line of the palate,
the palatal notch extending anteriorly to an imaginary line
connecting the centre of the eyes. Supranasals present ... Dfabuia.
OF BRITISH TERRITORIES IN EAST AFRICA. 857
Lb. Pterygoids in contact (at least quite anteriorly) mesially,
the palatal notch not extending anteriorly to between the
centre of the eyes.
Hyelids movable ; digits, if Beran, * with non-retractile
GIES “sencotobee ; tetsreseee eee Leygosoma.
Kyelids GaGetbie: tranenarene covering shen OVC) Seances Ablepharus.
II. Nostril pierced in the posterior border of the rostral, or between a nasal or 2
Jabial and the rostral.
A. Palatine bones in contact on the median line; nostril pierced
between the rostral and a se small nasal, which may be
reduced toa narrow ring ........ : .... NSeelotes.
RB
V3. Palatine bones separated on ‘thet median nine prance
present; the first upper labial entering nostril.
1. Nostril pierced between the rostral, the supranasal, .
the postnasal, and the first labial; nofrontoparictals Sepsina.
2. Nostril pierced between the rostral and the first labial. Melanoseps.
ILI. Nostril pierced in the very large rostral, with the posterior border of which it
is connected by a horizontal suture.
Mowwersen chiduscall yasancereeereneneeereent neater cere e ee neeetees mas GOLECOSE DS:
Lower eyelid transparent......... ccccscccceeseessecreeeeseeeees. s400ntias.
1. Maputa.
Mabuya, part., Fitzing. N. Classif. Rept. p. 23; Blgr. Cat. Liz.
ii. p. 150.
Synopsis of the Species.
1. Scales on the soles not spinose.
A. The subocular is distinguished from the (other) Inbials only by size and
position, and is not or scarcely narrowed eens
a. All the dorsal scales with four or more keels.
4to 7 keels on dorsal scales; 34 to 36 scales round mid-
body ; 5 or 6 supraciliaries ......... comorensis.
7 keels on dorsal scales ; 30 to 32 erates coutid ital boty: ;
4 to 5 supraciliaries.. Bee maculilabris.
7 to 9 keels on dorsal Beales: £ 30 o 32 ales roma Amie
body ; 4 supraciliaries ......... boulengeri.
b. Most of the dorsal scales Morons (planifrons with
5 keels on some scales). :
30 to 82 scales round mid-body; ear-opening with 2 or —
3 short obtuse lobules anteriorly, Prafrontals form-
ing a short median suture............ wu Orevicollis.
28 to 32 scales round mid- Ivars ear-opening ain
A projecting lobules anteriorly. Praefrontals Seqavated
by truncated end of frontal, in contact with fronto-
Nasal wes seq eces u planifrons.
28 to 32 scales fond ati Tee ; Lar nguniie win 20 or
3 long, pointed lobules anteriorly. Darietals forming
a suture behind interparietal.......00.....c.00eeeece ee. Homalocephala.
29 scales round mid-body ; ear-opening partly concealed
under the rounded scales of its anterior border ......... taitana.
ce. Dorsal scales bicarinate or tricarinate ; 80 to 32 scales
round mid-body; ear-opening with 2 to 4 short,
obtuse lobules anteriorly ............ cece cee tee eee ne eee eee quinqueteniata.
d. Dorsal scales smooth; 24 to 26 scales round mid-body ;
car-opening ovoelnanes Dice ca ny one or more
SCAlC8 vie. cies ea: uw. megalura,
13. The abenla een ane nheenioniy? parties che
reaching the lip; 30 to 34 scales round mid-body ;
ear-opening with 3 to 5 long, pointed lobules anteriorly. bayonit.
858
MR, A, LOVERIDGE: LIST OF LIE LIZARDS
II. Seales on the soles spinose and sharply keeled.
Ve
i |
30 to 36 scales round mid-body; dorsals tricarinate;
subocular largely bordering the lip. Frontoparietals
distinct or sometimes fused i WN OWNS: coonoocdosonveconodouane | CUUROMS
36 scales round mid-body ; dorsals bicar ate. subocular
reaches the lip but is much narrowed inferiorly ...... brawni.
32 to 36 scales round mid-body; dorsals tricarinate ;
hind limb reaching elbow in male and wrist in female. obséi.
32 to 38 scales round mid-body; dorsals mostly tri-
carinate, sometimes many quinquccarinate; sub-
ocular, if reaching lip, much narrowed inferiorly ...... striata.
31 scales round mid-body; dorsals tricarinate; sub-
ocular not reaching lip; frontonasal broken up into
SiahiclAs:. cs scscenctanese waceousdesusestne cen suoeseiacaceeenhes DANRCORLCIES:
Masuia comorensis Peters,
Blgr. Cat. Liz. ili. p. 163.
Kenya Colony; Tanganyika Territory; Zanzibar.
MABUIA MACULILABRIS Gray.
Bler. Cat. Liz. iii. p. 164.
Uganda; Tanganyika Territory.
MABUIA BOULENGERI Sternf.
Sternf. 8.B. Ges, naturf. Berlin, 1911, p. 248.
Tanganyika ‘Territory.
MABUIA PLANIFRONS Peters.
Blgr. Cat. Liz. ili. p. 167.
Webern diesnert Sternf.S.B. Ges. naturf. Berlin, 1911, p. 248.
Kenya Colony; Tanganyika Territory.
. MABUIA BREVICOLLIS Wiegm.
Blgr. Cat. Liz. iii. p. 169.
Kenya Colony; Tanganyika Territory.
MABUIA HOMALOCEPHALA Wiegm.
Blgr. Cat. Liz. iii. p. 170.
(Specimen in Museum of KH, A. & U. Na t. Hist. Soc, with-
out loc.)
ManutiA 'rarraAna Peters.
Blgr. Cat, Liz. iii, p. 171.
Kenya Colony.
MaBUIA QUINQUET&ENIATA Licht.
Blgr. Cat. Liz, iii. p. 198.
Uganda; Kenya Colony ; eanseaniiicn Territory ; Zanzibar.
Oo
OF BRITISH TERRITORIES IN EAST AFRICA. 85
9. MABUIA MEGALURA Peters.
Blgr. Cat. Liz. iii. p. 195.
Uganda; Kenya Colony ; Tanganyika Territory.
10. Maputa BAyontt Bocage.
-Blgr. Cat, Liz, i. p. 201.
Kenya Colony; Tanganyika Territory.
11. Mapura varia Peters.
Blgr. Cat. Liz. 111. p. 202.
M, asselit Peters in Blgr. Cat. Liz. ii. p. 201.
M. hildebrandtii Peters in Blgr. Cat. Liz. ili. p. 207.
Kenya Colony; Tanganyika Territory.
12. Masuta spraunt Torn.
Torn. Zool. Jahrb. Syst. xv. p. 585, 1901-02:
Tanganyika Territory.
13. Mapura opsri Wern.
Wern. Mitt. Nat. Mus. Hamb. xxx. p. 43, 1913.
Tanganyika Territory.
14. Maputa srriara Peters.
Blgr. Cat. Liz. ii. p. 204.
Uganda; Kenya Colony; Tanganyika Territory: Zanzibar.
15. ManuiA rrrEe@uLAnis Lonnb.
Lonnb. Arkiv for Zoologi, Band 14, no. 12, 1922.
Kenya Colony.
2. LycGosoma.
Mabuya, part., Fitzing. N. Classif. Rept. p. 23; Blgr. Cat.
Liz. iui. p. 209.
Synopsis of the Species.
I. Limbs well developed; the length of the hind limb exceeds the distance
between the centre of the eye and the fore limb. Tympanum distinct. No
supranasals, Rostral forming a suture with the frontonasal. One or more
pairs of enlarged nuchals. (Liolepisma D. & B.)
Frontoparictals much larger than the interparietal ......... clathrotis.
Frontoparietals subequal or smaller than the interparietal... kutwensis.
II. Limbs short or rudimentary (except fernandi). Ear distinguishable. Supra-
nasals present (or the nasal partially divided longitudinally and forming a
suture with its fellow). Preefrontals well developed. (Riopa Gray, p. 220.)
A. Jhimbs pentadactyle.
32 to 36 scales round the body; scales feebly tri- or quin-
QUECATINATEs cas eneheeenaeee ee eeernn oeceev catinteee cece eoreteecenen ERMAN.
26 to 28 scales round the body; smooth or feebly tri- or
quinquecarinate: 4th toc equal to or a little longer than
tTeH Sr Rig seers ester eecnccearni ie cmceeaueateaisccseesd SUNdevallic:
26 scales round the body; smooth; 4th toe much longer Domine
Ghanaians sere eee vecereeecsevseeeenee Serrandgi.
B. Limbs two-toed. Bodv elongate, serpentiform.................. anchieta.
860 MR. A. LOVERIDGE: LIST OF THE LIZARDS
III.
i)
Or
Limbs more or less developed. Ear covered with scales or very minute. No
supranasals, Pygefrontals (in the species with short limbs) minute or absent.
(Siaphos Gray.)
A. Limbs pentadactyle.
13 to 15 infradigital lamella; 24 scales round body ; ear-
opening much Tar ger than the nostril Ra veevee Kilimensis.
12 infradigital lamella; 24 scales round the eae ear-
opening not much larger than the nostril .................. thomasi.
14 infradigital lamellw ; 22 scales round body; ear-opening
not much larger than the nostril .............0... aloysii-sabandia.
B. Digits 5 or 4. Toes 5-5,
q to 11 infradigital lamellz under Ath toe.
2 to 24 scales round body ; ear-opening minute ......... graueri.
ie with 6 fingers ...........csecee cee oee grauert quinquedigitata.
Digits with 4 fingers 0... grauert quattuordigitata.
C. Digits 4-4. 12 infradigital Tania 9 in longest; 22 scales
round the body; ear- ee not much larger than the
NOs bri Wasser cs Saeko eneie Os snare “on Mee atem aaah, Oe Coe es mR OLE ROI IS:
D. Digits 3-3, 22 scales and the baiys « ear- ase much
larger than the nostril .....0 0... cece eee esveeeee ODlochmanni.
LyGosoma cLaturotis Bler.
Blgr. Ann. & Mag. N. H. (7) vi. p. 194, 1900.
Kenya Colony.
Lyaosoma kutrurNsiIs Lonnb.
Loénnb. K. Sv, Vet.-Akad. Hand]. Band 47, no. 6, 1911,
Kenya Colony.
. LyGosoMA FERNANDI Burton.
Blgr, Cat, Liz, iii. p. 304.
Uganda; Kenya Colony.
. LyGosoMA SUNDEVALLII Smith.
. Blgr. Cat. Liz. iii. p. 307.
L. leviceps Peters, Mon. Berl. Ac. 1874, p. 571, pl. fig. 3.
L. modestum Gthv, Aun, & Mag. N.H. (5) vi. p. 235, 1880.
Uganda; Kenya Colony; Tanganyika Territory; Zanzibar.
. LyGosoMA FERRANDIL Blegr.
Blgr. Ann, Mus. Civ. st. nat. Genova, (2) xviii. p. 718,
pl. 9. fig. 2, 1898.
Tanganyika Territory.
. LYGOSOMA ANCHIETE Bocage.
Blgr. Cat, Liz. iii. p. 316.
Kenya Colony ; Tanganyika Territory.
LYGOSOMA KILIMENSIS Stejn.
Stejn. Proc. U.S. Nat. Mus. xiv. (no. 862), p. 405, 1892.
Kenya Colony ; Tanganyika Territory.
8.
10.
ale
Bie
OF BRITISH TERRITORIES IN EASL AFRICA. 861
Lyeosoma 'rnomast Torn.
Torn. Zool. Jahrb. Syst. xix. p. 175, 1903.
Kenya Colony.
. LYGOsOMA ALOYSLI-SABANDIA: Peracca.
Perac. Boll. Mus. Zool. Tor. xxii. (no. 553), 1907.
Uganda.
LYGOSOMA GRAUERI QUINQUEDIGITATA Sternf.
Sternf. Ergebn. Deut.-Zentral-Afrika Exp. 1907-08, iv. (Z. 2)
p. 241, fig. 36, 1913.
Tanganyika Territory (?).
LYGOSOMA GRAVERI QUATLUORDIGITATA Sternf,
Sternf. Ergebn. Deut.-Zentval-Afrika Exp. 1907-08, iv.
(Z. 2) p. 242, fig. 3a, 1913.
Tanganyika Territory.
LyYGOsoMA MELEAGRIS Blgr.
Blgr. Ann. & Mag. N. H. (7) xix. p. 488, 1907.
Uganda.
LyYGOsoMA BLOCHMANNI Torn.
Torn. Zool. Jahrb. Syst. xix. p. 173, 1903.
Tanganyika Territory.
3. ABLEPILARUS.
Fitz. Verh. Ges. Naturf. Fr. i. p. 297, 1824; Blgr. Cat. Liz.
ii. p. 344.
Synopsis of the Species.
J. Frontoparictal and interparietal united into a single shield.
II, Frontoparietal single, interparietal distinct.
IN
26, rarely 24 or 28 scales round mid-body ..................... boutonii.
22 or 24, rarely 26 scales round mid-body ..................... boutondi var.
[peronii.
24 scales round mid-body ; tail as long as head and body. wahlbergii.
20 scales round mid-body; tail nearly three times the
length of head and body..........-.c::cceceeceeeeesseeeeterereeee megalurus.
ABLEPHARUS BOUTONIL Desj.
Blgr. Cat. Liz. ii. p. 346.
Tanganyika Territory ; Zanzibar.
ABLEPHARUS BOUTONII, var. PERONII Coct.
Blgr. Cat. Liz. iii. p. 347.
Kenya Colony; Tanganyika Territory.
862 MR. A. LOVERIDGE: LISt OF THE LIZARDS
3. ALBEPHARUS WAHLBERGI Sinith.
Bler. Cat. Liz. iii. p. 350.
Kenya Colony ; Tanganyika Territory.
4, ARLEPHARUS MEGALURUS Nieden.
Nied. Mitt. Mus. Berlin, vii. 1913, p. 89.
Tanganyika Territory.
4, ScEeLorgEs.
lVitz. N. Classif. Rept. p. 23; Blgr. Cat. Liz. iii. p. 408.
1. SCELOTES EGGELI Torn.
Torn. Zool. Anz. 1902, p. 700.
Tanganyika Territory.
5. SEPSINA.
Bocage, Jorn. Se. Lisb. i. p. 62, 1866; Blgr. Cat. Liz. ii. p. 417.
1. Supsina TETRADACTYLA Peters.
Blgr. Cat. Liz. ii. p. 420.
Tanganyika Territory.
6. MELANOSEDPs.
Blgr. Cat. Liz. ili, p. 422.
1. MELANOSEPS ATER, var. LONGICAUDA Torn.
Torn. Zool. Jahrb. Syst. xiti. p. 602, 1900.
Tanganyika Territory.
7. ScoLecosers.
Loveridge, Proc. Zool. Soc. 1922, p. 159.
1. ScoLtEcosers acontias Wern.
Melanoseps acontias Wern. Jahrb. wiss. Anst. xxx. (2) p. 19,
1914.
Tanganyika Territory.
8. ACONTTAS.
Blev. Cat. Liz. iii. p. 427.
1, Aconrras MELEAGRIS Gray.
Blgr. Cat. Liz. ii. p. 427.
Kenya Colony.
OF BRITISIT TERRITORIES IN EAST AFRICA. 863
Family ANELYYROPID”®.
One genus.
1. Wryprnta.
Gray, Cat. Liz, p. 129, 1845; Blgr. Cat. Liz. iii. p. 431.
J. Pryninta currort Gray.
Blgr. Cat. Liz. iii. p. 431.
Uganda; Tanganyika Territory.
¥ fab
ies i ah WY aug
¥
iF
SOME MAMMALS FROM JUGOSLAVIA. 865
42. On some Mammals from Jugoslavia.
By Ivor G.S. Monvacu, F.Z.8.
[Received August 27, 1923: Read November 6, 1923. ]
SysTEMATIc INDEX.
Page
Talpa ceca ceca SaVi oi. cci cts eeceet cence cnneesaneees 866
Glis glis postus, subsp. n.. Sar atece aaa la aetna OOO
Bvotomys glareolus sant, emeinart h. seohuecesen OOM
Evotomys gorkd, 8p. Ne wi. cecccecceree eee cencenseereveeeess 867
Microtus agrestis punctus, SUSE [Ns “coosneacgeannacens Shots)
Microtus arvalis arvalis Pallas ............c0000. 868
Microtus arvalis levis Miller ..............:00:000-0200 869
Microtus arvalis calypsus, subsp. n. ........-..-...... 869
(Measurements throughout are in millimetres.)
Among the areas least fully inspected in the preparation of
G.S. Miller’s Catalogue * (1) were those parts of the Balkans
now grouped together as Jugoslavia. In consequence of the
generosity of Mr. Uvaroff, of Lord Swaythling and Mr. Cotton,
material from these regions has recently been added to the
British Museum collection. The former made possible the pur-
chase of a part of the collection formed by V. Martino in Croatia,
Montenegro, and Serbia; the latter made possible a small scale
expedition to Slovenia and Croatia, undertaken by I. Montagu
and W. K. C. Cotton.
In addition to the specimens described here and by V. and EK.
Martino in (2), and to specimens not of immediate interest, these
collections include abundant series of Apodemus. No account is
given here of these series, since they are all referable to the races
A, s. sylvaticus, A. s. dichrurus, A. agrarius, and A. epimelas, and
do not materially affect existing knowledge of the range of those
forms.
The described material gives a number of noteworthy results.
It indicates the presence in the Croat highlands of 7’. ceca, of a
peculiar dormouse resembling that of southern Italy, and of a
remarkable new species of Hvotomys. An EH. glareolus of the
Croat lowlands, resembling that of the western Alps, is recognized
as peculiar, as is also a race of J/. agrestis from the Slovene Alps.
The presence of the northern and western J. a. arvalis in the
Croat lowlands, and that of the southern and eastern JZ. a. levis
in the Slovene and Croat highlands, together with the occurrence
of a new form resembling the former in eastern Serbia, throws
important light on the distribution of these two subspecies.
These circumstances may indicate an overlapping of the ranges
of the two forms, but, in the writer’s opinion, more probably
* (1) Catalogue of the Mainmals of Western It urope. BM. '(N.H), 1912.
(2) Annals and Magazine of Natural History, 9th ser. ix. p. 413, 1922.
866 MR, I. G. 8, MONTAGU ON SOME
reveal the presence of a pocket of JZ. a. arvalis extending along
the Slavonian plains.
TALPA CACA CECA Savi.
L'ype locality. Italy. ©
Material examined and Dimensions. Two adults (1 o,19)
collected by Montagu in June 1923 in Kupjak, Croatia.
HB. Th HE.
BSe Adult AeouGso3y oy eee agate 24 17
GOs) See OM OSLO OSU Nec NN acu alee 275 17
Description. Both the skulls are typical in all respects except
size. The skull dimensions throughout, and the length of the
hind feet, while less than those of 7’. ewropeus, are larger than
is usual in 7’. ewca; they afford, indeed, new maxima for the
latter form. The mesostyles of m2 and m3 exhibit in each case
the characteristic deeply-notched apex.
‘The presence of this species in the Croat highlands tends to
confirm the view that its range is continuous across the Balkans,
extending from northern Italy and Switzerland into Asia Minor.
GLIS GLIS POSTUS, subsp. n.
J'ype locality. Veliki Dergonel, the Gorski Kotar, Croatia.
Material ecamined and Dimensions. Three adults (2 ¢,1 ¢)
collected by Montagu and Cotton in July 1923 on Veliki
Dergonel.
HB. T. EE a ga
Go neAdulte gs. 2n7Aog rae eee BT 761 31 19°5
314, Seren a laree canoe odeloo odode 181 171 816 21
Type 86. Be Sob iBe- Hl ban caedbo 178 150 32 20
1 Damaged.
Description. The dorsal skin colour is yellow-grey, slightly
darkened by the longer blackish hairs; the tail is essentially
concolour with the dorsal fur. The animal in dorsal colour thus
differs noticeably from G. g. glis, showing no trace of the pinkish-
brown tint of that race, and approximates to G. g. italicus. The
tail of the latter, however, is characteristically darkened by heavy
_ pinkish-brown wash, and is accordingly readily distinguished
trom that of G. g. postus.
‘he dimensions of hind foot and ear are less than the average
dimensions of G. g. italicus, but larger than those of G. g. glis.
The skulls are in no case complete, but examination of the
available fragments indicates robust cheek-teeth and a general
large size of the constituent bones, recalling the characteristic
properties of G, g. italicus.
In view of the parallelism between the dormouse of the Croat
highlands and that of southern Italy, it is well to recall that
dormice from north Italy and Trieste, localities intermediate in
point of space, resemble neither in any way and are exactly
typical G. g. glis.
MAMMALS FROM JUGOSLAVIA. 867
EvoroMys GLAREOLUS SOBRUS, subsp. n.
Type locality. Rescetavi, Nova Gradisca, Croatia.
Material ecamined and Dimensions, Two adults (@ ) collected
by Martino in September 1921 in Rescetari.
HB. ft 180 185
elena, AACS 25 ISL ELPA coopcsdnanan cee 93 43 18 13
Type 21.12.1.16. Bt pis LOS iat bey es sue Nee 101 46 18 13
Description. 'The dimensions indicate that this lowland bank-
mouse 1s a small one. ‘The dorsal colour is yet more drab than
that of the typically dull skin of HZ. 9. helveticus. From this and
other forms the present one is readily to he distinguished by the
shortness and peculiar coloration of its tail. In both the speci-
mens this appendage is dorsally of a pale dust colour and ventrally
a nearly clear white. In all described neighbouring races, Z. g.
helweticus of the western Alps, Z. g. nageri of the eastern Alps,
and #. gorka of the nearby highlands, the tail is dorsally a dark,
neatly seal, brown and ventrally a distinctly yellow-buff.
The skull of this animal is in no way distinguishable from a
typical #. ylareolus skull of small size, such as that of EH. g. hel-
veticus. In both specimens the third inner angle of m3 is
present.
EVvoroMYs GORKA, sp. 0.
Type locality. Zalesina, the Gorski Kotar, Croatia.
Material examined and Dimensions. One adult (2) collected
by Montagu in June 1923 in Zalesina.
HIB. ba 2) ae cee oh
Fy Gils NGI) Or FAC BLED occencuocoschsosce ) LEY/ 54 19 12°5-
Description. This specimen is a peculiar one, differing in its
characters of skin and skull no less markedly from Z. glareolus
sobrus of the nearby plains than from previously described and
more distant races of Hvotomys.
The body size and skull length are equal to those of the largest
European forms of Hvotomys (LH. rufocanus, KH. cwsarius, H. glareo-
lus nageri, and /. g. hallucalis). ‘The tail is relatively short.
The fur is dorsally long and shaggy and of a bright rufous
colour. The flanks are without the usual grey shade, and:in fact
are so bright an orange-brown that the dorsal stripe pattern
typical of the genus is obscured. ‘The grey colour of the belly is
a trifle darker than that characteristic of neighbouring /’. glareo-
lus forms.
The skull of this specimen, a fully- developed adult, shows a
long palate and bull of great size. The nasals * are longer and
* The nasal measurements given by Miller for H. g. nageri are doubtless over-
estimates. His dimension 8°0 mm. seems to the writer barely to equal 7°8 mm.
868 MR. I, G. 8. MONTAGU ON SOME
narrower than those of /. g. nageri, the diastema is long, and the
incisive foramina are of great length. The length of the skull
from the condyle to the anterior end of the bulla is 9°25 mm.
In the largest specimens of Z. glarcolus this dimension’ attains
only 8°65 mm., and in YZ. cesarius only 86 mm. The enamel
pattern of the teeth exactly resembles that of a typical LZ’. yla-
reolus, in which the third re-entrant angle on the inner side of
m3 is present.
MICROTUS AGRESTIS PUNCTUS, Subsp. n.
Type locality. Bled, Slovenia.
Material examined and Dimensions. One young adult (2)
collected by Cotton in June 1923 in Bled.
Hb. Ts Hr. E.
Tyne 304. Young adult 9, 16.6/23. ........... 87 37 19 12.
Description. The dorsal colour is almost black, relieved on the
shoulders and face by a slight tawny peppering. In this respect
the animal nearly resembles young specimens of J/. a. levernedii.
In AL. a. levernedii, however, the flanks shade gradually into the
ventral greyish-yellow wash; in the present specimen the flanks
are tawny, and readily contrast with the ventral white. The
white of the belly, though interrupted here and there by the
appearance of the slaty bases of the hairs, shows no trace of
yellow. ‘The tail is bicoloured and the feet dark.
Measurements of the reconstructed skull indicate the presence
of a long brain-case like that of AZ. a. levernedii. The skull pre-
sents a unique character in that the anterior edge of the mas-
seteric plate is inclined forward. ‘The distance from the incisors
to the lower anterior corner of the plate is accordingly less than
in similarly sized skulls of other forms. ‘Che pattern of the teeth
is that of typical .W. agrestis, in which the fourth inner angle of
m1 is not present.
MIcROTUS ARVALIS ARVALIS Pallas.
Type locality. Germany.
Material evamined and Dimensions. Seven adults (2 ) collecte|
by Martino in August and September 1921 in Nova Gradisea,
Croatia.
HB. 4k HF. K.
91.12.18. Adult 9,17.8.21. wo... 100 37 15°5 11
DUTONOs Usa) OPM BL Ollie cee 100 35 16 11
DMO TIO ay Pepe Gholi en mn eh 104 3450 164
CHIU Dy WG, Uscceneen ye JOD 37 158 103
ERI, iy OR TEMM Ils cis 97 35 15 11
OMAR, Beal OME TS a a Sercicco hoo 110 36 165 12
aga ee OGLE LOI a Lemon) a koy) 36 155 114
Average of seven adults ................c0 00 10271 35°8 155 111
MAMMALS FROM JUGOSLAVIA. ° 86Y
Description. The dorsal fur is a lighter shade of yellow than
that usually seen in more northerly specimens. The belly colour
is white, often with the slaty bases of the ventral hairs readily
perceptible; in three specimens there are traces of yellowish
wash.
‘The skulls show the characteristic short brain-case and arc-
like zygomas of Jf. a. arvalis.
Microtus ARvALts Levis Miller.
Type locality. Rumania.
Material examined and Dimensions. Two young adults (1 ¢,
1 2) collected by Montagu and Cotton in June 1923 in Bled,
Slovenia. One young adult (¢) collected by Montagu in June
1923 in Kupjak, Croatia.
brEp. HRB. AN HI. Yr.
47. Young adult gf, 16.6. 23. ............ 102 29 16) pA,
305. 5 APN, BL On Os dere nace waee 88 28 14, 11
Kupsak.
65. Young adult ¢, 26.6. 23. ........... 99 30 18 12
Description. The dorsal brown coloration of the skins is
noticeably bright. In each example, and particularly in No. 47,
this brightness is present also on the flanks in such a manner as
to form a line of demarcation more definite than is usual in
M. arvalis. The belly is silvery without yellow wash. The feet
are nearly seal-brown.
The skulls of these specimens are those of typical I. a. levis,
showing the characteristic long brain-case and straight parallel
zy gomas.
MICROTUS ARVALIS CALYPSUS, subsp. n.
Type locality. Nova Varos, Serbia. ;
_ Material examined and Dimensions. One adult (3%) collected by
Martino in June 1921 in Nova Varos.
HB. 4 HF. KE.
Type 219. Adult 6 ?, 18.6.21. ...........0-6 101 38 15 ll
Description. The dorsal colour is pale yellow, distinctly lighter
than that of I. a. arvalis, slightly overlaid with grey in a median
line. The ventral surface is clear silver. ‘The feet are whitish
yellow and the tail bicolour, dorsally seal-brown, ventrally
whitish yellow.
The size of the skull and hind feet resembles in its smallness .
M. a. arvalis rather than J, a. levis.
The skull shows the remarkable combination of a long brain-
case and arc-like zygomas. In other respects it is not peculiar.
Proc. Zoou. Soc.—1923, No. LVI. 57
-870 SOME MAMMALS FROM JUGOSLAVIA.
Skull dimensions,
To. ceca. 1. 2. 3. 4, 5. «6. 7. 8.
63. Adult 9,317 Vt 15°75 946 425 206 12:0 12°3.
60. ,, 9,318 1085 158 9°5 425 20°38 12°25 125
EL. gorka. 1. 2. 9 10. 11. 12. 18. 6. Ta. 8a.
Type Gl. Adult 9,26°65 1386 43 122 68 79 82 166 GO 57
E. g. sobrus.
21.12.1.15. Adult $, 244 13°8 $1 6117 645 715 70 150 56 53
Type 21.12.1.16. ,, 9,234 131 40 111663 67 655 140 6°75 5°5
1. Condylobasal length. 8. Mandibular tooth-row (ex. inuci-
Q. Zygomatie breadth. ‘ sors),
3. Breadth of brain-case. 8a. Mandibular check-teeth.
4. Depth of brain-case. 9. Interorbital constriction.
5. Rostral breadth. 10. Occipital breadth.
6. Mandible. 11. Occipital depth.
7. Maxillary tooth-row (ex. incisors). 12. Nasal.
Ta. Maxillary cheok-teeth. 13. Diastema.
NOTES ON EAST AFRICAN SNAKES. 871
43. Notes on Bast African Snakes, collected 1918-1923.
By Artuur Loventiper, IF.H.S., C.M.Z.S.
[Received August 14, 1923: Read November 6, 1923.]
The 221 snakes collected during 1915-1917 have already been
dealt with in a paper written in the field during the Hast Africa
Campaign*. The present paper deals with some 290 specimens,
of which 286 were collected by the writer or his native collectors
during the years 1918-1923. Some few earlier captures are
referred to but not included in the figures.
The present collection contains representatives of all the Kast
African families. 34 of the 54 genera, and 57 of the 140 seeped
species. These numbers are distributed as follows:— |
Family. Genera, Species. Pes
Tvphlopidse ............ 1 8 23
Glauconiide ............ 1 4 15
TENET HD Gopdoacaeoe 1 1 8
Colubridsy Vee ee 28 : 41 201
Wirperidset il? 2.0.00) ee 3 6 39
Total ... 286
Only two new species resulted from this material, viz. Typhlops
excentricus Procter, and Geodipsas proctere Loveridge, which
have been described elaewhere.
The principal localities mentioned in the following pages are :—
Kenya Cotony.
Frere Town, Nairobi, Uasin Gishu Plateau.
TANGANYIKA TERRITORY.
Dar es Salaam District.—Dar es Salaam.
Lushoto District.—Usambara Mtns., Gonja.
Moshi District.—Kahe, Garagua.
Arusha District—Arusha, Longido Mt.
Morogoro District.— Dakawa, Bagilo, Morogoro, Mkata River,
Rudewa, Ilonga, Kimamba Stn., Chanzuru, Tindiga,
Kipera, Madazini, Kilosa, Kideti.
Iringa District.—Rumruli.
Dodoma District—Pwaga, Itende, Mpanira, IJkikuyu,
Kidenge, Mpwapwa, Dodoma, Suna, Gwao's.
* Loveridge, “Notes on Snakes in East Africa.’ Jour. E. A. & U. Nat. Hist.
Soc. No. 13, 1918, pp. 315-338. :
bY
872 MR, A. LOVERIDGE: NOTES ON
Kondoa-Irangi District—Mtali’s, Mkalama, Zengeragusu,
Usshora, Ulugu.
Tabora District.—Tabora, Izikisia, Luguo.
Mwanza District—Sanga, Shanwa, Sagayo, Nyambita,
Mwanza.
Bukoba District.—Bukoba, Kabare.
PorruaueEsE Hast AFRICA.
Lumbo on mainland opposite Mozambique Is.
With six exceptions the identifications were all carried out
in East Africa, and I take this opportunity of acknowledging
my indebtedness to Dr. G. A. Boulenger and Miss Procter for
examining these reptiles and confirming determinations of others.
My thanks are also due to Dr. H. A. Baylis and Mr, 8S. Hirst
for determining the parasitic worms and ticks.
As a general rule measurements are not given unless they
exceed the maximum given in the Catalogue of Snakes. They are
then given in inches, followed by snout to vent and tail measure-
ments in millimetres.
TyPHLOPIDEA,
TyPHLOPS MANDENSIS Stejneger.
Blgr. Cat. Snakes, iii. 1896, p. 587.
A single specimen was dug up in some building operations near
Government House, Morogoro (23.ix.17), and’ brought to me
alive. As the species was described from the Isle of Manda oft
the Lamu Coast of Kenya Colony by Stejneger as long ago as
1894, and no second specimen has been taken to my knowledge,
this new record from the interior of ‘l'anganyika Territory is of
considerable interest. It measured 184 inches (463+7). Sex 9.
Alive it was almost colourless except for a little buff on the belly,
whereas Stejneger’s specimen was ‘“‘ Uniform pale greenish grey
above, pale buff beneath.” Iam indebted to Dr. Boulenger for
making the identification, and the specimen has been given to
the British Museum.
lypuLors puNCTATUS Leach.
Blgr. Cat. Snakes, i. 1893, p. 42.
Three specimens from Dar es Salaam and Tindiga.
The former measured 1384 2°5 and 130+2 mm., with diameter
at mid-body of 4 and 5 mm. respectively, so that the diameter
goes into the body length 26 to 35 times. Grey above with or
without small black blotches and dorsal stripes following the
edges of the scale-rows.
EAST AFRICAN SNAKES. 873
Tyrunors DINGA Peters.
Blgr. Cat. Snakes, i. 1893, p. 45.
A specimen measuring 33g inches (850410) was caught on
27.xii, 22 at Kilosa, having one above ground after the first
heavy downpour of the lesser rains. In colour it was almost
identical with the figure in Peters’ ‘ Reise nach Mossambique,’
but a week later it was a dark metallic silver, doubtless a sign
that it was about to cast its skin.
TyPHLOPS MUCRUSO var. HUMBO Peters.
Bler. Cat. Snakes, i. 1893, p. 46.
Thirteen specimens from Ilonga, Kilosa, Kipera, Madazini,
Kadenge, Mpwapwa, and Usshora.
The largest female of these measured 232 inches (590+5)and a
male measured 183 inches (460+6); it was unearthed by some
pave digging a trench at Kilosa on 27.i.21, when I made a note
“‘whitish or flesh-coloured, eyes completely hidden.” It had 35
mid-body scale-rows, diameter went 36 wna into total length
and the tail 77 times.
On 18.11.21, having turned the “ sclouilees Typhlops” out of
the earth where I had been keeping it, I was astonished to find
that it was now the colour of ordinary Kilosa specimens of var.
humbo and the eyes distinct having taken up pigment. I do not
know what the explanation was unless that it had sloughed just
before capture and pigmentation was delayed in the new skin, but
I am perfectly certain that there was no trace of the eyes on
27.i.21, as I examined it very closely with a lens believing I had
something new. As the visibility or otherwise of the eyes is an
important key-character this is rather interesting.
The coloration was now as follows:—Black above, marbled or
speckled (the specks being sometimes confluent) with pale bluish
grey. Beneath, the pale blue is about equal to the black in
extent. A young specimen had the pale blue brighter, almost
bluish slate. It is the constant colouring of all my specimens
from Morogoro, Kilosa, and Mpapua.
The eye in my specimens is partly under the przocular, not as
shown by Peters* or figured by Sternfeld t for Z. mucruso.
Both these authors show one scale (preocular) between the ocular
and the nasal in forma typica, whereas in specimens of var.
humbo from Tanganyika Territory there are distinctly two. In
one young specimen the nasal is in contact with its fellow behind
the rostral, and nearly so in the adult mentioned above.
'The Usshora specimen was swallowed by a Bird Snake (7. kirt-
landii) with which it shared the vivarium.
* Peters, Reise Mossamb. iii. p. 95, pl. xiii. fig. 3 :
+ Sternfeld, Fauna der deutsehion Kolonien, Reihe Ill. Heft 2, p. 12, fig. 8.
874 MR. A. LOVERIDQE: NOTES ON
TyPHLOPs EXCENTRICUS Procter.
Procter, Ann, & Mag. N. H. ser. 9, vol. ix. p. 685, June 1922.
As only the type-specimen was taken during the two yeas
T resided at Kilosa it must be decidedly uncommon.
GLAUCONIIDS.
GLAUCONIA DIsTANTI Bler.
Blgr. Cat. Snakes, i. 1895, p. 62.
As the type-locality of this species is Pretoria it was somewhat
of a surprise to find it so far north, but Dr. Boulenger has
kindly eximined the Morogoro specimens for me. - ‘Thirteen
specimens in all were taken from Morogoro and Kilosa.
The largest measured 6 inches (139-+ 12), but the largest BE
six Kilosa “snakes was only 32 inches (78 +6).
The tail of the type was ineluded 12 times in the total length,
but the range of Kilosa snakes is from 9 to 13 times, one speci-
men having a body length of 65 mm. had a tail 7 mm. long.
The diameter of the body into total Jength ranges from 48 to 56
as against 65 times in the type. Whilst the type had five lower
labiuls, two Kilosa snakes have four and three others only three!
GLAUCONIA MERKERI Werner.
- Wern. Jahresh. Ver. Nat. Wiirttemb. Ixv. 1909, p. 61,
A single female fron Mtali’s, Mkalama Dist., where it was
found wriggling on the surface of the ground beneath a fig-tree
about 9 p.m. on 10.x.22. The species is, I believe, only known
from the types which came from Moshi much farther east.
My specimen is a female and is larger than the type by 52mm.
Length of head and body 212 mm. Length of tail 15 mm. Dia-
meter of body 3°5 mm. The diameter, therefore, goes 64:8 times
into the total length as agninst 80-87 times in the types. ‘Lhe
tail goes 14 times as against 114-134 times in the original spe-
cimens. It hes 4 upper labials.
The colour is uniform blackish, the borders of the scales lighter.
eee red wath a black ring.
Guaneones EMINI Blegr.
Blgr. Cat. Snakes, i. 1895, p. 64.
A specimen found in a heap of dry manure on the golf links at
Dar es Salaam, 4. xii. 22, agrees with Boulenger’s description of
G'. emini except in the following points :—
(i.) Flesh-colour. In life, however, semi-transparent, the
dorsal scales being minutély stippled with brown, the beily scales
quite transparent. A pinkish tinge imparted to the whole by the
blood-vessels which ‘can be plainly seen. ‘The type of G. emini
EAST AFRICAN SNAKES. 875
uniformly blackish, similar to specimens collected by the writer
at Nairobi, but there is a specimen from Kosa Kola, Lake Nyasa,
in the British Museum, which is also pale tending to orange on
the back and has been referred to G. eminz by Dr. Boulenger.
(ii.) Posterior border of the rostral does not nearly extend to
the level of the eyes as against “not extending quite to the
level of the eyes.” In the Catalogue description the Lake Nyasa
specimen is like mine in this respect also.
(iii.) The diameter of the body goes 57 times into the total
length as against 50-55 times.
I think there is no doubt, however, that I am correct in
referring my specimen to G. emini, which has already been
recorded from the extreme west of the Territory, the type-locality
is Victoria Nyanza. ‘The present record shows that its range
extends right to the coast in the east.
Length of head and body 102 mm. Tail 13 mm. Diameter
of body 2mm. The length of tail goes 94 times into the total
length.
GLAUCONIA LONGICAUDA Peters.
Blgr. Cat. Snakes, i. 1895, p. 66.
Four specimens taken at Lumbo, Mozambique.
The longest of these measured 4? inches (110412), all were
females. ‘This snake is far more slender than an average earth-
worm and of a very transparent flesh-colour. All were taken
among the roots of bushes when some land was being cleared of
stumps for a camp site.
One which was brought to me in the early morning was
dropped into a cigarette tin containing several Lycophidium
semiannulis and Aparallactus capensis from the same habitat.
On opening the tin about 6 p.m. the worm snake was found to be
missing, but by holding up the other snakes to a strong acetylene
licht it was located in the stomach of a Cape Black-headed Snake.
It was the work of a few minutes to chloroform the latter,
but the worm snake was already too far digested to be worth
preserving.
PYTHONIDA.
Pytnon sepxz Gmel.
Blgr. Cat. Snakes, i. 1893, p. 83.
Kight specimens, Rumruli, Ilonga, Chanzuru, Kilosa, and
Usshora. ‘
Three of these were young specimens sent me by Mrs. Billinge
- of Rumruli, Ivinga, who wrote that the natives had killed a female
python, whose unstretched skin measured 20 feet, and which had
34 young ones with it. She adds that the natives never attack
876 MR, A. LOVERIDGE,: NOTES ON
the pythons unless they have first taken a dog or goat-kid,.and
to judge by the number of pythons so killed they must do an
appreciable amount of damage in the district.
A native youngster coming up from Kilosa village heard an
outcry by the roadside which he thought was a young kid.
Parting the grass about a bush he saw a large python which was
swallawing a reedbuck. He took me to the spot, which was a
very likely one, but the. python had gone; I think it was more
‘probably a young bushbuck that was taken.
Many parasitic worms were found in the stomachs of my
specimens, viz, Lothridium pythonis Blainv., Polydelphis attenuata
Mol., Ophidascaris jfilaria (Duj.), and other indeterminate im-
mature Ascarids. A Jayval Filarian was also found coiled up
in pimples which it causes on the skin. Snakes so affected
rapidly lose their glossy and iridescent appearance, refuse to
feed, and succuub within a few months. ‘The disease appears
ta be infectious to the extent that another snake introduced into
the saine cage, after the removal of the dying snake, became
infected.
CoLUBRIDA.
‘TROPIDONOTUS OLIVACEUS Peters,
Bigr. Cat. Snakes, i. 1893, p. 227.
Four specimens from Dar es Salaam, Bagilo, and Kabave.
Found in swampy grass-lands and along river-banks.
All the specimens were rather small. A Dar es Salaam female,
kept in captivity, laid eight eggs measuring 17x8 mm. on
15.vi. 18. A Bagilo specimen killed 22, vii. 21 had three eggs in
the ovary measuring 226 mm., this was an extremely small
specimen measuring 262 mm. in head and body; the tip of the
tail was missing, as is often the case with snakes of this species.
It would be interesting to know what enemy it is that deprives
them of their tails.
Bagilo specimens have the borders of the ventrals bright mauve
and not olive.
The Dar es Salaam snake swallowed two frogs (anu nutti),
hind-legs first, on 22. vi. 18, and another, head first, on 24, vi. 18.
Boopon tingeatus Dum. & Bib.
Blgr. Cat. Snakes, i. 1893, p. 332.
Thirteen specimens from Mkata River, Kimamba, Pwaga,
Ikikuyu, Gwao’s, Sanga, Shanwa, T'abora, and Bukoba.
The House Snake is usually found about the habitations of
man, among rubbish in outhouses, under old sacks, or beneath
sheets of iron in the open. It is not infrequently met with
crossing paths, and when detected doing so it usually remains
EAST AFRICAN SNAKES. 877
perfectly motionless until very closely approached, One large
specimen on path entered a fissure in black cotton soil and on
being dug out disgorged a large rat (Rattus coucha microdon).
A bright orange-coloured specimen was met with at Dodoma
(1 accidentally trod upon it in coming out of my room). I have
never met with an orange variety before, and it faded to straw-
eolour when preserved. Young specimens tend to be rufous and
have fine reticulations which disappear in the adult, which is
plumbheous in most of the above specimens.
The reason for their frequenting houses is to be found in
their almost exclusive diet of rodents. One such specimen had
swallowed a rat and was found with a foot of its tail hanging
out of a crevice in the stone basework of the house. The snake
was so wedged in that the stone masonry had to be removed
hefore the snake could be extricated. A 2 feet 8 inch snake at
Gwao’s, with greatly distended stomach, held two unstriped grass
rats (Arvicanthis abyssinicus newmanni) in its stomach.
On a kopje at Tabora I came upon a Striped-bellied Sand
Snake swallowing a young House Snake. When I appeared the
Sand Snake disgorged its prey, which I put iu my pocket, but
found it dead on reaching home, presumably killed by the venom
of the back-fanged species.
Vicks (Aponomma leve Linn.) were found in one Kilosa
specimen.
@ Lycoriipium acurinosrre Giinth.
Blgv. Cat. Snakes, 1. 18938, p. 338.
The day before sailing from Dar es Salaam (25. v. 23) I en-
countered a snake on the sea-front road by the golf links; it was
holding its snout downwards, after the manner of an Atractaspis
viper, and pushing against a fragment of coco-nut shell as if
seeking shelter beneath it from the sun. The snake showed no
sense in pushing round and round the fragment. 1 confess
I mistook it for A. restrata, picked it up by its stwmpy tail,
dropped it in a bag, and sent it on board.
T did not examine it again until the second day at sea, when
I found it not only dead but very decomposed. Close examina-
tion showed at once that it was a Lycophidium, and according to
Boulenger’s key * it was E£. acutirostre, hitherto only known from
Zanzibar Is. As I was unable to count the ventrals or compare
it with the full description, I have marked the determination
with a query. The specimen was too decomposed to preserve.
_ It seems to me from its striking superficial likeness to Aéract-
aspis rostrata that this species of Lycophidium has adopted similar
burrowing habits, and it may be on this account that it is so
rarely met with, less than a dozen specimens being known I
believe.
* Boulenger, “A List of the Snakes of East Africa... ete.” P.Z.S. 1915, p, 620.
878 MR. A. LOVERTIDGE: NOTES ON
LyYCOPIHIDIUM SEMIANNULIS Peters.
Blgr. Cat. Snakes, i, 1893, p. 339.
Eight specimens were taken at Lumbo, where they lived in the
surface soil amongst the roots of grass.
The largest male measured 7? inches (170+21) and female
93 inches (203435). None of the specimens attempted to Lite
when handled.
LyYCcOPHIDIUM CAPENSE Smith.
Blgr. Cat. Snakes, i. 1893, p. 339.
Two specimens from Kilosa, found on paths.
The largest specimen I have yet come across was taken at
Morogoro, 183 inches (405 + 52).
A nematode (Physaloptera sp.) was taken from a Kilosa snake
Lycopnipium gacksontr Bler.
Blgr. Cat. Snakes, i. 1893, p. 340.
A single specimen was taken in a tent at Longido after a night
of heavy rain. The type-locality is Kilimanjaro, so the present
record of this rare snake extends our knowledge of its westward
distribution somewhat.
It was taken in the act of attempting to swallow a skink
(Lygosoma ferrandii), which was rather a big mouthful for it.
SIMOCEPHALUS NYASS.z Giinth.
Blgr. Cat. Snakes, i. 1893, p. 347.
A single specimen at Lumbo during the demolition of a
termite hill. Female 22 inches (433 + 122).
CuLonoryuis NEGLECTUS Peters.
Blgr. Cat. Snakes, ii. 1894, p. 94.
Five specimens from Nairobi. As a variant from its usual
habitat one put its head out of a leather hair-brush case, having
reached the dressing table from an open window.
One was seen swallowing a frog—its usual diet—but another
had no less than three skinks (Ablepharus wahlhergit) in its
stomach.
An immature ascarid was also found in one specimen.
CHLOROPHIS IRREGULARIS Leach,
Blgr. Cat. Snakes, ii. 1894, p. 96.
Single specimen from Bukoba, collected by Mr. N. C. Miller.
EAST AFRICAN SNAKES, 879
PHILOTHAMNUS SEMIVARIEGATUS Smith.
Blgr. Cat. Snakes, ii. 1894, p. 99.
Twelve specimens from Dar es Salaam, Ilonga, Kimamba,
Kilosa, Kabare, and Lumbo.
None of these exceeded in size a female from Morogoro,
measuring 513 inches (837+466), taken 25.xi.17,, One speci-
men had only 149 ventrals and two had 159 sub-caudals.
The coloration of Lumbo specimens was strikingly different
from the grass-green examples, of which a long series were taken
previously. In Lumbo specimens the head and neck were pale
green; body mauve; both freely speckled with black. The tail
was plumbeous; throat china-white; belly and gub-caudals
whitish with a mauve tint. Ventral keels of a mauve colour.
The tongue at tip and base was black, but the middle portion
bright Cambridge-blue. All four specimens taken at Lumbo
were the same.
Attracted by the outcry raised by birds, I captured the Frere
Town specimen whilst it was descending the almost vertical tree-
trunk, taking advantage of every irregularity in the bark.
A Spotted Wood Snake was found swallowing a gecko (Hemi-
dactylus mabouia) in the fowl-house and only the tail was to be seen.
Immediately it was approached the snake disgorged the gecko,
and on being seized by the tail inflated the neck and anterior
portion of the body vertically, and struck at my hand repeatedly.
Here then is another tree-snake with the same habit of inflation
when alarmed or annoyed as the Boomslang and Bird Snake.
Another specimen on being caught in a thatch disgorged a
gecko of the same species, and a third snake had swallowed a
young toad (Bufo regularis).
During a flood one specimen was found on a maize stalk,
five feet above the water. A young one was recovered from
the stomach of a One-streaked Hawk (Kaupifalco monogrammica).
TuRASOPS RoTUSCHILDI Mocq.
Mocq. Bull. Mus. Paris, 1905, p. 286.
A specimen sent me from Bukoba extends the range of this
scarce reptile westwards; it was described from British Mast
Africa. I have also had the pleasure of examining three more
specimens collected by Mr. J. A. ‘Turner in the Yala River region
near Mt. Elgon.
RHAMNOPHIS JACKSONII Giinth.
Blgr. Cat. Snakes, iii. 1896, p. 632.
One male from Kabare.
A specimen of this snake was killed at Muthaiga, near Nairobi,
in 1919, by Mr. A. J. Klein, and is now in the Nairobi Museu.
It measured 7 feet 53 inches (16714584), and eight specimens
were collected by Mr. ‘Turner on the Yala River.
889 Mh, A. LOVERIDQE: NOTES ON
I have never seen the resemblance of this snake to the black
phase of Dispholidus typus remarked upon. Their similarity cf
appearance is extraordinary and when alive they are indistin-
guishable, except perhaps for the slightly larger eye of the
Boomslang—both are tree-snakes and found in the same locality at
times. Like all snakes that are large and black they are indis-
criminately called ‘‘ Black Mambas” by the European residents.
CoRONELLA SEMIORNATA Peters,
Blgr. Oat. Snakes, ii, 1894, p. 195,
Two examples from Dar es Salaam and Sagayo, Mwanza.
The latter specimen is very juvenile (210 mm. over all), and
has a small seale between the rostral, internasals, and nasals,
apparently split off from the rostral.
CoRroNELLA SCIIEFFLERI Sternf,
Sternf. Sitzb, Ges. Nat. Fr. Berl. 1908, p. 93.
I have not been so fortunate as to collect this rare snake
myself, but a specimen in the Nairobi Museum was obtained by
Capt. Rainsford just south of Lake Rudolph in 1918. ‘This new
record is, therefore, far north of the type-locality Kibwezi on the
Uganda Railway.
GRAYIA THOLLONI Mocq.
Mocq. Bull. Soe. Philom. (8) ix. 1897, p. 11.
Mr. N. C. Miller sent me a specimen for determination which
he had collected at Bukoba, 1922.
TfoMALosoma LuTRIXx Linn.
Blgr. Cat. Snakes, 11. 1894, p. 273.
A single specimen from Kabare, Bukoba, 10. i. 23.
This female, though small (290+ 38 mm.), was very bloated,
having no fewer than ten eggs, measuring 128 mm., in its
oviduct which extended forward to the region of the heart.
PROSYMNA AMBIGUA Bocage.
Blgr. Cat, Snakes, ii. 1894, p. 248.
Three specimens. One of these was taken in a termite nest at
Lumbo; the other two were found in a bottle without data in a
German House at Morogoro.
One of these had paired priefrontals and 155 ventral scutes.
DasyYPELtis scaBRA Linn,
Blgr. Cat. Snakes, ii. 1894, p. 354.
Six specimens were collected from Kilosa, Kipera, and Zenger-
agusu in Tanganyika Territory and Lumbo in Portuguese East
Africa,
EAST AFRICAN SNAKES. 881
The Kilosa and Kipera specimens are of the type that appears
to be a mimic of Causus rhombeatus, but C. rhombeatus is
not found at Kilosa, and where this Night Adder is common at
Nairobi you get an all-black or all-brown variety of the Egg-
eater. The Zengeragusu reptile was a most interesting variety.
Colour iron-clad grey with a single row of brick-red dorsal spots.
This snake was literally covered with ticks (Aponomma lcve
Linn.), seventy-five of which I collected.
Gropirsas rroorers Loveridge.
Love. Proc. Zool. Soc. 1922, p. 313.
Since my description of the type a second specimen has been
taken in the same locality by my collector.
Male. Snout to vent 350mm. Tail 70mm. Mid-body scales
17; ventrals 153; caudals 40; upper labials 8. This specimen
agrees with the type in all essential respects, excepting that the
3rd, 4th, and 5th upper labials enter the eye as against 4th and
5th in the type. It has also 5 more ventrals and 9 caudals less.
The following markings are also distinguishable. <A black
collar on nape or back of head touching posterior border of
parietals, but not reaching to ventrals ; it is very broadly V-shaped.
‘'welve or more black saddle-like markings, two scales deep, on
anterior third of body, separated from one another by two-scale
intervals which are approximately 2 mm. interspaces. White
stipplings on sides of anterior third of body sometimes falling on
outer edges of ventrals, which are grey and thus differ from the
colouring in the type, as they are very distinct from the blackish
dorsal colouring.
TARBOPILIS BEMIANNULATUS Sinith.
Blgr. Cat. Snakes, ii. 1896, p. 51.
Five specimens from Morogoro and Lumbo.
The Morogoro specimen is so much larger than the dimensions
given in the Catalogue that I give it here—353 inches (735+ 159).
A Lumbo specimen had only 198 ventruals.
It is curious how one may be in a locality for many months
without meeting with a particular species of snake, and then
several turn up within a few days, followed by a long period
when none are met with. Such was the case with this species at
Lumbo, where I caught two females in one evening and a third a
few days later, the circumstances of capture being as follows :—
_ At 7 p.m. I was hastily summoned to the Mess, where it appears
that the adjutant was about to take his seat when the snake was
noticed entwined in the wicker back of the chair. After putting
this Tiger Snake into a bag [returned to my tent, and within an
hour was summoned to capture a second, which was slowly moving
across the are of illumination cast from a lighted tent. Both
were females, and there was only a millimetre difference in. their
lengths (2 feet 6 inches) ; they were taken about 20C-300 yards
apart.
882 MR. A. LOVERIDGE: NOTES ON
The larger had ten eggs in the ovary, the smaller nine. Both
stomachs were empty of food, but the one was afilicted with
immature and, therefore, indeterminate worms of the genus
Physaloptera, as was also a specimen taken a few days later.
LEPTODIRA HOTAMBEIA Laur.
Bler. Cat. Snakes, iii. 1896, p. 89.
Two specimens from Garagua and Mwanza.
The latter had 20 and not 19 scale-rows at mid-body. Instead
of the white frecklings usual in Kast African Herald Snakes it
had transverse wavy white lines, one-scale wide, all along the
back, which gave it ‘a’ very Causus-like appearance. When
annoyed these Herald Snakes flatten their heads till the white
lips can be seen from above; the body is also depressed and flung
about spasmodically, giving the reptile a very viperish aspect.
I have always found them vicious and ready to bite on
the least provocation ; one specimen introduced into a vivarium
where there were already a couple of larger snakes (/’. sibilans
and 2. oryrhynchus) bit them both, but neither appeared to sufler
any ill effects.
CUAMZTORTUS AULIOUS Giinth.
Blgr. Cat. Snakes, iii, 1896, p. 98.
T have only come across three specimens of this scarce snake in
the past eight years: one was much battered in the roadway at
Mwomero, another was found, without data, in a bottle in a
German house at Morogoro in 1916, and recently (26. iv. 21)
a young one at Chanzuru near Kilosa which had the remains of
a gecko (apparently Lygodactylus sp.) in its stomach.
AMPLORIIINUS NOTOTANIA Giinth.
Blgr. Cat. Snakes, ii, 1896, p. 125.
Two specimens. A young male from Kilosa, and a female
from Mkata River. The former was crossing open ground, and
the latter was twisted into the side of a grass hut, about 6 feet
from the ground. In its ovary were two eggs, 12 mm. long
(27. viii. 21). In its stomach the remains of a gecko (Lygo-
dactylus pictwratus), and it was apparently stalking another
which was close to it at time of capture.
TRIMERORHINUS TRITENIATUS Giinth.
Blgr, Cat. Snakes, iii. 1896, p, 139.
_ Two specimens from Arusha and Nairobi. I take this oppor-
tunity of correcting an error of mine when I referred these
specimens to Psammophis brevirestris* Peters. The Nairobi
exainple was taken from beneath a latrine seat!
~* Journal E. A. & U. N. H. Soc. No. 18, 1918, p. 320.
EAST AFRICAN SNAKES. 883
A fortnight after its capture it attempted to swallow a dead
female Mabuia striata which was put into the cage. ‘The skink
was much too large for it, but the snake persevered for thirty
minutes, but could not get past the front legs; it finally disgorged
the whole. The Ar usha specimen had a skink in its stomach
when taken. ‘This snake is extremely gentle.
‘RHAMPHIOPHIS OXYRHYNCHUS Reinh.
Blgr. Cat. Snakes, iii. 1896, p. 146.
Hight specimens, Dar es Salaam, Chanzurn, Kilosa, Tindiga,
Pwaga, Dodoma, Luguo, and Mwanza District (1920-22), as
against twenty-six from Morogoro and Jumbo (1915-18).
Several were dug out of gerbil (7’. swaythlingi) holes and one
was found occupying the same hole as a mungoose (//. ivorit).
The largest male (whose tail-tip was missing) measured
567 inches (1014 + 425), and largest female 5 feet (1640 +
469), nearly six inches longer than the maximum measurement
given in the Catalogue. Half-a-dozen specimens had more
than 110 subcaudals, the highest number being 116. As might
he expected, a specimen from the sandy thornbush country of
Dodoma was very pale sand-colour.
At Kilosa these snakes were pairing during December in 1920.
Three very large ones were seen many times in the grass during
November, the rains having commenced on the 1st, but it was not
till the 4th of December that I surprised a pair in coitwu; one of
these I caught, and the other the following day. It was about
5 feet long. Ten eggs measuring 40 x 20 mm. were laid by one
of this pair in the vivarium between 1-7. viii. 1921, so that the
period of gestation would be about three months. Hight eggs
(15x 8 mm.) in a Dodoma specimen taken 11. vii.21. I have
already recorded: 13 eggs laid in October by a half-grown female
of this species.
In diet they are omnivorous; a small specimen endeavoured to
swallow a large rat, but could not get past the front Jegs and was
forced to disgorge (12. iii. 21), but a large specimen took a young
rat (15. 6. 21), and a wild specimen disgorged a shrew (Crocidura
Jjlavescens) on being caught. The natives savy that a tame guinea-
fowl chicken was taken by the largest of the three snakes which
were seen in September. The chicken was heard calling in the
grass, but as the boys were afraid to go into the grass, it is a |
matter of supposition. In captivity I feed them on M/abuia varia,
which they take well. It might be of interest to herpetologists
to know that a snake should not be fed on recently chloroformed
food. Wishing to pack some of these snakes to go to Europe
I chloroformed some geckos and skinks and introduced them into
the mouths of the snakes, which swallowed them, but a few
moments later showed signs of poisoning, one rolling on its back.
The bite of this species would not appear to be dangerous to
human beings, as a small native boy who caught a 4-foot snake
884 MR. A. LOVERIDGE: NOTES ON
was much bitten on the hands (which bled freely), but suffered
no ill effects. After the first excitement, incidental to capture,
they settle down to a life of confinement and are the gentlest of
snakes.
I found a young specimen in the stomach of a hawk (Butastur
rufipennis) at Morogoro on 31,1. 18.
Parasitic worms (Jilaria sp.) were found encysted in the skin.
Ticks (Aponomma leve Linn.) were found on a Kilosa snake
(20. v. 23).
PsAMMOPHIS SUBTENIATUS Peters.
Blgr. Cat. Snakes, iii. 1896, p. 160.
The Stripe-bellied Sand Snake called Sangaraza in Kiswahili
is so widely distributed that I have not troubled to catch many
this tour, as forty-six were collected in 1915-19; about a score
were taken 1920-22 from many fresh localities near Dakawa,
Mkindo Mkata Stn., Ilonga, Kimamba, Chanzuru, Kilosa, Itende,
Kidoti, Suna, Mtali’s, Sekenke Rd., 20 miles from Mkalama,
Usshora, Wembere, ‘labora, Sanga, Lalago, Sagayo, Nyambita,
and Lumbo.
This snake is an expert climber; one flashed across my path
and was 20 feet up in the topmost twigs of a stunted tree ina
moment. My attention was drawn to another five feet up ina
maiombo bush, but so well did it blend with its surroundings that
T could not distinguish it until it moved, though I was within
ten feet of it allthe time. I caught several in the thatches of
native huts, where they had gone in search of skinks I believe.
My largest male measured 49? inches (861 +4399) and female
453 inches (775+ 383). Both from Morogoro.
Whilst the colouring in specimens from one locality seems
constant, a great deal of variation occurs between examples from
different localities. ‘he characteristic black ventral lines were
indistinguishable, except with the closest examination when
traces could be seen, in the Kidete specimen collected by Mr.
©. F. M. Swynnerton. Four snakes from Mtali’s had pure white
instead of deep yellow bellies. At Suna. the coloration is adapted
to the sandy thornbush in which they live, and which seems to
influence the coloration of its typical mammals and, to a less
extent, birds.
This species will eat mammals, birds, snakes, and lizards.
A specimen taken at Frere Town after a hard chase was placed
in an ordinary small biscuit tin, about 8 x5 inches in size. The
following day a native brought me a Warbler (Prinia mystacea)
with its head almost knocked off, this I dropped into the tin,
and on opening it the following day found a bulge in the snake
and no visible bird, This freshly-caught diurnal snake hail eaten
a damaged dead bird in the darkness of a small tin on the day
succeeding capture, ‘here is no saying what a snake will do,
The following note-was made at Kimamba on 15. viii. 21:
EAST AFRICAN SNAKES. - 885
“ Between 5.30 p.m. and sundown I saw a Stripe-bellied Sand
Snake lying in the roadway; three times I disturbed it and it
returned each time. I noticed that a number of the Small
Weavers (Lagonosticta sp.) were picking up grass seeds and hopping
about unconcernedly close to it. I think one might assume that
the snake was lying there with the object of securing a bird.”
I have already mentioned under B. lineatus how one of this
species was found swallowing a young House Snake at labora.
Their commonest food, however, is the striped skink, and this
they very soon dispose of, as the following timing will show :—
12.44.50. Snake seized skink,
12.47.10. Head of skink was in throat.
12.48. Hind-legs enveloped.
12.48.10. ‘Tail disappears.
There is usually a pause for rest after the body has passed into
the gullet, ond the tail of the meal sticks out of the snake’s mouth
as if it had been having an after-dinner cigar!
At Frere Town (1. vii. 19) my attention was drawn to one of
these snakes which appeared to be playing. When first seen it
was stretched out at full length excepting that its head was
turned round in the direction of its tail. 1 then passed its head
beneath its body, then over the back, then beneath the body
again, and so on, traversing its own length towards the tail.
This was not done hurriedly, but in dallying fashion, with
oceasional withdrawing of the head. Unfortunately, a native
running up at this moment disturbed it, so that I was unable to
see the end of the performance. =
At Kilosa a captive specimen was seen to rub first one side of
its mouth against its side, then the other side of the mouth
against the opposite side, repeating the operation a score of times
as it slowly worked along its own length to the tail. I feel sure
this caressing movement was only play. Having reached the tail
it moved slowly away in the grass (Kilosa, 2. v. 23). This snake,
nearly 3 feet in length, was killed and three-quarters eaten by a.
baby lemur (Galago panganiensis) occupying the same large roomy
cage (Kilosa, 4. v. 23).
At Lumbo camp I often wondered what chance a snake would
have of being in the camp without my being informed by one of
the hundreds of natives employed about the place. With the
object of deciding this I released a Stripe-bellied Sand Snake
near my tent. This tent was situated beneath a mango and a
coco-nut palm, but had a cleared space of tifty yards of sand on
three ‘sides of it, on the fourth side were several buildings. The
snake was caught by me on a railway embankment half-a-mile
away, escaped, and was recaught by the reed fence enclosing the
tent. Next day I liberated it at Ga.m. At 8.30 a native came
running to say a snake was in his tent, which lay a little more
than fifty yards to the west of mine, and was ‘the first cover in
that direction after crossing the open sand. TI recaptured it and
Proc. Zoot. Soc.—1923, No. LVILI. 58
886 MR, A. LOVERIDGE: NOTES ON
a week later released it at lla.m. At 1.30 p.m. a European
corporal hurried up requesting me to catch a snake which was
under his bed; the marquee in which he slept was due north
from my tent. A week later I again released it at 6 a.m. and
at 8.45 a.m. found a man trying to head it off from entering a
patient’s marquee, almost due south from my tent and the first
cover in that direction. It had now tried in three different
directions, which was rather remarkable, and had been appre-
hended at the first tent in the direction taken. Twice more 1
liberated it, the last time at 10 p.m., to give it a night’s start, and
1 saw its tail disappear beneath my bed, As I heard nothing
more of it fora fortnight I thought it had at last won clear of
the camp, but on Oct. "10 it was “found at noon in the tent of
some A.N.M.C. boys, one of whom struck at it with a stick, so
injuring it that 1 killed it and threw it into a case of Lesser
Mungoose (//elogale ivorii Thos.), one of which seized it, first
erune hell its head, and then swallowed it whole. This sripilee had
a portion of its tail missing, so it was unmistakable. 1 concluded
from the results of this little experiment that [ made a pretty
exhaustive snake survey of the area covered by the camp when I
collected sixteen species in it during five months.
Many kinds of worms were found in the stomachs of various
specimens, including two new species in a Mombasa snake, viz.,
Oochoristica crassiceps Baylis, and Ophidascaris mombassica Baylis.
Physaloptera affinis Gedoelst, from a Kilosa specimen. Lhysalo-
ptera sp. and Ascaris sp. from Mombasa specimens. | am of the
opinion that these worms may cause the death of the host, for
in two instances the snake turned over and over and died. In
the case of the Mombasa snake the feet and claws of the Prinia
it had eaten just a month before were still in its stomach.
PSAMMOPHIS SIBILANS Linn.
Blgr. Cat. Snakes, iii. 1896, p. 161.
Nine specimens from Chanzuru, Kilosa, Wembere, and Lumbo
Lunges male 583 inches (1120 +4362) and largest female
593 inches (1100+ 404), but the tail tip is missing. The tail is
proportionately longer in the females of this species. Both these
records are of Morogoro specimens, being the best of seventeen
specimens collected 1917-18.
This snake feeds upon both mammals and reptiles, a very
large mouse being taken from the stomach of one snake and
several records of geckos (HZ. mabowia) being taken by them.
The eagle in whose gullet a full-grown specimen was found
has since been identified as the Black-breasted Harrier Eagle
(Circaétus pectoralis Smith). I have since taken a younger snake
in the stomach of another species of hawk.
At Lumbo (19. viii. 18) some natives killed a fine Cobra
(WV. nigricollis) in the act of swallowing a large Hissing Sand
Snake. Thecobra measured 503 inches and had alre eady swallowed
EAST AFRICAN SNAKES. 887
28 inches of the sand snake whose body measured 3323 inches;
unfortunately, in the excitement of killing the cobra the greater
part of the tail of the sand snake had been cut off, but other
specimens of this body length had tails of between 12 and
13 inches, so that a 504-inch snake was engaged in swallowing :
46 inch snake and would doubtless have succeeded.
From one of these Lumbo snakes I removed three worms
(Polydelphis quadricornis Wedl.) which measured 14mm. x 6 mm.
A Morogoro specimen, which was ailing and would not feed,
spued up a quantity of watery and slimy matter on the morning
of 9.iv.18 and died the same evening. There were no visible
parasites in the stomach, but one lobe of the liver appeared to be
full of cysts.
When crossing the Wembere Flats (9. xi.21) I came upon a
sand snake at 8 a.m. that was apparently dying of thirst. It
offered no resistance to being picked up and died shortly afterwards.
The stomach was clean and empty. It had dusky ventral stripes
like the brighter ones of P. swbteniatus.
PSAMMOPHIS BISERIATUS Peters.
Blgr. Cat. Snakes, 111. 1896, p. 168.
Three specimens from Kahe, Ikikuyu, and Dodoma. This
species seems partial to sandy or rock-strewn desert country.
The Dodoma snake, though only measuring 12? inches, had a
skink (Lygosoma ferrandii) in its stomach, as had also the
Tkikuyu specimen.
PSAMMOPHIS ANGOLENSIS Bocage.
Blgr. Cat. Snakes, ili. 1896, p. 170.
Six specimens from Kilosa and Izikisia,and two which were
without locality in a bottle at Morogoro,at which place they were
probably taken. Three were taken on paths and one twisted into
the grass of a banda five feet from the ground. This last had
nine upper labials on the left lip, the right having the normal
number of eight.
‘TMELOTORNIS KIRTLANDIL Hallow.
Blgv. Cat. Snakes, ii. 1896, p. 185.
One specimen from Zengeragusu, also seen at Liunbo.
‘The largest Morogoro specimen measured 58} inches (919+
548); the Zengeragusu female is 533 (933+ 415).
The Morogoro specimen Jaid 8 eggs on 16.1.18. These
measured 27X15 mm. The yellowish parchment-like shells were
irregular in outline with many concavities. ‘They were laid
singly at intervals of five to ten minutes. ‘The Zengeragusu snake
laid two eggs on 24.xii.21 which were dry when found, they
then measured 38x15 and 34x14 mm. respectively. As the
08*
888 MR. A. LOVERIDGE: NOTES ON
snake escaped the following day it is probable she would have
Jaid more.
This is an essentially arboreal species and offers one of the
finest examples of cryptic colouring amongst Kast African snakes.
The long and slender body is vine-like in its proportions, and
coloured for the part. In Morogoro specimens the crown of the
head was leaf-green and not unlike a leaf in shape; the Zenge-
vagusu snake had the crown of its head the same colour as
the body however. The tongue is bright red with a black tip.
When annoyed or scared they inflate their throats vertically like
~a Boomslang does.
I havealready mentioned the swallowing of a 7'yphlops mucruso
by this Zengeragusu snake in captivity. Two months after its
escape it was killed in a tree only 200 yards from its cage and
then had a Chameleon dilepsis in its stomach.
DisPHoLipDuS TYPuUS Smith.
Blgr, Cat. Snakes, iii. 1896, p. 187.
wo specimens of the Boomslang or Ngole, as it is known in
Kiswahili, were taken at Kilosa and Lumbo respectively.
‘The largest male measured 56 inches (1027 + 393) and female
692 (1269+489), the latter being 258 mm. longer than the
maximum given in the Catalogue. ‘These two snakes wero
the finest of a series of eleven collected at Morogoro. Of the
two snakes under consideration, the Kilosa one was bright green
and the Lumbo snake brown.
Tn captivity they ate sunbird, wagtail, weavers (Ploceus, Uregin-
thus, Lagonosticta), warbler, bulbul, and swallow. The Iilosa
snake when shot had the remains of a black swallow (Psalido-
procneé sp.) in its stomach.
The Lumbo Boomslang, 683 inches in length, had a chameleon
(C. dilepsis) in its stomach. At Morogoro a Brown Boomslang
fell out of a tree in the avenue, together with a chameleon.
A green Boomslang and a chameleon of mine escaped, and later
T recaptured the snake close by with what was probably the same
chameleon in its mouth. A few days later a second brown
Boomslang fell out of the same trees with a chameleon, All
these incidents occurred within a mouth of each other, showing
that though the Boomslang may like a chameleon diet it has
considerable difficulty in mastering its prey, seeing that no fewer
than three fell out of trees while attempting to do so.
I therefore introduced a chameleon into the cage of a very
large and black Boomslang. The snake immediately approached
the chameleon, sliding silently towards it with raised head; the
chameleon thereupon raised its occipital flaps, inflated its throat,
and swayed about from side to side, suddenly lunging forwards
with widely gaping mouth and uttering a hiss. I then removed
the chameleon from the cage, but fed several snakes on cham-
eleons later when pushed for other food.
EAST AFRICAN SNAKES. 889
CALAMELAPS POLYLEPIS Bocage.
Blgrv. Cat. Snakes, 111. 1896, p. 246.
Two specimens, one dug from a termite mound at Lumbo, the
other found in a bottle at Morogoro without date, but as all
the other material in the bottle was local I have little doubt that
it was collected in Tanganyika Territory. This species is only
known from Nyasaland and Angola before.
Lumbo male measured 17% inches (405-447). Scales 19, 176,
30,6. The maximum number of caudals given in the Catalogue
is 27.
RIWNOCALAMUS MELEAGRIS Sternf,
Sternf. Mitt. Zool. Mus. Berl. iv. 1908, p. 244.
Found on a path at Gonja (29. v. 16), measures 113 (269 + 25) ;
originally described from Lamu Is., K.C. I believe the identifi-
cation to be correct, though the number of ventral scutes cannot be
determined as the snake was stepped on and slightly damaged.
It has 15 scales at mid-body, 28 subcaudals, 7 labials, and was pre-
sented to the E. A. & U. Nat. Hist. Society at Nairobi, in whose
museum it now Is.
APARALLACTUS JACKSONII Giinth.
Blgr. Cat. Snakes, iii. 1896, p. 256.
The longest of five specimens collected at Longido West
measured 10% inches (228+ 48).
APARALLACTUS WERNERE Blgr,
Blgr. Cat. Snakes, iii. 1896, p. 257.
Single individual from Bagilo, Uluguru Mts., measured 104
inches (203+ 45).
APARALLACTUS CAPENSIS Smith.
Blgr. Cat. Snakes, iii, 1896, p. 259,
Seventeen specimens collected at Luimbo about the roots of
grass or shrubs, or on the surface of the sandy soil in the early
morning. The largest of this good series was 47 mm., less than
the maximum given in the Catalogue.
Earlier in this paper I have referred to a worm snake ‘oatine
eaten by this species in confinement,
ELAPECHIS GUENTHERI Bocage.
Blgr. Cat. Snakes, 11. 1896, p. 359.
One female from Nairobi (21. viii. 19) measuring 232 inches
(545 + 43), in whose stomach were four lizard’s eggs measuring
8x4mm. Is it possible that the snake had swallowed a pr egnant:
lizard and that the gastric juice had not acted on the covering of
the eggs ?
890 , MR. A. LOVERTDGE: NOTES ON
EXLAprEcHis NIGER Giinth.
Blgr. Cat. Snakes, i. 1896, p. 359.
A single specimen from an ant hill at Lumbo; another was.
killed but badly smashed in the same hill. I also encountered a
third specimen in November 1920 just before dusk on the Uasin
Gishu Plateau near the Burnt Forest. TJ held it down with my
cycle pump till IT had had a good look at it. As I had lost my
way I had other things to think of than its preservation, and so.
released it. Lumbo female 204 inches (488 + 32) with 142 ventral
scutes.
NAIA nNigricoLtiis Reinh.
Blgr. Cat. Snakes, iii. 1896, p. 387.
Twelve specimens of the Black-necked Cobra, known as Kigau
in Kikami:and Pera in Kiswahili (not Pili as given in sundry
dictionaries, which is the Puff Adder), were met w ith at Morogoro,
Rudewa, Kilosa, Mtali’s, Tabora, Mwanza Dist., Frere Many
Nairobi, and Lumbo.
The Lumbo specimens are referable to Peters’ var. mossambica,
Nairobi ones to forma typica. Mtali snake wasall black with no.
throat markings. Mwanza Dist. example was black above,
mottled black and white beneath, no red on throat. Tabora
specimen had the throat red- banded, being young it was plumbeous
above. I came upon it as I was clambering over a kopje, and it
came straight for me for the best part of a yard with hood spread,
spat at me, then turned into a crevice, which was the reason it.
approached, “ home” being between us. It struck me that the
red and black bands on the throat are obviously “ warning
colours.”
It is curious that of a score of specimens collected all have been
females hitherto ; one had 180 and another had 181 ventrals, two
had 51 and two 53 subcaudals.
During September 1920 an Indian juggler who had a captive
cobra brought me ten eggs measuring 35 x17 mm. recently laid
by it. The eggs are usually deposited in old termite hills.
Mammals, birds, reptiles, and amphibia are alike included in
the dietary of this species, and no doubt this fact explains its:
wide distribution and numbers. On arriving at Frere Town
(17. v.18) I was informed by my host that he had been much
plagued bya cobra in his fowl-house. During the three previous
nights it had killed six pigeons, two pigeous, and one pigeon.
respectively. At seven o’clock that evening I received a message
that the cobra was now in the fowl-house coiled : among therafters
about twelve feet from the ground. On arrival I saw a capture
was impossible and sent back fora ‘410 gun, from which I delivered
a charge of No. 10 shot. The concussion in the small iron
building made such a dust that we had to retire: a flop, and down
fell the cobra at our feet still writhing and striking this way and
EAST AFRICAN SNAKES. 891
that. It measured 644 inches, and in its stomach was a young
pigeon, whilst two pigeon’s eggs were in its gullet.
Twelve days later, just after dark, there was another great
outcry amongst the poultry, and my host running out with a
stick surprised a 663-inch cobra swallowing a chicken, taken
thus at a disadvantage it was easily dispatched. It was of course
stated that it was the mate of the first, come in search of its
fellow, but unfortunately for this popular belief both were
females.
At Morogoro, a native brought me a chicken coop containing
one dead cobra, one fowl! ditto, and three chickens, one of which
was headless. A few days before, this snake was supposed to
have taken three chickens from the same coop. This time the
boys hearing cackling ran out and killed the snake, but not before
it had bitten the fowl and her brood. Opening the snake I found
one chicken in its stomach and a chicken’s head in its gullet, and
so surmise that it was in the act of swallowing it when struck by
the boys, who probably knocked off the chicken’s body in their
attempts to strike the snake’s head. Length 51 inches.
T have already spoken of a cobra killed whilst swallowing a
sand snake. I have known one to eat four toads (B. regularis)
in a fortnight, another three in one day, and a still more
remarkable case of gluttony occurred at Morogoro. I went out
with a lamp and put two toads in the snake’s cage, which was
occupied by a half-grown cobra only. It seized the first toad and
on my return was chasing the other round the cage with the first
in its mouth; it struck at it again and again, but of course
without effect. It then paused and swallowed very energetically.
When the first was disposed of the second was bitten in the
abdomen, held for half-a-minute, then released, but as it began to
hop it was seized by the hind leg, and for nearly twenty minutes
the cobra attempted to swallow it hind end first. At the end of
that time it took the head in its mouth and swallowed it with
ease. Precisely the same thing occurred with the third toad,
which speaks badly for the reputed intelligence of the cobra.
A fourth and fifth toad followed, but I did not stay to witness
their engorgement. Six days later this half-grown cobra had
resumed its normal proportions.
Its dietary leads it to frequent the haunts of man, where it is
frequently found in sheds, fowl-houses, rubbish heaps, and tents.
In the bush they prefer to take up their abode in termite heaps,
upon which they lie and bask in the morning sunshine.
I know of a family residing near Nairobi in which nearly
every member, and many of the employees about the farm, have
been spat at in the eyes at one time or another. I wish to
emphasise this point that the cobra deliberately aims at the face,
as only the other day I read in a journal that it was a matter of
accident when the venom reached the eyes. The lady of the
household referred to, on going to the fowl-house, where it was
none too light, saw something dark in one of the boxes, and
supposing it to be a fowl she bent over it and received a charge
892 Mn. A. LOVERIDGE: NOTES ON
of venom full in the face, the resultant shock and pain was so
severe that she sank down on her knees and called for help.
This cobra rarely bites, so the following ease, which occurred at
Kilosa on 26.v. 21, is of interest. A very intelligent native in
iny employ was returning home one evening at dusk (6.30 p.m.)
when he stubbed his foot against what he thought was a stick
lying across the road. Next moment with a short hiss the cobra
Jaunched at him and struck with both fangs just above the left
ankle—his feet being bare of course. ‘The snake withdrew
immediately and set off in the direction of the railway line; he
ran after it, and the snake rose and spread its hood; he looked
about for a stone, but his friends called to him to withdraw or he
would get bitten again. He rejoined his companions, and very
soon began to feel sick, so he went to a native (Wanyimwezi)
“doctor,” who first applied a ligature above the knee and then
made from nine to a dozen horizontal incisions above the site of
each fang-marls, 7.e. between the bite and the heart, and into
these rubbed some “ medicine.”
Ramazan was taken home by his friends, and on arrival ate
some mealie-meal (usual evening food), but threw it all up.
‘Every time he attempted to eat the folloving day he was unable
to keep anything down, and he said he felt successive waves of
venom come up from the leg as far as his throat and then recede
again, By the 28th he was well enough to return to work.
I met a weird old “snake charmer” in Mwanza Dist. who had
in his possession a 68-inch cobra, which he kept in a small bark
basket. he first time I saw him playing with it, the snake slid
out when he came to put it back in its box; this happened two or
three times, and he slapped it on the head, when, quick as thought,
it apparently attem»ted to spit in his fuce (and as he was stoop-
ing over the box his face was not two feet from the snake). I
was within four feet myself and remarked that no venom accom-
panied the open-jawed hiss, and he replied with a laugh that the
venom was finished. He elected to become a camp follower for a
small consideration. A few days after, when he was holding his
daily display—tying the cobra round his naked waist or wrapping
it two or three times round his neck and flinging its head over
his. shoulder, so that it struck his back with a resounding whack,
—my curiosity was so piqued that I bought it from, him and
chloroformed it. ‘Lhe poison teeth and parotids were intact !
Among the parasites of this snake were worms (Diaphano-
cephalus simus Daubeney 1923 and Polydelphis quudricornis
(Weadl.), also ticks. he tick (Aponomma exornatum) was taken
on a Nairobi cobra.
Nata ware Linn, | |
Blgr. Cat. Snakes, iii, 1896, p. 374.
_ Though this snake ranges from Arabia and the borders of the
Sahara northwards to Zululand, neither Tornier nor Sternfeld
give any definite localities in Tanganyika Territory for the
PAST AFRICAN SNAKES. 893
Egyptian Cobra, and the present specimen from Kilosa is only the
second which I have come across. It was shot by Capt. Turnley
beneath a rock in a dry ravine called Mbweni and is now in the
Game Dept. collection.
Male just under 6 feet (1475 +325). Scales 17, 108, 69, 7;
the second labial on the right side is divided.
The stomach was found to contain a mass of mammal fur and
a. piece of tree bark measuring 31 x 26 mm. obviously swallowed
with the food.
DeENDRASPIS ANGUSTICEPS Smith.
Bigr. Cat. Snakes, iii. 1896, p. 437.
Six specimens from Morogoro, Chanzuru, and Kilosa, and a
7-foot specimen killed in the passage of a house at Mombasa.
The largest male was 94 inches (18804 502) and largest female
was 974 inches (2332-4139), Morogoro. Both IGlosa snakes were
bright leaf-green and under 6 feet.
‘wo newly-hatched young with the umbilical cord still unhealed
were found on March 5th and 31st respectively, both at the
same spot. :
The one’ Kilosa specimen which was brought to me alive fed
with avidityon dead rats; in fact, I think being nervous of large
rats, mambas prefer them dead. As with my Morogoro examples,
it would not commence to feed if anyone was about, possibly
realising that it was at a disadvantage ; however, by returning
quietly | succeeded in watching it feeding.
VIPERIDS.
Causus rHomBEATUS Licht.
Blgrv. Cat. Snakes, iii. 1896, p. 467.
Three examples of the Rhombic Night Adder were taken in
Nairobi and the Ngong Forest. .
During one week one of these adders swallowed a largish frog,
three small toads, and nine very small toads. On being chloro-
formed a week later and its stomach examined, all were found to
have been completely digested except the feet of the frog.
Another snake was seen by me to take seven small toads one
after the other, each about the size of a thimble.
Causus resimus Peters.
Blgr. Cat. Snakes, iii. 1896, p. 468.
Two were taken at Luguo and Sagayo.
‘The Luguo specimen, though on reddish soil, was a beautiful
grass-green ; the Sagayd specimen, on the same kind of ground,
was the usual greyish olive. Both with the normal markings.
The former had 7 labials on the right lip and normal 6 on left ;
the latter 26 pairs of subcandals. jprtarti, oe
894 MR. A, LOVERIDGE: NOTES ON
Causus DEFILIPri Jan,
Blgr. Cat. Snakes, i. 1896, p. 469.
A single specimen from Bagilo, Uluguru Mtns.
The largest of 15 specimens collected at Morogoro measured
16% inches (2934382); this series shows that the number of
ventrals may range from 110 to 129.
CAUSUS LICUYENSTEINI Jan.
Blgv. Cat. Snakes, ii7, 1896, p. 470.
This specimen was one of nine collected on the Yala River by
Mr. H. J. Allen Turner and is now in the Nairobi Museum.
Typical specimens are olive-green in life and plumbeous or lead-
colour when preserved ; the specimen under consideration agrees
in every way with the published description of C. lichtensteini,
except in the unusual markings and number of ventrals; but
typical specimens from the same locality have also the same
number.
Head and body 200 mm.; tail 15 mm.; costals 15; ventrals
158; caudals 17; labials 6.
General colour grey (brilliant leaf-green in life like Chlorophis,
but with a velvety tone—Turner). Head black. White line
commencing at the first scale behind the last labial following the
outline of the head passing along the edge of thesupracculars just
above the eyes, and finishing on the corresponding scale on the
other side. Lower portion of all the upper labials and whole of the
3rd china-white. An inyerted V on nape, apex just reaching to
posterior border of parietals. Throat pure white with a black U
corresponding to outline of throatand reaching to oval margin on
the second labial only. Third and fourth eular scales (or ventrals)
black, 5th and 6th white, 7th, 8th, and 9th black, 10th and 11th
white, remaining ventrals grey with faint white transverse bars
on every 13th ox 14th scale. Black V-shaped markings along
whole of dorsal surface at intervals of seven to nine genltas ; nine
very distinct white Jateral marks corresponding to bars on the
belly, the 10th just above vent forms a complete white ring three
scales wide, white spot near tip of tail, white freckles on anterior
part of black.
‘Another specimen Mr. Turner tells me was quite black in life,
similar to its preserved colour ; a brown stripe from the posterior
border of the eye passes along the whole outer row of body scales ;
the only other marking is a transverse brown band four scales
wide across the back abovethe vent. Its formula is as follows :—
267 mm.; 26 mm.; 15.144 ..20.6.
Brris anrerans (Merr.).
Blgr. Cat. Snakes, iii. 1896, p. 493.
Twenty- one Puff Adders from Nairobi, Morogoro, Kilosa,
Kipera, Mpanira-kwa- Sagoi, Mpwapwa, Usshora, and Lumbo.
EAST AFRICAN SNAKES. 895.
Most were taken lying on paths, but some in clearing or burning
grass.
The largest males were 434 and 403 inches from Kilosa, the
largest females 414 and 39 inches from Morogoro, but none of
these exceeded Catalogue dimensions. It is interesting to note
that the skin of the largest when dried measured 52 in.
The coloration is extremely variable in East African Puff
Adders, which may be lemon-yellow, chocolate-brown, or brick-red,,
or intermediate between these three main types. A young reddish
Puff Adder cast its skin on Nov. 7th and appeared in a cream-
and-brown skin ; exactly three months later, 7.e. Feb. 9th, he
sloughed again. On 17. viii.21, I made a comparison of two.
young local snakes. A nut-brown one had a black spot on the
lateral portions of each 5th ventral scale approximately. The
reddish viper, on the other hand, had the whole of the under-
surface chequered with black like a Tessellated Snake.
The smaller of the two males whose measurements are given
above was most unusually coloured : the black V-shaped markings
had a lighter outer edge of old gold; posteriorly there were
rectangular patches of the same colour, Another snake from
Kipera which was driven out by fire from a woodland of maiombo:
bush, and which harmonised most wonderfully with the reddish
soil and fallen brown and yellow leaves, had the V-shaped
markings vermilion, black, and cream, the rest of the back being
brown.
Some small boys informed me that two Puff Adders, which are:
known to the Swahili as ** Moma” or “ Pilipili,” were mating in
the scrub not far from where I was. There was only the male
to be seen when I arrived, a very fine one, and it lay with hemi-
penes extruded perfectly motionless, nor did it move when
I quietly placed a forked stick on its neck, not till I picked it.
up did it commence to struggle. Pairing was taking place on
Aug. 20th. A female very heavy in young was killed on
13.1. 20.
The Mpanira Adder had swallowed a largish bird whose quills.
were undigested. A younger specimen had an orthopteran in
its stomach in addition to ayoung toad. Toads (Bufo regularis),
but more often rats (2. c. microdon), are their principal food ; so
many persons have kept and recorded the feeding habits of this.
species that there is no object in my doing so here.
Three notes on the effects of their bite may not be amiss.
A fine conditioned male Bushbuck was picked up at Kilosa on
26. 111.21. There was a single puncture on one haunch, and from
the fluid condition of the blood and general hemorrhage I should
certainly say that the buck had laid upon (?) and been bitten by
a Puff Adder that morning.
When in the bathroom, my wife saw a rat entering by a hole
which drains away the bath-water. When just clear of the hole, it
struggled violently as if to free itself, and she thought she saw a
snake's head and called to me that there was a snake there. ‘The
896 MR, A. LOVERIDGE: NOTES ON
rat, meanwhile, ran up to the window-sill and entered a blind hole ;
by the time I reached the room, the rat had jumped to the wash-
stand and was lying dead with blood flowing from its left hind
foot. As Puff Adders are very common here and Cobras rare,
1 aim fairly certain it was the former. This occurred about 3 p.m.,
and 1 laid the rat by the hole, when, sure enough, it disappeared
at dusk, as I supposed it would.
Just before dark one evening I was passing the cases containing
snakes when I saw that a 2-foot Puff Adder had forced its head
through a broken corner of the glass door. It could not get out
further as it was too fat, nor could it withdraw itself owing to
the triangular shape of the head. Taking hold of its neck with
forefinger and thumb, I eased the belly scales past the glass
with the left hand, then, shifting my grip to the head, was pressing
on the quadrate bones and was on the point of letting go W hen
the snake twisted round and drove its left fang down my thumb,
scoring it so that it at once began to bleed. I imagine no venom
was liberated, as I suffered no serious consequences, though all
precautions were immediately taken, my native helper lancing
and ligatuving my thumb, which was immersed in a solution of
permanganate so strong that it took all the skin off. I might
add that I put a vat into this Puff Addex’s cage the following
morning ; the snake bit it, and the rat died very promptly and
was duly eunllowed Whilst we were in camp at Lumbo a native
died in hospital from a Puff Adder’s bite.
A very large percentage of Puff Adders are infected with
nematode worms, which I believe at times are the cause of death,
thus: Kilosa, 28. vii.21—Young male adder found dead about
fifteen feet from the kitchen door. Its stomach and intestines
were very full of rat’s fur; in the esophagus were a large number
of immature ascarids which Dr. Baylis states are probably Poly-
delphis quadricornis (Wedl.).
Kilosa, 19, vii. 21—One of my Puff Adders not having fed for
four months and being obviously unwell, I killed it. Beneath
the skin it had four nodules or Haven Jena of flesh-like substance
about 3 inch long, 3 inch wide, and } or 2 inch thick; these united -
the skin with the spine so that the re ptile could not be skinned.
Another snake killed the same day had a large number of
minute nematodes in the wsophagus which Dr. Baylis states are
Diaphanocephalus sp.,and adds that the species is being described
from other material by Daubeney under the name of D. obliquus.
In another the viscera was teeming with P. guadricornis, and
there was also a sapeworm in the stomach.
Birts casontca Dum. & Bibr.
Blgr. Cat. Snakes, iii. 1896, p. 499.
My native collector shot a very fine specimen in the Usambara
Mtns. and preserved the skin. I examined a second specimen
from Kilwa, which is on the East Coast south of Dar es Salaain.
EASY AFRICAN SNAKES. 897
ATRACTASPIS ROSLERATA Giinth.
Bigr. Cat. Snakes, iii. 1896, p. 514.
Six specimens of the Snouted Burrowing Viper were collected
at Lumbo from August to October 1918, and a female at Kilosa
in April 1923.
All but one were females, and the largest of these measured
243 inches (614+8); a male had 21, not 23, scales round mid-body.
These snakes are surprisingly quick, and when going to pick
one up in a tent (it having been discovered by the moving of a
box) 16 struck at my thumb. As I saw the snake lunge I was
also quick in withdrawing and just felt a prick on the top of my
thumb, which at the time happened to be protected with some-
what horny skin; no venom was visible.
iia ‘
; se nO: ey Be e ioe) ; ‘ fs pet
NOLES ON EAST AFRICAN BIRDS. 899
44, Notes on Hast African Birds (chiefly nesting habits
and endo-parasites) collected 1920-1923. By ArrHur
Loveripg4, F.E.8., C.M.Z.8.
[Received August 14, 1923: Read November 6, 1923.]
The collection for which the birds mentioned in this paper
were shot consists of 1558 skins representing 57 of the 63
families of East African birds, 311 of 398 genera and 761 species.
Only 150 eggs were taken in the peviod covered by this paper.
As before *, a few notes on some of the European migrants
have been included, but with this exception simple records are
avoided, though many rarities such as Cossypha albigularis Rchw.,
Callene sharpei Shell., and Zurdinus stictiguia Rchw. were
collected.
Dr. Hartertt has described the following species and races
from the present material :—Cinnyris loveridges, Arizelocichla
neumanni, Hremomela jlaviventris' tardinata, T'urdus milanjensis
aubugure, Tioptilus stierlingi uluguru, and Saxicola torquata
promiscua ; and Dr. Neumann}, Anthreptes longuemarei neylectus.
My grateful thanks ave due to Dr. Hartert and Mr. Arthur
Goodson for going through a great deal of the collection and
making the necessary, determinations. To Dr. Baylis also for
much kindness shown in the identification of the parasitic worms
mentioned in the following pages. fs
The only localities outside Tanganyika Territory referred to are
Frere Town and Nairobi. As difficulty may be found in locating
some of the other places (English and not German spelling of
place-names is given), I have arranged them according to their
districts and as far as possible Saonn east to west and south to
north, z.e. working from the Central Railway.
Dar es Salaam District.—Day es Salaam, Pugu.
Morogoro District.—Bagilo, Morogoro, Mkindo River, Wami
River, Mkata River, Tindiga, Kimamba, Ilonga, Kipera,
Medlazinn Kilosa.
Dodoma Dosiian —Dodoma, TRilenmattindle, Mahaka, Mbonoa,
Ndogwe, Suna, Mbulu’s, Mdjengo’s.
Kondoa-Irangi Disnich __Mtali’s, Zengeragusu, Usshora, Ulugu.
Tabora District.—Tabora, Tzilsisin, TNaale, ‘Simbo. Tamibali:
Luguo, Wembere Flats.
Mwanza District—Mwadira, Shanwa, Sagayo.
I witnessed an interesting display of curiosity exhibited by
birds at the sight of tortoises. A cage containing tortoises was
# Loveridge, P.Z.S. 1922, pp. 837-862: “ Notes on East African Birds collected
1915-19.” ‘
+ Hartert, Bull. Brit. Ornith. Club, xlii. p. 49; xliii. p. 149; xliv. p. 6, 1923.
if Neumann, Sonderabdruck aus Or nithologische Mon. xxx. p.1 (1922),
900 MR. A. LOVERIDGE : NOTES ON
placed on the path one afternoon at Nairobi, and the tortoises
were trying to get out. Some forty birds collected, and evinced
the greatest interest, craning their necks this way and that and
uttering cries. Twice a sunbird hovered close to the cage the
better to observe. I noted ten species of birds through the
glasses, and they remained for half-an-hour, and only left because
disturbed ; therefore I had ample opportunity of watching them.
The species represented were :—
Nectarvinia hilimensis Shell.
Parus albiventris Shell.
Serinus icterus Vieill.
Serinus striolatus affinis Reichn.
Passer griseus suahelicus Neum.
Ploceus reichenowit Fisch.
Ploceus spekei Heugl.
Cossypha heuglini Harti.
Dioptrornis fischert Rehw.
Oolius striatus hikwyuwensis van Som.
NECYTARINIDSA,
CINNYRIS SENEGALENSIS INASSTIMATA Hartert.
A nest containing two fresh eggs of the East African Purple-
shouldered Red-breasted Black Sunbird was blown down from a
mango-tree in which it had been built (Kilosa, 18. vii. 21).
A nest which was being built last month in a rhododendron-
bush, within six feet of the verandah and seven feet from the
ground, held two perfectly fresh eggs of a very pale colour to-day.
Unfortunately in replacing it I eracked one. The bird, however,
continues to sit on the remaining egg. She leaves it a great
deal during the day, and on returning flies straight into the nest
from another bush thirty feet away; hitherto I have never seen
her perched on the home bush (Kilosa, 27. xii. 21).
Latterly I have seen the bird cling to the front of the nest
before entering. Within the past few days the egg has been
blown out of the nest ! (Kilosa, 1.1. 22).
A month after the egg was blown down, the nest itself suffered
the same fate. The bird is now sitting on her new nest fastened
to the tip of a branch of a rubber-tree and about twelve feet from
the ground (Kilosa, 12. ii. 22). A nest containing young is sus-
pended from the tip of a branch of a big tree (Kilosa, 15.1. 23).
A nest with two fresh eggs (Ilonga, 19. iv. 23).
Crynynis LovEntpert ITaytert.
During the month of May 1922 my native colleetor obtained
two adult males and two immature females of this species, hitherto.
only known from a single male collected on 24. vi.21. These
four specimens come from Bagilo, Uluguru Mtns., Tanganyika
EAST AFRICAN BIRDS. 901
Territory. Some adult females were afterwards obtained ; they
are very similar to the male. As the immature plumage has not
been described, I do so here.
Upperside olive- green, with slight bluish-grey iridescences
becoming so pronounced on head as to form a smoky-blue crown ;
tail-coverts yellowish-olive; wings blackish-brown; outer webs
of the primaries with very narrow, inner with wider olive-
yellow outer edges; lesser and median upper wing-coverts like
back, longest like primaries; sides of head like crown; throat
greyish anteriorly, greenish-yellow posteriorly and on forebreast ;
breast, abdomen, and under tail-coverts whitish-grey with yellow
wash.
Wing 54mm. in this specimen, which is being presented to
the Tring Museum; 56mm. in second specimen in my own
collection ; tail 38 mm.; outer tail-feathers 28 mm.; bill from
forehead 21 mm.; tarsus 20 mm.:
PARIDS.
ANTHOSCOPUS CAROLI subsp.
A nest of the Penduline Tit containing almost fully-fledged
young was found in a tree about 20-25 feet from the ground.
Tree of the maiombo type in scattered bush-country on a, hillside
(Kilosa, 27. xii. 20).
MovTAcILUIDS,
Morvracinua AGuimMP Dumont.
A nest of the African Pied Wagtail (in the thatch of a hut)
which held one egg on the 12th had three to-day (Simbo,
12. xi. 21).
A nest with three young several days old was found in a
thatch, the young being fed on. termites which were flighting
strongly after recent rain (Ndala, 15. xi. 21).
A bird was seen to leave nest in thatch containing one egg
which had a fairly advanced embryo in it (Tabora, 18. xi. 21).
Young ones were found in a nest in a thatch only eight feet
from the ground a few days ago and flew to-day (Kilosa, 29. vii. 22).
Yesterday being the first rainfall of the lesser rains, I examined
the nest referred to in the last note, and was surprised to find the
_ bird brooding three eggs in the old nest. The species is obviously
double-brooded. I was particularly pleased, as the young were.
taken by a native from the last sitting, and I returned them
amidst much clamour from the old birds, which show confidence:
by re-using the same nest (Kilosa, 18. xii. 22), Young standing
up in nest (31. xii. 22). Nest empty (4.1.23). Fresh nest built
alongside but four inches from the old one. Bird pulling grass.
from thatch to build with (15. ii. 23). Three eggs in nest (early
lii, 23). Grits
Proc. Zoo. Soc.—1923, No. LIX. 59
902 MR. A, LOVERIDGE: NOTES ON
FRINGILLIDA,
EMBERIZA FLAVIVENTRIS Steph.
A nest of the Yellow-breasted Bunting was found yesterday ; it
contained two eggs, and is situated in a maiombo-bush five feet
from the ground. ‘The third egg completing the clutch was laid
to-day. The eggs are very similar to those of the English Bull-
finch. ‘The nest is built of dry grass-stalks lined with very fine
roots or root-fibres; it is rather untidy and exposed. The bird
is extraordinarily wary, and will not approach the nest when.
anyone is in the vicinity (Kilosa, 24. xii. 20).
SERINUS SULPIHURATUS SHELLEYI Neum.
Three eggs were found at Kabare on 11.i.23. The nest was
made of fine rootlets and lined with down from plants. | Bird
shot.
PLOCEIDA,
UR2aGinvilus NIASSENSIS Rehw.
Two nests of the Nyassa Blue Waxbill, each with two eggs
(Kilosa, 5.11.21). Nest and two eggs (11. iv. 21). Nest and five
fresh eggs (30.v.21). Two nests near house contain young
(19. iii.21). Two more nests found; one the bird has not
finished building and is still carrying grass; the other bird is
sitting on a clutch of -eggs (25.11.22). The bird which was
building on the 25th has now four eggs. The nest, which is built
in a lime-tree six feet from the ground, merely rests on the
branches, and can be lifted off and replaced without disarranging
it; it is neatly lined with fowl feathers (5. iv. 22).
Casually looking out of the railway-carriage window at the
telegraph posts (which are girders set upright with a smooth
surface on the west and a concavity on the east), I was struck by
the number of posts carrying nests (presumably last year’s), and
from the time I started counting till we reached the station,
T had counted 47, which did not include the remains of old
nests. It interested me, as it showed the adaptability of the
species to modern conditions, for they had seized on this siteinsuch
numbers that two out of three posts were occupied despite the
abundant bush close by. Another species of Weaver (Ploceus sp.)
had gone one better in selecting a snake-free site, and a number
of its nests were attached to the wires midway between the posts,
though some were against them, but not more than half-a-dozen
nests of this kind were seen (Pugu, 17. vi. 22).
LLAGONOSTICTA SENEGALLA 2
A nest built over that of some wasps was in a small tree and
contained young (Kilosa, 13. xi. 22).
EAST AFRICAN BIRDS, 903
PYTELIA AFRA CINEREIGULA Cab.
A nest containing four fresh white eggs measuring
16 x 12mm. was found in a maiombo-bush about seven feet from
the ground; the bird had just commenced sitting (Kilosa,
25.111. 21).
DINEMELLIA BOIMI Rehw.
Giant Weavers were usually met with in parties numbering
three to six individuals, of which one would be in a tree (?0n
guard) and the others feeding on the ground. They were extra-
ordinarily difficult of approach. They were seen building their
huge nests of twigs at Usshora (27.xi.21), Ulugu (7. xi. 21),
Wembere (8.xi.21), Luguo (9. xi. 21), Tambali (10. xi. 21),
Simbo (12. xi. 21). I have not seen them elsewhere.
ANAPLECTES RUBRICEPS Sund.
A Red-headed Weaver flew from nest containing one fresh egg
measuring 20x 14inm. and of a uniform pale blue colour. The
nest was situated at the extremity of an acacia-thorn branch
overhanging the road, about ten feet from the ground. Some
rain had already fallen in the district (‘Tambali, 11. xi. 21),
A nest was found containing two very unequal-sized young in
it (Izikisia, 16. xi. 21). meus
QUELEA SANGUINIROSTRIS INTERMEDIA Rchw.
From 4.30 to 6.30 p.m. a most extraordinary flight of Southern
Masked Weaver-Finches took place; millions of birds must have
taken part, and I have never seen anything like it. They were
in-a series of huge flocks which had almost the appearance of
smoke as each thinned and thickened in the undulating flight.
‘The intervals between the flights were very small, so that there
was practically one continuous stream of birds following the
course of the river N.E. to S.W. to their sleeping places in the
reed-beds (?) of a large body of permanent water called “ nyanza”
by the local natives (Simbiti River, 13. x. 22). About 7 a.m. we
were fortunate in witnessing part of the return flight, but it was
‘a mere nothing compared with the evening flocks (14. x. 22).
CoRVIDA.
CorvULYTUR ALBICOLLIS Lath.
A pair of White-necked Ravens made a great outcry on my
approaching a rocky kopje where a nest was found under an over-
hanging rock at a great height. The young appeared to have
flown, but there was no sign of them (Mbulu’s, 11. x. 21).
A nest was found in an almost inaccessible situation under an
overhanging rock on some rocky hills. It contained four very
59*
904 MR. A. LOVERIDGE: NOTES ON
markedly different eggs and a newly-hatched young one (Mtali’s,
22.x. 21). ,
T visited the Jast-mentioned site a little earlier this year and
found five eggs; the bird had only just commenced to sit. My
collector obtained the eggs by fastening his fez to a withy and
fishing them ont one at a time, which he necomplished success-
fully though in a very precarious position himself all the time
(Mtali’s, 9. x. 22).
Corvus scaPuLAtTUS Daud.
The nest of a White-bellied Raven was found in a bussu-palm
about forty feet from the ground, the palm heing in open bush-
country. The nest, which was on a heavy base of twigs, had a
lesser foundation of bark cord commonly used, and discarded, by
porters. On this was a thick lining of goat’s hair and a few ends
of string. Jt contained four typical crow eggs measuring
41x28 mm., much smaller than the raven’s, which were
50x 33mm. (Izikisia, 16. xi, 21).
Nest at the top of a slender fir-like tree in the town contained
three young. Nest was fully sixty feet from the ground (Kilosa,
28. xi. 21).
Building in a baobab at Pooma (4.x. 22). Four newly-finished
nests in stunted trees growing from the rocks at Shanwa
(21. x. 22).
DICRURIDA.
Dicrurus AFER LUGUBRIS Hempr.
When collecting lepidoptera I caught a Black Drongo in my
net ; its stomach contained beetles (Frere Town, 29. vii. 20).
Flushed a bird from its nest built in the fork of a branch of a
low tree at a height of seven feet from the ground. ‘The nest -
was made of fibres and grasses fastened together with spider-web
but unlined. It held three perfectly fresh eggs, one of which
was misshaped with a protuberance at the lower pole. The
ground-colour of these eggs was slightly pinkish, upon which were
superimposed brownish-red blotches, particularly thick around
larger pole. I watched a male displaying a few days ago, rising
and diving before the female which was sitting on a tree (Tamballi,
Allo, i)
LANIIDE.
Lanius cananisi Hartert.
Nest built entirely of rootlets (coarser used for exterior, finer
for lining) measured 5x44 inches outside, 4x34 inside, and
13 inches in depth. Clatch consisted of three fresh eggs mea-
suring 26x19mm. Pale olive ground-colour, with blotches of
' purplish-brown and brown grouped around the larger pole; a few
scattered elsewhere (Tindiga, 26. i, 22).
EAST AFRICAN BIRDS. 905
LANIUS coL~uuRIO Linn.
A Iippoboseid (Olfersia sp.) was taken on a shrike at Kimamba,
3.1v.23. Worms (Milaria nodulosa Rud.) from a Dar es Salaam
specimen, 24.1.19.
HLARPoLustEs SENEGALUS ORIENTALIS Cab.
The Coastal Large Bush-Shrike. was nesting at Kilosa on
5.1.23. Full clutch of two eggs in a maiombo-tree at a height of
seven feet from the ground.
MALACONOTUS POLIOCEPHALUS BLANCHOTI Steph.
A nest of the Brown-breasted Giant Shrike with three eggs
was found by my collector (Kilosa, 16. xii. 21).
CAMPOPHAGIDS.
CoRACINA PECTORALIS Jard. & Selby.
A female White-bellied Grey Cuckoo-Shrike, which is the first
I have obtained in this district, had a nematode (Physaloptera sp.)
in its stomach (Kipera, 8.ix.22). Mr. Goodson notes that the
wing and general size is smaller and somewhat paler than in
typical West African C. pectoralis.
PYCNONOTID®,
PYCNONOTUS TRICOLOR MIcRUS Oberh.
A nest of the Kilimanjaro Yellow-vented Bulbul with two
eggs in lime-tree seven feet from the ground; bird sitting (Kilosa,
ol. xii. 22). Nest with three eggs in a thorn-bush six feet from
the ground (Kilosa, 7. i. 23).
I encountered some natives chewing the red berries of some
wild fruit which was very gummy and had a nasty dry taste and
which they called Onembo in Kinyaturu. I thought they were
eating this fruit and attempted to doso myself, but presently
learned that they were on their way to trap birds, and this con-
stituted their “ bird-lime,” for, when chewed, it becomes a very
thick gum. ‘This they smear on a fine thread stretched near the
edge of the water-holes, which in this arid district are few and
far between, and consequently much resorted to by birds, which
collect to bathe and drink.
_ In about two hours these men returned with some twenty
bulbuls and a weaver, all of which they had plucked, except for
the heads which they had inserted through a string worn below
the knee, the naked bodies depending. The result was quite
ornainental, but they were destined to be eaten shortly. ‘These
men said that the Dongerero (local name for this bulbul) got
stuck when shaking the water from their wings after bathing
(Ndogwe, 1. x. 22).
I was sitting beneath a tree when suddenly, with loud cries,
906 Mr. A. LOVERIDGE: NOTES ON
a very frightened bulbul fell at my feet and commenced to hop.
and scramble away across a patch of open ground to some rank
grass fifty feet away. I had to run to overtake it, and, when
caught, it was found to have its left foot and wing firmly in the
grasp of a very fine leaf-insect (Zabalius ophthalmicus Wall.)
some three inches in length. ‘The strong grip of the insect was
astonishing, and considerable difficulty was experienced in dis-
engaging it from the bulbul, which I then released (Madizini,
3. ili, 23).
Pycnonorus DODSONI subsp.
A nest of the Mombasa Yellow-vented Bulbul containing two
partly-incubated eggs was found on the mainland, Mombasa, on
27. xii. 20
dlibde x eo Ve
PHYLLASTREPHMUS CABANISI sUCcosUS Rchw.
A nest containing two eggs was taken at Nabare on 12.1. 23.
It was composed of a foundation of dry sedges and leaves and
lined with very coarse rootlets.
SYLVIIDA.
EREMOMELA FLAVIVENTRIS TARDINATA Hartert.
A nest measuring 60x 40mm. deep inside was composed of
fine fibres, grass, and raw cotton, lined with very fine grass and a
little cotton. It contained two white eggs finely speckled with
black and purple spots, especially thick around the larger poles.
They measured 15x 10°5 mm. My collector shot both parent
birds (the types) from the nest (Sagayo, 2. xi. 22).
OrsricoLa NANA Fischer.
The hen bird was shot from a nest of not very definite shape
loosely woven of dry grass, lined with finer grass and much down
of the Javan silk-cotton tree, or kapock as it is locally called.
The three eggs measured 14x11 mm., and were white finely
streaked and speckled all over with pale pink. ‘There were very
small embryos in the eggs (IXilosa, 17.1. 22).
‘ ACROCEPHALUS GRISELDIS Hartl.?
I have queried this identification only as regards the nesting
note, as the nesting bird was not shot, but appeared to me to be
the Lesser Great Reed-Warbler (A. griseldis), which I had already
collected at the spot on 12.ii.21, but the Great Reed-Warbler
(A, arundinaceus) and the Sedge- Warbler (4. schenobenus) were
collected in the same swamp on 3. i. and 22. iv. 21 respectively.
‘Two purple-blotched eggs in a nest suspended from four sedges
in a swamp. Bird sitting very close (3l.v.21). One egg
hatched (Iiloga, 1. vi. 21). ;
EAST AFRICAN BIRDS. . 907
HirnuNnNDINIDS.
RIPARIA FULIGULA RUFIGULA Fisch. & Rchw.
A nest of the Brown-throated Rock-Martin built under the
eaves of a house contained two very bad eggs on which the
bird was sitting on 17. ii.22. On 17. iii. 22 the nest was again
examined and a clutch of two fresh eggs found and taken. On
4 & 5 iv. 22 the birds had but one egg in the nest; this was left.
On 1. v.22 the bird was again sitting on three eggs measuring
19x8mm. White with brownish or reddish speckles over the
whole surface, with a marked tendency to grouping round the
larger pole (Kilosa, 1.v. 22). In iv.23 the nest was examined
several times and the bird found to be sitting on a single
infertile e388:
HIRUNDO sMItHit sMITHit Leach.
A Wire-tailed Swallow’s nest containing young under eaves
of my house (1. xu. 20). - After a night and morning of heavy
rain I found a newly-fledged Wire-tailed Swallow flapping wearily
on the ground. After a few hours’ drying in the house it was
able to fly away (13.111. 21). I notice a pair of these swallows
have neatly trimmed off the broken edge of a nest of A. puella
abyssinica which contained young in January and have laid two
eggs (27. iv. 22); this was the full clutch (28. iv. 22). Bird just
begun to sit on clutch of three eggs (31. iv. 23). Three nests each
with three young (IGlosa, 6. v. 23).
Of nests examined to- day one was ready for eggs, a second
held three fresh eggs, a third three hard-sat eggs, two nests held
newly-hatched young, and one three fledglings almost ready to fly
(Frere Town, 30. v. 23).
HIRUNDO PUELLA ABYSSINICA Gueér.,
Northern Stripe-breasted Swallows nesting in bath-room ; young:
fledged to-day (13.111. 21). Commenced building on verandah
(26. xi.21), Feathering nest; each of the three adjacent houses
has aiso a nest (26. xii. 21), I saw one swallow on entering nest
break off the entrance tunnel, which had buta poor base of attach-
ment in the enamelled ceiling-boards (5.1.22). As I was passing
beneath the nest three-quarters of an eggshell was ejected. I
therefore investigated two other nests, and found one to contain
incubated white eggs without markings, the other young (6.1. 22).
The first heavy rain falling since the catastrophe of the 6th
inst. set the swallows to work at repairing the damage. It is
interesting to see the patchiness of the building materials, both
grey and. red mud in alternating patches according to the source
from whence fetched (10.i. 22). ‘Another eggsheli was found:
beneath the nest mentioned on the 6th. ‘lhese swallows are’
having trouble with a third, which sometimes succeeds in Smee
their nest:(17. i. 22),
908 MR. A. LOVERIDGE: NOTES ON
A nestling found dead beneath this nest (18.1.22). The two
remaining young ones found dead beneath nest this afternoon.
They appear to have been dead some time; one had congealed
blood upon its back. I surmise that they were killed by the
swallow which entered the nest some days ago and caused such a
commotion, and they have been thrown out of the nest by their
parents (19. 1. 22).
About this time there was a fresh egg lying broken beneath
nest (1.ii. 22). The tunnel and anterior third of the nest have
been broken down, so I have removed the rest (3.xi. 22). I
revisited this house after an absence of some six months and
found that these persevering birds had rebuilt the nest, which had
also fallen down again and again to judge by the débris beneath.
Their efforts had at last been crowned with success, as I picked up
a fledgling twenty feet away. It rested quietly on my hand until
its parents came sweeping around with encouraging cries, where-
upon it flew for fifty yards with them flying around it all the
time (31. xii. 22).
There is another nest of this species in which, the tunnel
having been broken off, a pair of Swifts (C. affinis) built their
feather and straw nest on to that of the swallows. My attention
was drawn to this nest by a pair of sparrows hanging to the
outside and screaming! Beneath the nest was a freshly-broken
sparrow’s egg, and inside the nest was a hen swift with enlarged
ovules, ready to lay perhaps in a week’s time (5. v. 22). A month
or so later a swift’s eye was found broken beneath this nest.
Continual disputes between the sparrows and the swifts take
place.
A Northern Stripe-breasted Swallow was found sitting on two
eges (31. xii. 22). Another nest, which has been a long time
in building, hatched out young to-day (7.1.23). These young
when disturbed at night by a light make a noise not unlike that
of a Square-marked Toad, and most unbirdlike, Another nest ;
hatched eggshell found beneath (5. iii. 23).
CYPSELIDS.
APUS AFFINIS Gray. |
As many scores of Square-tailed White-rumped Swifts were
flying about the boma and in and out of their nests, I examined
dozen of the latter. They were all empty and, I believe, unlined
(Kilosa, 4. vii. 22). A broken egg lying beneath a nest may have
been there for a couple of months whilst I have been away.
Some natives drew my attention toa swift caught in the web of a
spider (ephila sp.) some three feet below the guttering and two
feet from the wall and fully thirty feet from the ground. ‘The
bird was perfectly helpless, spinning round and round in space.
On getting it down, its feet were found to be firmly bound to the
body by the viscid web, and it was so wrapped in webbing that I
thought it best to chloroform it. Male. ‘Testes small (Kilosa,
16. xii. 22),
EAST AFRICAN BIRDS. 909
Of thirteen occupied nests examined to-day, two had one fresh
egg apiece, another held two fresh eggs, whilst a fourth had two
hard-sat and a fifth three hard-sat eggs. The remaining eight
held young in all stages. ‘he parent birds all allowed themselves
to be lifted off their nests. Fully nine of the nests had the nest
of a Wire-tailed Swallow for a foundation. Many other nests
were empty (rere Town, 30. v. 23).
CAPRIMULGIDS.
CAPRIMULQUS EUROPAUS MERIDIONALIS Lartert.
A male Mediterranean Nightjar was shot at Kilosa on 4. ii. 21,
and a female at Suna, Singida, on 27. ii. 22.
CAPRIMULGUS FOSSEI MOSSAMBICUS Peters.
Two eggs of the Mozambique Nightjar were brought to me by
my collector at Kilosa, 26.xi.20. They measured 28 x 20mm.
The one was addled, whilst the other held a very small embryo.
CoRACIIDS.
CoRACIAS GARRULUS GARRULUS Linn.
A. female of the European Roller was shot at Kiloga on 1. ii. 21.
They were observed to be very common in the Tabora District
during November of the same year. Fairly common and feeding
on grasshoppers at Ilonga, iii. 23. Parasitic nematodes in stomach
were in too poor condition for definite determination, but were
possibly IZadjelia sp.
MEROPIDE.
MELITTOPHAGUS PUSILLUS MERIDIONALIS Sharpe.
Two newly-hatched young of the Little Southern Bee-cater
were in a hole in the side of an ant-bear burrow which was
occupied at the time by a pair of porcupines (Sagayo, 4. xi. 22).
MEROPS APIASTER APIASTER Linn.
Shot several European Bee-eaters, which have been numerous
here for the past week or two. Their long tail-feathers are just
beginning to show (27. xii.20). ‘These birds are still here; on
several occasions during the past fortnight I have thought that
they were assembling for departure. They are in excellent.
plumage (5.11.21). The bee-eaters are still abundant here
(Icilosa, 13 & 30. iii. 21). :
Many seen flying (Luguo, 10. xi. 21). On my return I heard
the bee-eaters almost daily last week (Kilosa, 26. xii.21). The
bee-eaters are very busy with the millions of bees at the rubber
blossoms (Kilosa, 6.1.22). I heard the bee-eaters about a,
fortnight ago for the first time this year (Kilosa, 1.ii. 23); still
here (Kilosa, 1. iii.23). I may add that df. nubicus, persicus,
and superciliosus are all found here quite commonly.
910 MR. A. LOVERIDGE: NOTES ON
BUCEROTIDA,
_ Loprocrros sJacksont O.-Grant.
The Wazigoor call Jackson’s Hornbill ‘‘ Kwenabe,” which.
may be specific, as they call Bycanistes * Hondohondo” and
Bucorvus ‘‘Mkinga.” They eat this Zophoceros. ‘wo males
were shot at 12 p.m. and 6 p.m., and their stomachs contained
large numbers of black ants, other insects, and seeds of fruit. A
female contained a stick insect, beetle elytra, seed of the
“ mziga tree,” and fruit of the ‘““mkongo.” <A third male had in
its stomach a stink-ant (Paltothyreus tarsatus?), a black carabid,
and both seeds and fruit like the female (Wami [iver camps,
27. vili.—2. ix. 21).
A LCEDINIDA.
? HALCYON LEUCOCEPHALUS CENTRALIS Neum. |
' A kingfisher left its burrow in a bank at the bottom of a
ravine. The nest-hole was just ten feet from the bottom of the
water-course, which is now dry. In the nest-hole were two
roundish white egys approximately 25 mm. diameter and on the
point of hatching. Two examples of this species were found
drowned in a water-butt! (Glosa, 18. xii, 20).
CucULIDA.
CENTROPUS SUPERCILIOSUS INTERMEDIUS van Som.
Caterpillars, grasshoppers and beetles, and two species of para-
sitic worms were found in. the stomach of a female Hackle-necked
Coucal (Nairobi, ix. 19).
CENTROPUS SUPERCILIOSUS LOANDA C. Grant.
Worms in a female were identified as Davainea sp. (Kilosa,
17. viii. 20). Another coucal was heavily infected with lice.
One of these birds was also found drowned in the same butt as
the kingfishers mentioned in a preceding paragraph.
CUCULUS CANORUS GULARIS Steph.
At 6 p.m.a Yellow-billed Grey Cuckoo was calling in the thorn-
bush near camp. Its note seemed to lack the rich ringing sound
it has in the spring-time at home, but was unmistakable. 1
answered it about twenty times, while my collector stalked and
shot it—a male in bright clean plumage with large testes.
(7x5mm.) he stomach contents, were too triturated to be
recognizable excepting for long caterpillar hairs (Zengeragusu,
1.xi,21). Many cuckoos flying (Luguo, 10. xi. 21). Two seen at
close quarters (Kipera, 22. ii, 23).
EAST AFRICAN BIRDS. : 91}
CAPITONIDSE.
LYBIUS ALBICAUDA ALBICAUDA Shell.
Two Black-billed White Barbet nestlings were found in a hole
in the banks of the (dry) Shimiyu River at Sagayo on 8. x1. 22.
PICIDA.
CAMPOTHERA spp.
My attention was attracted by the noise made by two young
woodpeckers in a knot-hole of a thorn-tree in open thorn-
bush country. The young were practically fledged and making
a hissing sound; the nest-hole was only two feet from the ground
(Mtali’s, 21. x. 21).
Birds of another species, called “ Kinente” in Kinyaturu, were
seen feeding their young in a nest-hole which they had made in
a branch some thirty feet from the ground. Both species had
red-headed males and this species a black-headed female (Mbonoa,
DON ix. 22)
CoLUMBIDS.
TRERON DELALANDEI GRANTI van Som.
A new worm (Ascaridia fasciata Baylis) was found in the
co-type specimen of the Coastal Green-tailed Green Pigeon shot
at Dar-es-Salaam, 7. xi. 18.
STREPTOPELIA SEMITORQUATA SEMITORQUATA Riipp.
A nest containing two eggs remarkably different in size. One
measured 3623 mm., the other 32x24 mm. Both slightly
incubated (Kilosa, 22. vi. 21).
PsItTACIDS.
AGAPORNIS PERSONATUS Rechw.
Four white eggs of the Yellow-breasted Love-Bird measuring
22x17 mm. were taken from a hole in a baobab-tree at- Mahaka,,.
Dodoma, on 17. iv. 22,
STRIGIDA.
Tyo ALBA AFFINIS Blyth.
A “nest” of the African Barn-Owl was found beneath the
galvanized sheets forming the roof of a house; the bird gained
admission through a small ventilating hole. Two fresh eggs
measuring 40x 31mm. were found on a huge accumulation of
pellets which were mostly disintegrated. Besides the very com-
mon local rats (22. rattus alexandrinus and R. couche microdon)
there were the remains of a shrew (Crocidura flavescens). Some
912 MR. A, LOVERIDGE:. NOTES ON
of the skulls were in very fine condition, beautifully cleaned
(20. v.21). The hen bird was transferred to an aviary, where she
laid an egg the following day, but instead of laying it in a big
box of pellets provided, she dropped it on the floor presumably,
for it was smashed (Kilosa, 21. v. 21).
A young owl, unable to fly, was picked up in the grass close to
this house the following year, on 31. vii. 22. Of fifteen parasites
from the stomach of this species Dr. Baylis writes that these
worms are Habronema sp., “‘ probably ‘ Spiroptera’ penithamata
Molin, although this is said to belong to a different genus from
Habronenca.”
BUBONIDA.
Orus LEUCcoTIs? Grant Kollibay.
‘Lhree White-faced Scops Owls, apparently birds of the year,
3 6 2, were flushed from a thorn-tree in fairly close thom-
serub. Hach had the remains of a single dark-coloured rat in its
stomach (Mtali’s, 19. x. 21).
GLAUCIDIUM PERLATUM Vieill.
A Pearly Owl had a large J. c. microdon in its stomach (Kilosa,
26. vii. 21).
GLAUCIDIUM CAPENSE SCHEFFLERI Neum.
A female with very enlarged ovules sitting in a thorn-tree in
bright sunshine. Its stomach contained a male gecko (Lygo-
dactylus pictwratus) and a grasshopper. This is the bird that
makes a peculiar bubbling note just before dusk (Wami River,
oe dx. 21)!
Buso Lacreus Temm.
IT was shown an empty ten-gallon petrol drum in which a
Milky Eagle-Owl had nested in the fork of a rubber-tree only
eight feet from the ground and fifty yards from the house. The
one end of the drum had been filled in with plaited grass to induce
bees to hive in it. On October Ist the Rev. R. Banks flushed an
owl from it, and found the nest contained two large white eggs.
To-day there was a broken shell on the ground showing faint
‘traces of incubation ; the other was missing—probably taken by
a native (Kilamatinde, 5.x. 21).
BuBO AFRICANUS AFRICANUS Temm.
A Lesser Grey Hagle-Owl banged against the mosquito gauze
which encloses the verandah. Was it trying to take some insect
which was coming to the light, and itself misjudged the distance ¢
I shot it, and on examining the stomach found it to contain
half-a-dozen beetle elytra and one cricket (Kilosa, 18.i. 21).
EAST AFRICAN BIRDS. - 913
FALCONIDA,
FALCO RUFICOLLIS Swains.
A male Red-necked Falcon shot at 8 a.m. was found to have
the remains of a mouse and the leg of a grasshopper in its stomach
(Izikisia, 16. xi. 21).
FALCO SUBBUTEO SUBBUTEO Linn.
Hobby Hawks are very common here just now, their coming
synchronizing with the arrival of the rains. A female shot.
to-day had its stomach very full of termites (Simbo, 14. xi. 21).
CERCHNEIS TINNUNCULUS 'TINNUNCULUS Linn.
The European Kestrelis very numerous here. In the stomach
of one were parasitic nematodes which Dr. Baylis says are.
referable to Acuaria (Dispharyna) sp., possibly A. (D.) recto-
vaginata (Molin) (Morogoro, 24. i. 18).
CERCHNEIS NAUMANNI Fleisch. |
A pair of Lesser Kestrels were killed with one shot. ‘lhe
stomach of the female was distended with black ants, whilst there
were very few in that of the male. 8 a.m. (Izikisia, 16. xi. 21).
ELANUS CERULEUS Desf.
My collector found a pair of Black-shouldered Kites building
at Tindiga in June 1920, but on his revisiting the place in July
found that the tree had been cut down by local natives with a
view to destroying the nest; two eggs were lying smashed upon
the ground. Lice (Philopterus sp.) were found on a specimen
shot at Nairobi.
MILVUS MIGRANS PARASITUS Daud.
Returning to camp long after dark, probably between 7 and
7.30 p.m., my attention was attracted by a huge flight of birds
which passed low over our heads with much noise and whistling
cries, and commenced settling in a baobab-tree some two hundred
yards behind me, ‘They were coming from an easterly or south-
easterly direction, and there must have been quite three hundred
of them. Having no idea what they were, I sighted two against
the sky and killed four with one shot. ‘The stomachs contained
(i.) a scarab and a great many small grasshoppers, (ii.) a single
grasshopper, (111.) two grasshoppers, (iv.) empty. I visited this.
tree again to see if it might be the nightly rendezvous of all the
local kites for roosting purposes, but there were none to be seen
(Sagayo, 27. x. 22).
A great many kites are about the camp, and to-day one was
picked up in an exhausted state with the skinned body of a mouse
(thrown out by the skinners) fixed to both upper and lower:
914 MR, A. LOVERIDGE: NOLES ON
mandibles. An examination of its claws showed that there was
but one (the third in each case) on each foot. The other toes
were stumps showing the metatarsal bones worn through, as if
the bird had frequently tried to seize things with them. The
loss of the claws was not of recent date. I kept the bird for
twelve hours, but it seemed unable to feed, and in so low a state
that I killed it and had the skin preserved (Sagayo, 31. x. 22).
IT was informed that one of these kites was building ina ss
tree at Dar es Salaam in November (Dar es Salaam, 3. xii. 2 2).
An Unstriped Grass-Rat (dA. a. newmanni) was found in the
stomach of a male, shot to-day (Mtali’s, 20. x. 21).
Two species of worms were found in the stomach of one kite,
viz. (i.) Idiogenes longicirrhosus (Fuhrm.) and (ii.) Choanotenia
infundibulum var. polyorchis Klavtocz (¢) (Morogoro, 23. i. 18).
THoreraA£Tus WAULBERGI Sund.
A single example of Wahlberg’s Brown Eagle was collected by
the boy, who stated that he shot it when feeding on insects
ou the ground. It was a male and its stomach found to be
distended with thousands of large yellow ants (Camponotus sp.,
I think) (Kilosa, 4. 11. 21).
In a great nest on the top of a high tree on the banks of the
(dry) Shimiyu River was found a downy nestling (31.x. 22),
‘This was removed on the 9th of November, and grew very slowly
though it fed well. On 25. xii. 22 I found that one of its legs
was broken, «nd not knit properly, owing to some carelessness
on the part of the native in whose charge it was during my
absence. A curious thing was that I never saw the parent bird
near the nest, even though I slept in a neighbouring tree one
nicht and visited it at the first streak of dawn. Another day I
posted my collector to watch, and he said that he saw the old
bird circling in the sky above, but it did not approach (Sagayo
and Kilosa),
SPIzATUS BELLICOSUS Daud.
An immature male of the Martial Eagle, measuring tw
feet nine inches from beak to tip of claws, and with a wing
expanse of six feet six inches, killed a vulture (WV. monarchus)
after a fight lasting half an hour which was witnessed by
Capt. Tor nley, who subsequently shot the eagle. On examining
its stomach [ found it to contain the foot of a “dove or pigeon and
its stomach, On opening this second stomach I found seeds and
a number of small white beads (Kilosa, 5, viii. 21).
Evuronmaiitus sertoczster Bp.
Just as I shot a Hartebeest this bird came sailing overhead
and alighted in a tree close by. Its stomach contained indeter-
minable rodent remains and a single nematode (Porrocecum
-depressum (Zed.)) (Kipera, 8..i1x, 22).
EAST AFRICAN BIRDS. :' 915
Burro aveun Riipp.
A male Augur Buzzard shot on a rock had a large black rat
(A. tenebrosus 2) j in its stomach (Mtali’s, 19. x. 21). Another male
shot in a tree had an Unstriped Grass-Rat (A. a. neumannt) in
its stomach and in its crop a lizard (Hremias spekii) (Ndala,
lexi 2D):
Two newly-fledged young were shown me by a native child
who was about to eat them. I sawa nest in a thorn-tree among
the rocks, but it was empty. The natives say that the buzzards
nest here every year (Mtali’s, 19.x.21). A nest was seen in a
tree leaning out over the edge of a rocky krantz. My collector
shot the hen bird and took the eggs, which were quite fresh and
measured 52x62mm. They were white with brown smudges
and reticulations, mostly at the lower pole, but also extending
towards the upper pole (Shanwa, 23. x. 22).
MELIERAX CANORUS METABATES Heuel.
An immature male Northern Chanting Hawk with two lizards
(27. spekii) in stomach (Ulugu, 7. xi. 21). Anadult male with a
pregnant lizard (Agama atricollis 2), grasshoppers,. and many
parasitic nematodes, which were not preserved, in its stomach
(Ndala, 15. xi. 21).
MELIERAX POLIOPTERUS Cab.
A nest containing a single nearly-fledged young of the White-
rumped Chanting Hawk was built in a tree growing. from a
fissure in the rocks. Its plumage was strikingly different from
the adult. The back plumage is nearly black; it has a central
gular streak, but the lower breast shows signs of barring like
the adult. When first seen it was henna fed on an agama
lizard by its parent, but it soon took to feeding itself on scraps of
meat cut up small (Shaniwa, 24.x.22). The stom a of another
specimen shot 2t Mwadira contained an agama umd grasshoppers
(19. x. 22).
Circaiirus FASCIOLATUS Gray.
The Banded Harrier-Eagle is, I believe, a somewhat scarce
bird; in the stomach of a male shot to-day there were remains of
a rat (2. c. microdon) and the scales of a lizard, apparently those
of Mabura striata (Mkindo. River, 8.ix. 21).
CIRCAETUS CINEREUS Vieill.
A female Brown Harrier-Kagle had snake scales in its stomach
(Mtali’s, 19. x. 21).
KAUPIFALCO MONOGRAMMICA Temm.
The stomach contents of a One-streaked Hawk, shot at 7 a.m.,
were a gerbil (Zaterona vicina ?) and a freshwater purplish crab,
916 MR, A, LOYERIDGE: NOTES ON
which (? the same one) I had encountered on the path at 4 p.m.
yesterday (Kilosa, 19. ii. 21). A male had the skull and fur of a
shrew (Crocidura flavescens) with many orthopteran remains and
a few parasitic nematodes (Kilosa, 26.iv.22). A very young
Spotted Wood-Snake (Philothamnus semivariegatus) and ginss-
hoppers comprised the stomach contents of another male
(Kimamba, 7. iv. 23). Yet another male shot at 5 p.m. had a
galeodes, a large locust, two praying mantids, and other insect.
remains. It had also two parasitic nematodes, identified by
Dr. Baylis as Physaloptera % acuticauda Molin (Kilosa, 6. v. 21).
Astur 2 POLYZONOIDES Smith.
A female with ovules somewhat enlarged. In stomach, the
fur and feet of a rodent that looked like Mus bella (Wami River,
9.ix. 21).
PoLyBoRIDES 'ryPIcus Smith.
Salimu shot a female Bare-faced Whistling Hawk in imma-
ture and interesting plumage. The primaries and two of the
secondaries in each wing were brown, and there were quite 2
number of light brown feathers among the inner wing-coverts,
The bare skin around the eye was light yellow, the base of the
bill between lores pink, the bill itself black. Eyes black. Legs
yellow. In its stomach was an adult lizard (Zonwrus tropido-
sternunr) (Kipera, 5. viii. 22).
Circus MAcRouRUS Gmel.
A Grey Harrier was infested with lice and in its stomach
were nemato les (Physaloptera ? alata Rud.) (Morogoro, 28.11. 18)..
VuULTURIDSA,
NECROSYRTES MONACHUs Temm.
A worm (Thelazia depressw Baylis, 1920) was taken from the
eye of a Brown Vulture.
TBIDID&.
THRESKIORNIS zrurorica Lath.
Just after sunset some fifty Sacred Ibises came flying across.
the plains to roost in a big tree overhanging from the rocks.
The tree is also occupied by a score or more Corvus scapulaius
and probably as many Milvus a. parasiticus (Shanwa, 20. x. 22).
HIAGEDASHIA HAGEDASH Lath.
My collector shot a female Great Glossy Ibis with a perfectly
formed and coloured egg in its oviduct. This egg measured
51x46 mm., and was slightly greenish-white in gronnd-colour,
spotted, blotched, or heavily smudged with brown, I told my
EAST AFRICAN BIRDS. 3 917?
boy to return and search the neighbourhood thoroughly for a
nest, and this he succeeded in finding in a tree growing from
a fissure in the rock-face about 20 ft. from the ground. It was,
very difficult of access, and was eventually reached by coming,
down the fissure in the’ rock-face, which only a native with bar a
feet could negotiate. He then removed his putties and, tying them’
together, lowered them to me. I tied a net on to these, and the
two eggs were safely lowered one by one. They measured 5d x 42,
and 61 x 42 mm. (Shanwa, 21. x, 22).
A male which was shot from a flock of three had the seed of
a rubber-tree and a peculiar looking dipterous larva in its
stomach (Iipera, 5. viii. 22).
PLATALEIDS. ;
PLATALEA LEUCORODIA LEUCORODIA Linn.
The European Spoonbill was shot at Kilosa on 4. iv. al, and its
stomach contained the head of a large dragonfly (Anaw sp.)
and the plastron parts of a big beetle (? Copris’ sp.) together
with a little green stuff. Its intestines were teeming with
tapeworms (Cyclorchida omalancristrota (Wedl.)).
PLATALEA ALBA Scop.
An African Spoonbill was shot ab Sagayo on 8. xi. 22 by oy)
collector. -
ARDEIDA.
Mrsoruoyx INTERMEDIUS BRACHYRHYNCHUS Brehm.
The stomach of a female Short-billed White Heron contained
a frog and no parasites (Kilosa, 28. vi. 21).
ARDEOLA RALLOIDES Scop.
The stomach of a Buff-backed Heron held insect remains and
three nematodes identified as Contraceccum mer ea (Rud. )
(Kilosa, 29. vi. 21). :
ARDEA PURPUREA PURPUREA Linn.
The stomach of a male Purple Heron contained a house-rat
(2. r. alewandrinus), several small fish, and a dragonfly with
black bands on its transparent wings. Parasitic nematodes were
Contraceecum microcephalun (Rud.) (Kilosa, 11. vii. 21).
ARDEA CINEREA CINEREA Linn.
A. male of the European Grey Heron was shot at Kilosa on
6. vil.21. A female at Sagayo on 7.xi.22. The latter was shot
at 2 a.m, when it was fishing in a pool of the Shimiyu River.
Proc. Zoou, Soc.—1923, No. LX. 60
918 MR. A. LOVERIDGE ; NOTES ON
- BurorreEs ATRICAPILLA Afzel.
Two eggs of the Green-backed Heron of a uniform pale green
colour and measuring 34x28 mm. were brought me from a
swamp by my collector. ‘They were blowable, but had under-
gone considerable incubation (ICilosa, 5. vii. 21).
‘In the stomach of a specimen were worms identified as
Hymenolepis sp. (near to H. wrilateralis (Rud.)); in the intestines
of the same bird was:another nematode—Contracecum micro-
cephalum (Rud.) (Morogoro, 15. i. 18).
Scoprips.
Scorus UMBRETTA BANNERMANNI Grant.
The East African Hammer-head is known to the Wazigoor as
“‘Mzeringee,” the Wahehe as ‘‘ Mgetu,” the Wanyimwezi as
“Mgenyi.” A cock bird, shot while flying overhead. this
morning, had a frog and a shrimp and a number of parasitic
nematodes (Contracwcum microcephalum (Rud.)) in its stomach
(Mkata River, 25, viii. 21).
A nest was found in the fork of a tree leaning over a lagoon
which was 200 yards long but rapidly drying up. The nest held
four young within a week of being fledged, and they throve in
captivity for six months, but died when transferred to Regent’s
Park. They would probably have died if left in the nest, as the
surrounding ground was swarming with soldier-ants intent on
destroying every living thing; many were already on the trunk
of the tree (Mkata, near Station, 14. ix. 21).
CIcONIID 4.
ABDIMIA ABDIMI Licht.
The Purple Storks always appear with the first rains, and are
therefore known to many tribes of natives as the ‘“ Rain-birds.”
They have a peculiar way of circling in the air like a column of
smoke, more particularly just before sunset. The stomach of a
male. examined contained centipede, grasshoppers, cockchafer,
and three other species of beetles (Simbo-Ndala Road, 14. xi. 21).
A great flock appeared at Sagayo camp during the first week in
November 1922.
E/pHirPloRHYNCHUS SENEGALENSIS Shaw.
A male shot at Tindiga measured 6 feet 4 inches from tip of
beak to tip of longest toe-claw and 8 feet 5 inches across the
wings; the two fleshy lappets depending from the base of the
lower bill are bright lemon-yellow, not “ bright crimson” as
stated by Stark and Sclater*. .
I also saw Saddle-billed Storks at Kilamatinde and Kipera, but
it is a scarce bird and goes singly or in pairs,
* Stark & Sclater, ‘Fauna of South Africa,’ vol. iv, p. 44, line 7.
EAST AFRICAN BIRDS. - 919
_ ANASTOMUS LAMELLIGERUs Temm. :
The stomachs of a pair of Open-billed Ibis shot here contained
some hard muscle-like lumps of matter, apparently from shell-
fish. No parasites were observed (Kilosa, 2. iv. 21).
ANATID,
SARKIDIORNIS MELANOTUS AFRICANUS Hartl.
’ Tho Knob-billed Goose is very common here. An immature
female shot to-day had down still mixed with its feathers. In
the stomach and crop there was a large quantity of maize (ICilosa,
30. v. 21).
CoLYMBID&.
PoDICEPS RUFICOLLIS CAPENSIS Salvad.
The following notes on the nidification of the Cape Grebe were
all made on a small swamp or accumulation of temporary water
which forms after the big rains and lasts about three months :—
Found a floating not containing two dirty-white eges
(14. iv. 21). Revisited the nest.at 5 p.m. on the 15th when
there were three eggs, and again to- -day when there were six.
The hen bird appeared flapping about in the water in an attempt
decoy us away. Not far away I found another nest with four
gs (18.iv. 21). The bird, which laid four eggs and whose nest
i abel laid a fifth egg yesterday on a few partly-submerged
reeds which it had collected together for the purpose some fifty
feet from the abandoned nest. To-day it laid its sixth egg in
the deserted nest of another bird. ‘he full clutch would appear
to be six eggs (20. iv. 21).
A new nest with two eggs (22. iv. 21); five in this nest, and one
of them partly incubated (25.iv. 21). A fourth nest found con-
taining two eggs (25. iv.21). The fourth nest held seven eggs,
some fresh, some incubated. Shot what I thought to be the hen
bird, but she had a pure-white egg in the oviduct measuring
36 x 24. mm. and two enlarged ovules in the ovary. The
“ colour” of this egg compared with the brown need: stained ones’
in the nest was very striking (3. v. 21). ik
Revisited the eae for the first time since 3.v.21. One
nest held two eggs. Two pairs of birds were swimming about
with their young; ‘their conduct was now very different ; pre-
viously they had only splashed and dived to decoy one away,
now they showed themselves boldly. From time to time one of
the birds would utter a sharp whistling note, and having thus
attracted attention, would dive with a splash ; after a few
moments the head would appear some distance off, and the cry
would be repeated to lead one away from the young. These
were entirely black, showing no rufous whatever; broken bits of
shell were left in the nest from whence they came. It was a
pretty spectacle to watch the old ones taking the young for rides
on their backs, themselves semi-submerged (30. v.21).
60*
920 MR. A. LOVERIDGE: NOTES ON
The nest, which held two eggs yesterday, had only one in it
to-day, the other having hatched out, as was evidenced by the
bits of shell left. The remaining egg had been carefully covered
with grass, as.is the custom with all but deserted eggs (31. v. 21).
This egg still in the nest ; another nest was found also with two
eggs, uncovered and obviously deserted, as they had never been
covered, being only stained on the lower surface. They were
very fresh however (Kilosa, 1. vi. 21).
‘'wo species of worms were found in the specimen shot, viz.
(1) Dicecocestus aspera (Mehlis) and (2) HWymenolepis sp.
RALLIDA.
GALLINULA CHLOROPUS BRACHYPTERA Brehm.
Three nests of the African Moorhen were found built in
clumps of grass or reeds in deep water. The first had two egps
on 22. yi. 21, the second two eggs on 6, viii. 21, the third five
eves on the same date, Like the Common Moorhen’s eggs, these
show considerable variation in size, »veraging about 41 x 30 mm. ;
the coloration is also identical with that of the Hnglish eggs
and, like them, varies, a good deal even in the same clutch,
(Kilosa, v, d.). |
GaALLINULA ? ANGULATA Sund.
A single egg of the Lesser African Moorhen, measuring
48x31 mm., pale in colour with fine frecklings more like a
coot’s egg (Kilosa, 22, yi. 21).
PHASIANIDA,
. NuMIDA CORONATA REICHENOWI O.-Grant.
Five eggs of Reichenow’s Helmeted Guinea-fowl measuring
48 x 36 mm. were taken at Suna in February 1922 by my native
collector. °
Numipa mirrata Pall.
The Coast Guinea-fowl is extraordinarily abundant between
Mkata and the Wami River and at many other places in the
Morogoro District. I have killed five with one shot, and a native
killed eight with two on the path just outside the house (Isilosa,
4. viii. 22). Lice (Goniodes nwmide Mjob.) were taken on a
Kilosa, bird on 11,111. 1921.
PARID2.
PHYLLOPEZUS AFRICANUS Gmel,
- Bits of quartz and grass seeds in the stomach of a male African
Jacana with large testes shot this day (IXilosa, 9, iv, 21).
EAST AFRICAN BIRDS. 921
Microrarra CAPENSIS Smith.
I shot a Pigmy Jacana from a flock of three seen on the
swamp; its stomach contained insects; ovules large as if the
breeding-time is near (Kilosa, 1. vi. 21).
CHARADRIIDEA,
Ruworri.us crnctus crnctus Heugl.
A. native brought me 2 Three-banded Courser cock which
he snared as it returned to an egg which he had found buried in
sand with only the top showing; the egg was well incubated.
Heads of soldier-ants were found in the stomach of a female
(Zengeragusu, 3. & 1. xi. 22),
BURUINUS CAPENSIS CAPENSIS Licht.
A young Cape Spotted Stone-Curlew in down was brought me
(Mdjengo’s, 24. x. 21).
STRUTHIONIDA,
STRUTHIO CAMELUS MASSAICUS Neum.
As we were making camp a hen ostrich was to be seen some
two hundred yards away casually feeding in the stubble of a
mealie-field. Later I met it a mile away and carefully stalked it
(without a gun). After I got within fifty yards my self-gratula-
tion melted away, and when I was within twenty yards I felt
sure the bird had some history. I enquired of an old man at a
neighbouring kraal, and he told me that it had been in the neigh-
bourhood for a year past, and was supposed to have strayed from
a mission at Itigi (twelve miles away) where some had been
reared. It lived all alone in the bush and came daily to the
shambas to feed or be fed. ‘ It will come if you call it,” said he,
and picking up an old maize cob from the ground, he held it out
and called as one might call chickens, when, sure enough, the bird
came running, but halted ten feet away on seeing that the object
proffered was not edible. The old man said it would feed from
one’s hand if matama was held out to it (Mbonoa, 29, ix. 22).
(ie:
ie
i sf
th te
ae
yee
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au ye i
P. Z. S. 1923, LOVERIDGE, PI}. J.
John Bale.Sons & Danielsson, UY
Norman C. Miller.
TESTUDO PROCTERA, sp.n.
(natural size of type which is not adult. )
he
ies,
Pie
LOVERIDGE,
og Sq VOLE,
iP.
John Bale.Sons & Danielsson, Lt?
sp.n
TESTUDO PROCTERA.,
NOTES ON EAST AFRICAN TORTOISES. 923
45. Notes on Hast African Tortoises collected 1921-1923,
with the description of a new species of Soft Land
Tortoise, By Arraur Loveripar, C.M.Z.S., FES.
(Plates I, & II.*) —
[Received August 14, 1923: Read November 6, 1923.]
The hundred tortoises dealt with in these notes were all taken
in Tanganyika Territory (formerly German East Africa) by my
native collectors; not more than half-a-dozen were found by
myself. ‘They represent three of the four families known to
inhabit Eastern Equatoria.
For the convenience of other herpetologists a key to the species
recorded from this region has been added to these notes. It is
largely adapted from keys in Dr. Boulenger’s ‘Catalogue of
Chelonia in the British Museum,’ 1889, with the recently
described species incorporated.
Taking them from Mast to.West, the localities mentioned in
the following pages are :—
Morogoro District.—Turiani,, Mkata River, Uliya,. Myombo,
Ruaha, Godegode.
Dodoma Dis ict.— Pwaga, Kidenge, Tieileuy oh Boden, Kis-
saki, Kilamatinde, Mbonoa, Singida, Mdjengo’s.
Kondoa-Irangi District —Mtali’s, Zengeragusu, Ulugu.
Tabora District—Tahbora, Simbo, Tambali, Luguo, Wembere.
Mwanza District. —Sanga, Sagayo.
My thanks are due to Miss Joan B, Procter, Yr. Z.8., for advice
and suggestions and the use of material in her charge, whilst
writing up my field-notes. I have named the new species of soft-
shelled land tortoise after her as some acknowledgment of her
careful researches into the morphology and relationships of the
strange group to which it belongs.
' Tamalso indebted to Sir John Bland-Sutton for his gift of X-ray
photos of the type which illustrate this paper and to Mr. Norman
C. Miller, F.1.8., for making the water-colour drawing which
forms the plate accompanying this article. Mr. ‘Miller was
staying with me at the time of the tortoise’s decease and spared
time from his work on the Orthoptera to make this sketch
for me.
Mr. Stanley Hirst has also kindly named rte few ticks found
upon the tortoises. It is interesting to note that the four
species of Testudinide, though. coming from three widely sepa-
rated localities, all bore the same tick, which I have never found
on any other reptiles or mammals.
*® Wor explanation of the Plates see page 933,
924 . MR. A. LOVERIDGE: NOTES ON
TERESTUDINIDS.
CINIXYS BELLIANA Gray.
Bler. Cat. Chelonians, 1889, p. 143.
More than fifteen specimens from Turiana, Mkata Ravens
Uliya, Myombo, Kilosa, Usikuyu, Pwaga, Godegode, and Simbo.
Bell’s Hinged Tortoise is very common at Kilosa, both on
the swampy grasslands of the lower ground and on paths in the
maiombo bush on the hillsides. They can live in water for days
without drowning, apparently resting on the surface without
effort. I was given one by an Indian who had dropped it down
a well two days before, where it was when I found it, none the
worse for its experience.
_ In the Kilosa sub-district, where the country is of the open
maiombo-bush type with rank grass growing between the bushes,
the shells are much deeper than in the rock-strewn or dense
thornbush country such as Pwaga, where the thornbush covers
many miles with a thick and impenetrable mat; rocks were
found in the valleys and scattered here and there upon the hill-
sides, but those I came across did not seem to afford much cover
for tortoises. The single Bell’s Tortoise taken at Pwaga was
very worn and depressed, as if it continually pushed its way
beneath boulders, The Ikikuyu country is of a similar type to
that at Pwaga, though there are more open, sandy, thornbush
stretches on the low ground; the single specimen from this place
was also depressed but not worn, the markings being distinct.
Those from Simbo—similar country to Ikikuyu though a hundred
miles or more west—were also worn and depressed, as has been
remarked on elsewhere.
Locality. Length. Breadth. Depth. Depth.
Kilosa...... 190 mm. 125mm. 90mm. 47:5 per cent.
Pwaga...... IG Guae OTe BYE he SACOM
Ikikuyu ... 150 ,, BY a BO) Bae Se aR
Simbo...... 148 =, 95 ,, De SoROlnes
PRUE Rt Tass DM, Dihits, Oe
My method of taking the depth was by placing the animal on
its side between two blocks. Miss Procter measured with
eallipers at a given point, and as a depression in the plastron is
frequent, our depths, and consequently depth per cent., are dif-
ferrent, though both ways serve equally well to emphasise the
depressed type of Cinixys inhabiting thornbush country.
The largest male (?) measured 190x125 x90 mm. deep, the
largest female 188 x 12080 mm. The smallest example was
taken at Uliya on 16.11.21, and measured 50x46x25 mm. It
is somewhat abnormal in that it has twelve instead of eleven
marginals on either side,
On two occasions I have found several score of Lesser Stink
Ants (Paltothyreus tarsatus) in the tortoise enclosure worrying
- BAST AFRICAN TORTOISES. ‘995
the reptiles. On the 23rd inst. two Bell’s Tortoises were on
their backs, and this morning another. This is either done in
desperation or as a means of defence (24.ii. 22). Nevertheless,
two were dead yesterday ; the two which I had found on their
backs, I believe (Kilosa, 26. ii. 22).
Though there was plenty of grass in their enclosure one of the
tortoises was found to be eating hard maize corn; it would pick
up one in its jaws five or six times and drop it, then apparently
swallowed it whole. Another was’ found nosing a larye yellow
carpenter bee (Xylocopa nigrita) and was munching something
at the time. One of the hind legs of the bee was missing, and
the insect itself was very freshly dead, having probably fallen
from the beams above. I watched the tortoise return again and
again to the bee, but it never bit it. One knows the preference
of the Greek Tortoise for yellow flowers such as dandelions and
buttercups. Isit possible that. anything ofa yellow colour attracts
tortoises ?
To judge by some of the damaged shells on tortoises which
have been brought me, these creatures can sustain very severe
injuries to the carapace and the latter heal up.
It is the rule rather than the exception at [Kilosa for Bell’s
Tortoises to carry some half-dozen ticks (Amllyomma mar-
moreum Koch), each adhering to the fleshy portions of the hinge
of their shells.
TESTUDO PARDALIS Bell.
Blgr. Cat. Chelonians, 1889, p. 160.
Thirty-three specimens were collected from Pwaga, Mbonoa,
Singida, Mdjengo’s, Mtali’s, Zengeragusu, Ulugu, Tabora, Simbo,
Tambali, Luguo, Wembere, Sanga, Sagayo.
The Leopard Tortoise is frequently found in grassy steppe
country where there are scattered bushes, but it is also commonly
met with on kopjes, and often at a surprising height, as in the
ease of one whose skeleton was found between two sloping rocks
ati the very top of what was practically a mountain. One wondered
how many years it had taken to reach the summit, seeing that
the mountain was largely composed of piled-up boulders; this
alventurous individual would appear to have met its death by
sliding down the smooth sloping rock and being unable to
clamber up again (Mtali’s, 9. x. 22).
_ At Pwaga, where I was only camped for one afternoon, a local
native brought (in addition to the depressed and worn C. belliana
already referred to) no fewer than five Leopard Tortoises whose
depth of shell was in no way less than those inhabiting grassy
savannahs, though here they were in the same dense thorn-scrub
as CO. belliana. The range of sizes was so varied that they are
worth quoting. The largest tortoise was a male; the others
would appear to be all females. (1) 250x150 mm., (2) 130~x
85 mm., (3) 88 x 65 mm., (4) 70 x 56 mm., (5) 47 x 40 mm.
The largest male collected measured 312 x 210 mm. broad.
926 MR. A. LOVERIDGE: NOTES ON
Pairing took place daily during the rains (December to
January) amongst specimens in captivity. ‘lhe process of egg-
laying was witnessed on 21. v. 22 , and the following notes were
made at the time :—
“The tortoise enclosure measures 20 x 20 feet, two sides are
high walls supporting the grass roof, the other sides are netted
in. On visiting this enclosure at 5. 15 p-m. I observed a tortoise
busily digging with its hind legs. The selected site was the angle
formed by wall and wire netting and a post which carried the
latter. The hole was about six inches from both wall and
netting.
‘The ground was very hard, but she had already excavated to
a depth of about two inches in a rough circle of about four inches
in diameter. Digging was‘ accomplished by inserting one hind
foot which was scraped round and brought up with a very small
quantity of'soil on it, which often fell back into the hole at the
last moment. Nothing like the quantity was lost that might
have been, however, had not the tortoise discharged a quantity of
urine from time to time, which resulted in binding the light
powdery soil and caking ‘it to her feet. Both hind feet were
used alternately for digging, with occasional pauses between
changing them.
“ At 5.30 p.m. she began to undercut the sides all round by
a turning movement of the foot so that the claws cut deeply into
tlie sides of the hole in a semicircle ; the back part, as far I could
see, was not much affected.
«At 5.50 p.m. the hole was approximately four inches deep
and retained its original diameter. ‘The urine was not dis-
charged as at first but continued to drip fairly regularly and
‘almost continuously. By treading with her feet the soil was well
puddled, and yet again and again “the caked soil would fall from
her foot just at the moment when she was clearing the brink of
the crater, and thus rendered her work much more laborious.
«* At 6 p.m. she scooped round and round the hole, which now
measured almost five inches in depth.
“ At €.30 p.m. darkness fell, and an acetylene lamp was
brought.
“At 7.12 p.m. the first egg was laid in a membrane which
lowered it quite easily and gently to the bottom of the hole.
‘* At 7.14.50 p.m. the second egg slid slowly down the mem-
brane which connects the first egg with the mother. Yet it
did not fall on the first egg but lies alongside of it.
“‘ At 7.15.50 she put her right hind foot down and pushed eggs
slightly to one side causing them to grind one against the other ;
we therefore felt and found them to be anes hard-shelled
when laid. .
“7.17.50 third egg laid.
— 7.19.10 fourth egg laid.
--“7,21 put her left hind leg down and felt around, a fifth egg
was laid about this time,,
“7,22 a sixth egg was laid,
EAST AFRICAN TORTOISES. 927
“7.22.50 put her right hind leg down and moved all eggs from
the centre of hole to beneath the ledge of undercut.
“7.24.10 seventh egg laid, which appeared to be slightly
smaller than its predecessors. I have omitted to mention that
she raised herself slightly before depositing each egg.
“7.25 p.m. put down her right ‘foot and pushed the eggs
about quite forcibly till they lay in a semicircle, three-quarters
of each egg being in the undercut.
‘€ 7.26.10 put down her left foot.
“7.27 put down her right foot.
“7.27.50 put down her left foot. It now became obvious
that.she was scraping down the sides of the crater, using her feet
alternately. ;
_.‘8p.m. She now commenced a very definite treading move-
ment with the object of pounding the soil. The action was very.
mechanical, and she dug two little pits with her hind feet to find
sufficient earth to cover the eggs rather than utilise the earth
already excavated which lay scattered about.
** Not once during three hours had she faced about to examine
the result of her labours or inspect her own eggs, which she had
buried without ever seeing. I was so struck by the mechanical
nature of the proceeding that I picked her up and removed her
a couple of feet away from the hole and on hard ground. ‘There
she continued trampling away without interruption—it was a
sorry spectacle to see so much misplaced energy.”
I replaced her over the original’ site and left her to trample
it to her heart’s content. In the morning she was resting
motionless on the spot and remained so till I left at 9 a.m., how
much longer I do not know. The next evening she returned and
slept on the spot, and I noticed she did so many times, probably
every night. \s
I had removed the fifth egg before she filled in the hole, and
this measured 35x32°5 mm. The shell was exceedingly hard,
thin, and brittle, and took half an hour to drill. The albumen
was turgid, coming forth in lumps. The yolk was an unpleasant
ochre, very different from that of a fresh fowl egg.
On the 12th of June I made a note to the effect that, in addition
to sleeping on the spot where her eggs were hidden, she relieves
herself and makes her droppings on the site frequently, if not.
daily. I sent her away about the end of the month.
On July 8th it occurred to me that it might require her assist-
ance for the young to escape from the “ nest,” as the surface of
the ground above them had set as hard as cement. I therefore
dug down to the eggs and found the top of the uppermost only
2} inches below the surface. They had not hatched, so 1 left
them till 21. vili. 22, 7.¢., three months from time of laying and
then dug them all up. They had not developed embryos (so
I conclude that they were infertile), nor had they decomposed
beyond a slight smell. HG °
In no fewer than six of the localities mentioned for this species,
928 MR. A. LOVERIDGE: NOTES ON
remains, generally of half-grown individuals, were found. One
at Mbonoa had definitely been killed by a beast of prey or a man.
The local natives (Wataturu) stated that they did not eat them,
but this was probably untrue, as they ate agama lizards, rats,
and other creatures.
The tick (Amblyomma marmoreum Koch) was found on a
Pwaga specimen.
TESTUDO LOVERIDGI Bler.
Blgr. C. R. Acad. Sci. Paris, 1920, t. 170, p. 264.
Love. Jour. EK. A. & Uganda Nat. Hist. Soc. 1921, p. 50.
Procter, Proc. Zool. Soc. 1922, pp. 483-526.
Thirty-six specimens from Dodoma and Tabora; remarks on
the habits of these have already been published as an appendix
to Miss Procter’s paper. A dozen of these tortoises escaped at
Kilosa, and it will be interesting to observe if they survive the
numerous carnivora without having any rock shelters under
which they can seek refuge.
No ticks were ever found on wild specimens, the sandy soil of
their habitat being unfavourable to these pests. A tortoise
which escaped at Kilosa picked up a tick (Amblyomma mar-
moreum Koch) within twenty-four hours.
TESTUDO PROCTERA, sp.n. (PI. 1., IT.)
? Two specimens from Ikikuyu and Kidenge in Dodoma Dis-
trict. The localities are about a hundred miles east of Dodoma,
the type-locality of 2’. loveridgii. Ikikuyu, which is to be con-
sidered the type-locality of the present species, lies some fifty
miles south of Igulwe (Gulwe) station on the Central Railway.
The country is different from that of Dodoma in that there are
no essentially rocky kopjes, though there are small hills with
fairly numerous seattered boulders.
Within a mile of the spot: where this specimen was found
there are precipitous cliffs topping the mountains, and the
slopes below are boulder-strewn and clothed with a dense thorn-
bush thicket which is almost impenetrable. The more level
country at the foot was, doubtless, similarly covered at one time
but is now sandy thornbush steppe, amongst whose bushes the
native-owned herds of cattle and goats wander. It was here
that the tortoise was found eating a mushroom at 6 a.m.
Superficially it is very similar to 7’. loveridgit, but differs in
the following points :—
(i.). Greater degree of ossification of the bony exoskeleton,
particularly the carapace (the diamond-shaped fenestra -
tion of the plastron is similar to that of 7’. loveridgii).
(ii.) Greater depth of shell in proportion to length and
breadth which may be best shown thus :—
EAST AFRICAN TORTOISES. 929
Breadth. Depth.
Species. Length cant ne Whereabouts. :
in inm. aU
mm. | percent.| mm. | percent.
T. loveridgii ...| 865 77 90°5 19 | 26 Tring Museum.
T. proctere ...| 8d 72 84°65 40 47 British Museum.
In the large series of 7’. loveridgii examined by Miss
Procter the greatest depth per cent. of any specimen
was 33:4 mm., and that was of the youngest example
only 42 mm. long. The depth per cent. decreases
with age, as has been shown by Miss Procter.
(iii.) Presence of a strong vertebral keel on all five scutes.
(iv.) Reversion of the marginals, which are turned up so that
their outer edge forms an obtuse angle with their inner
edge and in some an almost cup-like depression.
(v.) Coloration—The centre of each scale is dark sepia,
almost black, surrounded by a broad yellow margin; the
star-like radiations of 7’. loveridgiit are absent.
Owing to its immaturity it is impossible to say definitely that
it might not be referable to the genus Cinimys, as the character
istic hinge of the carapace in that genus is not developed in
young individuals. I am confident that it is correctly assigned
to the genus Testudo. C. belliana was taken at Ikikuyu on the
same day as 7’. procter, and though depressed the ossification is
fully developed. 7’. tornieri, it may be remembered, was origin-
ally mistaken for a pathological phase of C. belliana by Tornier *.
1’. procter I consider to be one stage nearer the typical box
tortoises than 7’. tornieri.
Sex ?. Half-grown.
Type-locality. Ikikuyu, Dodoma Dist., Tanganyika Territory,
12. ii. 23.
This specimen was kept alive for three months in captivity,
but unfortunately died, and decomposition had set in when it
was found. The reptile was immediately preserved, but in poor
condition. It has been presented to the British Museum.
The second specimen was found beside a smallstone at the foot
of a rock-strewn, scrub-covered mountain (Mt. Hundugula) at
Kidenge. It was so young that I hesitate to refer it definitely
to this species, excepting on the grounds of locality, as it was
not twenty miles from Ikikuyu. It measured 40 x 37x17 mm.
(greatest length, breadth, and depth of shell), and was taken back
to Kilosa, where it was left in charge of a native who lost it in
cleaning out the vivarium. So rarely are these creatures met
with that the local chief said that during the thirty years of his
residence he had never heard of a tortoise being found in the
neighbourhood.
A tick (Amblyomma marmorewm Koch) was on the type-
specimea when found.
* Tornier, Kriechtiere Deut. Ost-Afrikas, 1897, p. 2.
930 Mf, A. LOVERIDGE: NOTES: ON
CHELONID 2.
CHELONE MyDAS Linn.
Blgr. Cat. Chel. 1889, p. 180.
A carapace was obtained from fishermen at Dar es Salaam in
1922 ; these turtles are caught fairly frequently. A live one was
seen alongside the ship in Tanga: Harbour (27. v. 23); it thrust
its head out of the water close alongside us, and could be seen
to advantage as it swam about.
PELOMEDUSIDS&.
Prnusios Niaricans (Donnd.), _
Blgr. Cat. Chel. 1889, p. 195.
Three specimens from the Ruaha, Kilosa, and Kissaki in
Dodoma District. These were adult, half-grown, and immature
respectively, the smallest measuring 47 x 40x 13 nm.
PELOMEDUSA GALEATA (Schoepff).
Blgr. Cat. Chel. 1889, p. 197.
Ten specimens from Dodoma, Mahake, Kilamatinde, Mtali’s,
and Luguo.
Of these the largest male eietiedia lbs. and measured 200 x
135 x 68 mm., the largest female 142 x 112 x 29 mm., and a very
young one from Mahaka on 10. iii, 22, measured 49 x 39 x 18 mm,
After two heavy rainstorms on the two preceding days, which
heralded the advent of the rainy season, a P. galeata was found
at 8.15 a.m. busily engaged in trying to dig itself into the middle
of a sandy road. The rains had doubtless disturbed it and
caused it to wander. (Mtali’s, 20. x. 21).
My wife reported having seen a Lesser Stink Ant (Paltothyreus
tarsatus), which had fallen into the water-tanl, seized by one of
these tortoises. This is remarkable, as the ant has a formidable
sting, as well as good jaws and a protective smell. We therefore
experimentally introduced three more, which were taken without
hesitation by the tortoises, who snapped them in half. The head
and thorax were first crunched up and then the abdomen picked
up and eaten. Grasshoppers and spiders were also taken, one of
the latter being warningly coloured in black and yellow.
A Key to rue Tortoisrs or TANGANYIKA TERRITORY, KENYA
Cotony, AND UGanpba.
Synopsis of the Families.
I. Shell covered with epidermal, horny shields,
A. Pectoral shields of plastron in contact with the marginals.
1. Plastral shields 11 or 13 ............. ceeseceesseneseeeee LESTUDINIDA,
2. Plastral shields 12 as intergular eee Leevsteeeeeee LELOMEDUSIDR,
B. Pectoral shields widely separated from ‘te marginals;
limbs paddle-shaped, with one or two claws.. saeeee CHELONIDA
Il. Shell without epidermal shields ............ccccccceesereeeeeeeeeees RIONYCHIDR,
EAST AFRICAN TORTOISES. - 931
TESTUDINIDS.
‘T'wo genera.
Synopsis of the Genera.
Posterior portion of carapace hinged, movable in adulé ............0..... Cinirys.
Posterior portion of citapacé immovable oi eee eeeeeseeeeee Lestudo.
CInIXxys.
Bell, Tr. Linn. Soe. xv. p. 398 (1827); Blgr. Cat. Chel. p. 140
(1889).
One species.
1. Cinixys BELLIANA Gray. Bell’s Hinged Tortoise.
Bigr. ¢.c. p. 148. .
Tanganyika Territory, Kenya Colony, Uganda.
TESTUDO.
Testudo, part., Linn. 8. N. i. p. 350 EE) ; Blgr. Cat. Chel.
p- 149 (1889).
Jfour species.
Synopsis of the ee
I. Shell deep, box-like .. 000 secseccsecsessesss pardalis.
IJ. Shell depressed, wields ip slight ee
A. Vertebral shields keeled .. dadsssadeacsedsGerscsteossotdsusvessse-sse PUOCLENE
B. Vertebral shields not easly.
1. Narrower. Breadth of Vain three- ae the jones in
the type... ; Veseseseeseee tornicri.
2. Broader. Breadth of plastron poroe “han length in young
to 13/16ths i in adult... chauPenecidaucna Sines dasseenecsneoeen ness LOUGNIAGIt.
1. Tesrupo PARDALLIS Bell. Leopard Tortoise.
' Blgr. t.c. p. 160.
Tanganyika Territory, Kenya Colony, Uganda.
2. Testupo procrer& Loveridge. Keeled Soft-shelled Tortois e.
Love., present paper. ees ;
Tanganyika Territory.
3. TESTUDO TORNIERI Siebenrock. Tornier’s Montuices
Sieb. 8.B. Ak. Wiss. Wien, vol. exii. p. 443, 1903.
Tanganyika Territury, Kenya Colony.
4. Trsrupo LoveRrIpait Boulenger. Soft-shelled Land Tortoise.
Blgr. Comptes Rend. Acad. Sci. t. 170, p. 264, 1920.
Tanganyika 'Torritory.
*“PELOMEDUSIDS,
Two genera.
932 MR. A. LOVERIDGE: NOTES ON
Synopsis of the Genera.
Mesoplastra extending right across the Been: front lobe of
plastron movable in the adult ....c.ceccesse teessrecseeeeee Slernotherus.
Mesoplastra small and lateral; front Igbo. of bso without
in wey pny adwlbie ecerencseenee tere eaeeoe cece cere rae eeenrecenen eos see un PLZOLONPGWUSL
STERNOTH ZRUS.
Sternotherus, part., Bell, Zool. Jour. ii. p. 305 (1825); Blgr.
Cat. Chel. p. 191 (1889).
Three species, (S. sinuatus and S. nigricans doubtfully dis-
tinct and difficult of determination.)
I. Beak notched mesially, with a very slight, obtuse cusp on each
side of the notch; interorbital width considerably less than the
longitudinal suture between the frontal shields ..................... simuatus.
II. Beak neither hooked nor bicuspid; frontal suture non or p Tbe
slightly exceeding the width of the ‘interorbital space.
A. The length of the outer border of the hector) shields equals
that of the humeral ................ .. nigricans.
B. The length of the outer border of the ‘edie | is 5 eh ese
than that of the humeral, and does not exceed the length of
the inner border of the latter shield .. soHobapDbqac Bd Le Boabeo. | CHAR ATER DUI
1. SrernoTHERUS rs te Smith's Water Tortoise.
Blgr. t.¢. p..14...
Tanganyika Territory, Kenya Colony, Uganda.
2. STERNOTHARUS NiaRicANs (Donnd.). Black Water Tortoise.
Blgr. t.c. p. 195.
Tanganyika Territory, Kenya Colony, Uganda.
3. STERNOTHZRUS DERBIANUS (Gray).- Derby’s Water Tortoise.
Blgr. ¢.¢. p. 195.
Uganda.
PELOMEDUSA.
Wagl. Syst. Amph. p. 136 (1830); Blgr. Cat. Chel. p. 197
(1889). ‘
One species.
1. PeLomepusa GALEATA (Schoepff).
Blgr. é.c. p. 197.
Tanganyika Territory, Kenya Colony, Uganda.
CHELONIDA,
Two genera.
Synopsis of the Genera.
Costal shiclds in four pairs ........c.cccssseetereee sce see ssessssscseseeeeee Chelone,
Costal shields in five Or MOKe PAITS eee see epeeseeee Dhalassochelys.
EAST AFRICAN TORTOISES. 933
CHELONE.
Onelonia, part., Brongn. Bull. Soc. Philom. ii. p. 89 (1800).
Chelone Blgr. Cat. Chel. p. 180 (1889).
‘wo species.
Synopsis of the Species.
Carapace unicarinate in young. Limbs usually with a single claw... mydas.
Carapace tricarinate in young. Limbs usually with a double claw ... tmbricata.
1. Curtonr MypAs (Linn.). Green Turtle.
Blgr. é. c. p. 186.
Tanganyika Territory, Kenya Colony.
2. CieLone mBricata (Linn.), Hawksbill Turtle.
Blgr. t. c. p. 183.
Tanganyika Territory, Kenya Colony.
THALASSOCHELYS,
Vitz. Ann. Wien. Mus. i. p. 121 (1835); Blgr. Cat. Chel. p. 184
(1889).
One species.
1, THALAssoonELYs CARErTA (Linn.). Loggerhead Turtle.
Tanganyika Territory, Kenya Colony.
TRIONYCHIDA.
One genus. (Cycloderma in neighbouring territories.)
TRIONYX,
Trionyx, part., Geoffr. Ann. Mus. xiv. p. 1 (1809); Blgr. Oat.
Chel. p. 242 (1889).
1. Trionyx TRiuNGuis (Forskal). Freshwater Soft-Tortoise.
Victoria Nyanza,
EXPLANATION OF THE PLATES,
Prater I.
Type of 7. proctera, sp. n., from a water-colour drawing made after death.
Pratz II.
Type of T. procter, sp. n., dorsal and ventral views from X-ray photographs.
Proc. Zoou. Soc.—1923, No. LXI, 61
ee
Aaa ee
NOTES ON BAST AFRICAN LIZARDS. 935
46. Noles on Wast African Lizards collected 1920-1923,
with the Description. of two new Races of Agama
hionotus Bler. By Axruur Loveripes, F.E.8., C.M.Z.S.
[Received October 3, 1923: Read November 6, 1923.]
The following notes are based on a collection of lizards and
chameleons collected by the writer during six months in Kenya
Colony (formerly British East Africa) and two and a half years
in Tanganyika Territory (late German East Africa).
Representatives of all the families of East African Lizar ds were
again collected (with the exception of Anelytropid), more than
half the genera and a. little more than one-third of the species.
The numbers are distr ibuted as follows :—
No. of
Family. Genera. Species. Specimens.
Geckonidee 220.0203.) 02. 4 Tie UN 92
Avoid a eaves sae 1 4) 128
OMIA 22 Me eee 1 J 2
Waranidse een ee 1 2 7
Amphisbeenidee ............ ] 1 2
Macertidaee ae ee ee 5 ies 249
Gerrhosauridi ...........- 1 2 4
Scinerd:ey eee ewe 4 12 119
Chameleontidee ......... 1 7 42
Total... 645
Of the forty-eight species, however,’ no less than seventeen
were collected by the writer for the first time and of these two
were undescribed. Particular attention is directed to such
rarities as Pachydactylus boulengeri, Geocalamus modestus, Algi-
roides allen, Mabuia irregularis, Mabuia obsti, and Melanoseps”
ater var. longicauda. Several of these species have only bees
known from the types hitherto.
Taking them from east to west and south to north, the prin-
cipal localities mentioned in the following pages are :—
Kenya Conony. rachis
Kilindini, Mombasa Is., Frere Town, Voi, Nairobi, Eldoret,
Kisumu, Karungu Bay. ;
TANGANYIKA TERRITORY.
. Dar es Salaam District.—Dar es Salaam. ; ie
_ Morogoro Disirict.—Bagilo, Mt. Kirui, Mkindo River,.
. Wami River, Mkata River, Ilonga, Kiniamba Station,
» Tindiga, Kipera, Kilosa, Kidai, Kidete, Kisanga, Mbala. ,
61*
936 MR. A. LOVERIDGE: NOTES ON
Dodoma District—Lukole, Ikikuyu, Mpanira-kwa-Sagoi,
Kidenge, Itende, Pwaga, Igulwe, Dodoma, Kilamatinde,
Mbonoa, Ndogwe, Suna, Gwao’s, Mbulu’s, Pooma, Singida,
Mdjengo’s. at Bat Meer
Kondoa-Irangit District —Mtali’s, Mkalama, Usshora, Ulugu.
Vabora District.—Tabora, ‘Izikisia, Ndala, 'Tambali, Luguo.
Mwanza District.—Sanga, Lalago, Mwadira, Shanwa, Sagayo,
Nyambita, Mwanza.
Bukoba District.—Bukoba, Kabare.
The determinations and draft of this paper were made in Nast
Africa, but three days were spent in going over them again at
the British Museum, where I had the advantage of comparing my
specimens with allied forms in the National Collection. In this
connection I would like to express my most grateful thanks to
‘Miss Joan B. Procter, F.Z.S., for much help in examining dubious
determinations and for making notes for me on spécimens sent,
home. “py aa buct hi
I have given varietal names to two Agamas to emphasise their
very distinctive colouring. One of these forms | have already
referred to in a previous paper. “G00 ORL eho \
As before * measurements are given to the néarest eighth of
an inch followed by the exact measuréments in millimetres of the
length from snout tq vent, followed by thé tail length when intact ;
reproduced tails are denoted by an “r”., Maximum lengths are
only given when these exceed .thosé recorded in the previous
collection.. ;
Once again I am indebted to Dr. H. A, Baylis, F.Z.S., for
carrying out the determinations of parasitic nematodes found in
so many tropical lizards.
GEOKONIDSA.
HemipactyLus MABouia Mor.
Bler. Cat. Liz. i, 1885, p. 122.
A further large series of forty-five specimens of the House
Gecko. were collected from the following fresh localities :—
Zanzibar, Bagilo, Mkindo, Ilonga, Kilosa, Mbala, Mt. Hundu-
gula at 1200 feet, Itende, Kilamatinde, Mkalama, Kisumu
Frere Town, and Mombasa Is. | Sete ae om
Mjusi kafiri is the’Kiswahili name for this species. Kololo-
lumbusi in Kigiriama according to Koch, ae
,. On Mombasa Is. a. series was taken on baobab trees, where
they attain to a greater size than in houses, in such situations it
is exceptional to meet with them less than 80 mm. in length from
snout to vent. I am disposed, therefore, to think that they
desert houses as they grow older and take to trees. But the eggs
are common enough in fissures of the bark, though the young are
* Loveridge, “Notes on East African Lizards collected 1916-1919,” Proc. Zool.
Soc. 1920, pp. 131-167.
- - BAST AFRICAN LIZARDS.’ ” 937
only occasionally met with, so if my theory is ‘correct the young
must find their way back to the houses.. Young specimens aré
commonly: met with in. rubbish heaps or accumulations of
vegetable matter in waste land near houses. Examples caught
on bushes are olive-green in colour when found and look very
distinct. pil apn as ne
“he chin shields of the present series show considerable
variation ; normally only the first pair are in contact behind the
mental, sometimes the second pair are also, again in one Mombasa
specimen the shields forming the second pair are transversely
divided. There is little profit in discussing the abnormalities
of a species whose wide variation is evidenced by the dozen
names under which it has been described from time to time. -
I wonder whether it is generally known that daylight can be
seen through a gecko’s head if the ear-openings happen to be in
line with the light. A gecko used to lie on a certain bar just
at sunset at Kilosa, so I had the opportunity of seeing this
phenomenon many times. - hee ON ig
_ A six weeks old gecko measured 45.mm. over ‘all; only one of
the present series was longer than those recorded from my
previous’ collection, and ‘the tail was unfortunately reproduced
(87+60r), so that over all it was not greater. ‘Lhis fine speci-:
men came from Bagilo in the Uluguru Mtns. |, “3
« A Frere Town, specimen had a bifid tail, the regenerated.
portion being the longest (52456 & 59). A gecko with only
three legs was seen several times at Kilosa. — aOR As phe
> The following note on mating was made at Kilosa 1. vi. 21.:—
“ Whilst. working -by lamplight at 7:10 p.m., I heard a. noise
‘tchk, tehk,’ and looking up, saw two large geckos on the out-
gide of the mosquito gauze. which enclosed the verandah. | One
had seized the other by the right side of its neck: apparently just
at the monient my attention was attracted, a second Jater, there
was a rapid movement as the posterior portions of their bodies:
met and pairing took place. Moving the lamp possibly scared.
them, for’ they separated a minute afterwards, running away in
different directions.” Boaanes
«: Ib'seens probable that eggs are laid at.all times of the year, for
besides Mareh, J uly, October, December already recorded, a pair,
of eggs were found in a lock at Frere Town on 2.wiil. 20, five
beneath an earthen pot at Kilosa on 13. v: 22, twenty : beneath a!
pile: of rotting palm-leaves at Dares Salaam on A. xii, 22, five
beneath logs at Mbala on 26.ii.21; also at Dar es Salaam and
Frere Lown on’ v.23. The:Kilosa’ eggs hatched on 20 and
2}. v.22, and nine of the Dar es Salaam eggs within a fortnight:
of being collected. - 7 Teabe
: Dietetic records are best described ‘from my notes direct :—
- “Shot a gecko on trunk of palm-tree with a polydesmid in its
mouth.” (Zanzibar, 19.xi.20.) Beet a
. A half-grown gecko captured a tsetse (Glossina morsitans) on
mosquito-gauze of the verandah.” -(Kilosa, 1. xi. 20.).% + 4
}
938 MR,A. LOVERIDGE.: NOTES ON
. “A large mantis was resting on the outside of the gauze in
bright sunshine at 3. p.m. when a gecko darted upon it, but the
mantis managed to elude it and escape.” (Kilosa, 5. v, 21 .)
“At 7.45 p.m. I was sitting at work when I heard a sharp
rapping at.the far end of the verandah which sounded precisely
like the hammering of a rat’s tail when trapped, the volume of
sound being quite as large, On reaching the spot however, I found
it was a large gecko which had swallowed the greater part of a
centipede, but three- eighths of an inch remained in view together
with the caudal appendages. Disturbed by the light, the gecko
paused for a minute, then commenced spasmodically ham-
mering the side of its jaw on the wooden panelling, and then
disgor wed two inches of the centipede, which I judged by the
breadth .to be about three inches long. It rested for a while
with this object hanging. limp and motionless from its mouth
except for giving it.an occasional knock. Presently it ran.up a
post and along the spouting of the gutter, where with a few gulps
it re-swallowed the whole. From the time I became a spectator
fill all was over’ was ermera rs like ten minutes,” (Kilosa,
14, vi. 21.)
‘. “ Hearing a gecko thumping some abet against the wall out-
side, I found it to, bea large cricket. To prove what a noise they
can make against a stone ‘wall, I measured the distance from the
gene of the combat to where I had been sitting and found it to
be 23 feet.” (Kilosa, 9. vii.21.).
“A fly (Wegritomyia maculipennis), yhichia is an excellent 1 mimic
of one of our wasps, was crawling on a white and lighted ceiling
at 8 p.m., when 1 observed a gecko stalking it. When the gecko
was just about the right distance for a spring the fly gave a buzz,
and the lizard, after atudy1 ing it a moment longer, withdrew.”
(Kilosa, 1 i. oe
- § At sunset I observed a gecko o on the gauze facing upwards to
a large blue hernet, the hornet adopted a menacing attitude and
then flew a few inches up the gauze and again settled, the gecko
ran after it but halted an inch. away and did nob attack, ” (Kilosa,
29.i. 22.)
- “ Hearing a hammering a few nights ago, I ran out to find a
gecko with a large male siafu or soldier ant.” (KCilosa, 31.1, 22. Me
_ “A gecko was hammering a black field cricket this evening.’
(Kilosa, 11. 111.22.) - I might add here that a Mombasa fe
had a cricket in its stomach, and a Dar es Salaam specimen dis-
gorged one when caught.
ots My attention. was. drawn, to a. gecko which seemed to be
struggling with something on the eround., On fetching a light
it was found to be flinging itself about and squirming over in an
attempt to rid itself. of a solpugid (Solpuga darlings) which was
holding the: gecko in its jaws, having hold of the head between
eye and ear. The skin was all gone from this area and there
were signs of bites on the right side and near the base of the tail.
I surmise that the gecko had first attacked the solpugid, but the
EAST AFRICAN LIZARDS. 929
latter would certainly have killed the gecko, which already had
a large black field-cricket in its stomach.” (Kilosa, 12.iv. 22.) -
A spider, moth, and acridian were in the stomachs of Mombasa
and Frere Town specimens. It is strange that some people still
kill these useful little lizards on mere grounds of dislike.
They do not appear to have many enemies. I caught a
Spotted Wood-Snake (2. semivariegatus) in the act of swallowing
one in the fowl-house, and the gecko had not dropped its tail.
A month later I caught another Spotted Wood-Snake, and when
placed in the vivarium it disgorged a bolus of one of these geckos
which also had its tail intact. One was found in the stomach of
a Monitor (V. niloticus). :
When large numbers of these geckos live in a house it is quite
a remarkable sight tosec them leaving it at sunset to go foraging ;
they descend the walls from the eaves, and should there be shrubs
or tree-trunks within a foot or so, they reach them by a leap.
The natives put their own interpretation on these nocturnal
expeditions however, and informed me it was evidence of the
creatures’ shrewdness, as they feared to remain in the house
after dark lest a careless person should set it on fire!
According to Koch’s Vocabulary the natives say this lizard
dislikes man, and requests that God will destroy him in order
that the gecko may not be disturbed whilst eating sand. ~
ILnMipAcTYLUS SQUAMULATUS Torn.
Torn. Die Kriechtiere D. O. A. 1897.
One male and three females were collected at Kilosa on open
grass-free ground after dark; two were seen by the light of a
lantern as I was returning home, and the third was about nine
inches away from a giant field-cricket which was shrilling in a
most deafening manner. I wondered if the gecko was dazed by
the noise as it made no attempt to escape when approached.
Male measured 45 mm.+r, the Jargest female measured
4} inches (67440) and had two enlarged ovules in its ovary over
2 mm. in diameter (13. vii. 21).
The colour in life was pale reddish brown above with an
irregular network of brown lines on the body, and less distinctly
on the tail. The posterior end of the enlarged tubercles was
cream-coloured, very noticeable on flanks. A remnant of an
eye-streak is to be found in a small dot in front of eye anda
faint streak behind. Another specimen was dark nut-brown,
JIEMIDACTYLUS CITERNIL Blegr. :
Bligr. Ann. Mus. Gen. (3) v. 1912, p. 329.
Six specimens from Mbala, Lukole, Kidenge, and Itende, where
they were taken beneath stones on rock-strewn hillsides or, in a
couple of instances, beneath Jogs. ‘These records extend our
‘knowledge of the southerly distribution of this Somali species
‘considerably, - Sears A Eni hath aL
940 MR, A. LOVERIDGE: NOTES ON
The male referred to in my last. paper: has 7-7 upper labials
and 5-6 lower labials. -'.
The largest male measured 92 inches (836437 mm.). It is
a pale brown in life with darker brown spots tending to form
cross-bars.. Eyelids yellow, a good deal of yellow on labials,
many of the tubercles yellow.
Examination of stomachs negative.
‘“HEMIDACTYLUS BROOKII Gray.
Bigr. Cat. Liz. i. 1885, p. 128.
Five examples from Mkata River, Bukoba, Karungu Bay,
Kisumu,:and Mombasa; One was dug from a termite hill and
others were found beneath boulders on ne shore of Lake Victoria,
The female from Kilindini was beneath a tin near the shore.
One of these had 8-7 upper labials and only 6-6 lower. A male
larger than any I had taken before measured 44 inches (65452),
and caine from Mkata.; .
,. Both those from the termite hill and the boulders were quite
Pl ick when found but rapidly changed after capture.
Eggs, laid i in pairs, were found beneath boulders and measured
1] x "10 mm. One examined was found to contain a well-advanced
embryo (Karungu Bay, 16. xii. 22).
There is often considerable difficulty in distin guishing this
species from H. ruspolii, but.an examination of a. series of each
in the British Museum .shows. that whereas H. ruspolit has the
head covered with platés more or less equal in size and generally
keeled, the head of 2. brookit has enlarged, keeled, oval shields
scattered amongst very smal]. granules which cover the greater
Ey of its surface. .
A LyqopAcTYLUS FISCHERI SCHEFFLERI Sternf.
, Sternf. Ergebn. Deutsch-Zentr.-Afr. Exp. 1918, iv. 2, p. 206.
A single male from. Bagilo, Uluguru. It only differs from the
description of the type in having a lower number of prz-anal
‘pores—5 ‘instead of 6—and in the under surface being profusely
spotted except in the centre of the abdomen, as against “ Under-
‘parts pure white, at the most a few little ‘brown spots on the
chin-shields ” QE, ). ae ;
LyGopActTyLus GROTEI Sternf.
Sternf. S.B. Ges. naturf. Beylin. Nr, 4, 1911, p. 245.
Five specimens from. Kilosa, Itende, and. Tgulwe.
A male captured.on a stump in a rubber plantation at Kilosa
was coloured as follows:—“General colour grey. A. blackish
-stripe commencing at nostril passes through eye and almost
vanishes on groin though continued faintly on base of tail.
Thrée pale spots on line on flank: A second black spot on
’ BAST AFRICAN LIZARDS.. 941
canthus rostralis (forming with its fellow a A mark on snout)
passes above eye.'and breaks up into two lines dorsally which
vanish in lumbar region. ‘Tail reddish with faint greyish mott-
lings dorsally.” —
‘wo eggs (7x6 mm.) probably of this species were taken at
Pwaga near Itende. ‘Termites in stomach of Itende specimen.
LycopactyLus PicruRATUS Peters. :
Blgr. Cat. Liz. i. 1885, p. 161.
Seen at Mkindo River, Mkata Stn., Kimamba Stn., Kilosa,
Kipera. Madazini, and Kidebe. I had fired at a tree some
200 yards away at Kipera, and on reaching the spot picked up
the gecko, which had apparently been killed by the concussion
of the rifle-bullet as it penetrated the tree.
A male was found in a most unlikely place, viz. deep down in
a termite’s nest; in colour it was quite black, but soon regained
its characteristic yellow head when exposed to the light.
‘Two eggs measuring 5 x 6 mm, were found in a rotting pile of
palm-leaves at Dar es Salaam, 4. xii. 22, but did not hatch.
A. gecko of this species was found in the stomach of an
owl (Glaucidium capense scheffleri), Wami River, 3. ix. 21, and in
the stomach of a snake (Amplorhinus nototenia), Mkata River,
24. viii. 21. .
Thirteen specimens of the Frere Town var, were again collected
at that place and also at Kilindinion Mombasa Is. As previously
mentioned, the typical yellow-headed form was collected on the
Island in 1919, but a very thorough search for it proved
negative, and as it was caught at the station I imagine 16 wus
introduced. :
_ The largest. male measured 3 inches (37 +40) and female 2?
inches (39 + 40), being larger than the last record in body length
only, the tail being reproduced. .
PAcHYDACTYLUS BOULENGERI Torn,
_ Torn. Kriechtiere D. O. A. 1897, p. 26.
A single female collected near a dry watercourse on the Sagayo
road some fifteen miles west of Mwanza about 1 a.m. on 13, xi. 22.
_ Except for the blunter snout this gecko is superficially very
like H. mabowia, particularly in colour and markings. On closer
examination the feet of course are very dissimilar. This female
measured 22 inches (45+ 29).
“f PactypactyLus BIBRONII Smith.
|; Bligr. Cat. Liz. 1..1885, p. 201.
Kighteen specimens from Igulwe and Dodoma. | 0 re
The largest male measured 43 inches (60+ 56) and female 32
(51+46). One Igulwe specimen had a large cricket in its
stomach, the other five were gorged on termites.
949 MR... A. LOVERTDGN: NOTES ON
KLAsMoDAcTYLUS TRIEDRUS Bler.
Bligr. Rev. Zool. Soc. Afr. iii. 1913, p. 104.
Three specimens of this rare Congo Gecko were obtained at
Suna, in Singida, where they occur on the stems of the Bussu
Palma, where thore is abundant cover for thom among the
brinch- stalks and fibre. “6 .
The Wanyaturu call them Mwenkenyo and say that they in-
habit houses also, and when they groan someone falls sick.
I thought at first they were confusing it with //, mabouia, but
later saw Z. triedrus come out of the thatch of the hut 1 was
occupying and move down the wall to the entrance of a termite’s
nest in the floor, into which it vanished !
The largest of these males measured 5% inches (67 +82) and
the others 63+ 70 and 61+r.
One specimen had a plant bug in its stomach,
AGAMIDZ.
~ AGAMA IIISPIDA var. DISTANTT Bler.
Bigr. Ann. & Mag, N, H. (7) ix. 1902, p- 339.
The Wanyaturu, who have specific names for the various lizards,
call this one Lukhumbeli, and the Wanyiramba name is
JKimandagala.
Forty-three specimens were collected from Dodoma, Mbonoa,
Suna, Gwao’s, Mtali’s, and Mkalama in ‘Tanganyika ‘lerritory
and Delagoa Bay i in Portuguese Mast Africa (1914). This active
little Agana is essentially an inhabitant dof sandy thorn-bush
country, and though rocks may be abundant as at Gwao’s, it
leaves these to A. lionotus in the main and itself takes to trees if
chased. The five P. HK. A. specimens were taken as follows :—
‘'wo in sandy thorn-bush, one forty feet up a tree in the same
locality, one on a fence, and one under a heap of garden rubbish.
The largest male measured 84 inches (934125), and female
3 inches “(93-491). A young one taken at Delagoa Bay on
24 xi. 14, measured just over 2} inches.
The soil of Delagoa Bay being reddish the Iieavails harmonized
with it most wonderfully, nor did their colour fade on preservation
to any great extent, for nine years afterwards they are chestuut-
brown ‘and very different from the sandy-coloured form from
Gwao'’s. Colour notes made at time of capture read: ‘ Body
colour brick-red and brown, a cream-coloured dorsal line extends
from the parietal scale to beyond the hind limbs where it merges
into the cream shading of tail. ''wo of the smaller specimens
had bluish heads and « large blue gular spot surrounded by longi-
tudinally arranged series of smaller spots. ‘Ventral scales also
brick-red.”
' Gwao’s specimens were very ‘different. Male; “ Above sandy
or greyish fawn. A sienna-brown speck about centre of canthus
rostralis, two slightly V-shaped sienna-brown lines across crown
’ BAST AFRICAN LIZARDS. ‘943
between supraoculars. Similar markings, often interrupted on the
vertebral line, along back and tail, Four on back and nine on
tail and a very U-shaped one on nape. A few faint marks on
limbs. Below pure white excepting throat, which is dirty white
with eight longitudinal blue lines, and a large blue patch on
basal portion of throat.”
One female was uniformly sandy-buff, with only very faint traces
of tho-sienna markings of the male here and there. Uniformly
white beneath, the blue vermiculations found on the throats of
males are only faintly discernible on close investigation. Another
type of female had a reddish-brown ground-colour. Crown of
head dull bluish-grey with two very distinct cross-bars uniting the
supraocular regions. Cheeks brignt ultramarine. Throat cream-
coloured with rusty vermiculations and slightly discoloured basal
spot. Rest of under surface whitish. A slight cream dorsal
‘stripe from occiput to halfway along the tail bordered at intervals
by sepia blotches, of which there are five pairs on body and ten
pairs on tail.
One of my specimens has the occipital not enlarged.
Miss Procter observes that many lack the light vertebral line
and in some the ventral keels. are marked, both of which are
features distinguishing A. distanti from A. hispida. I therefore
consider that A. distanti is not a distinct species but only a geo-
graphical race of A. hispida.
Large white crickets were taken from the stomachs of two
specimens, ants were commonly met with, and once a ? glow- worm.
One of these lizards was found in the stomach of a Roller
(Coracias neevia). _
At Gwao’s they are commonly afflicted with a parasitic worm
which proved to be a new species (?hysaloptera gigas Spaul), and
also Strongyluris brevicaudatu Miiller.
AGAMA MOSSAMBICA Peters.
Blgr. Cat. Liz. 1. 1885, p. 363.
Four specimens from Kilosa, Izikisia, Ndala.
An immature male from Kilosa had 70 scale-rows, Izikisia
specimens 75, and the Ndala female 84. The ear-opening of the
last-named specimen was noé “much larger than the eye-
opening”; its third and fourth toes equal, also fifth toe not ex-
tending beyond the first. ‘Tail noé twice as long as distance from
gular pouch to vent, the distance being 60 mm. and tail length
100 mm.-In some specimens from Nyasaland in the British
Museum the ventrals are unkeeled, and in others from the same
locality they are keeled.
'. The largest male measured 104 inches (934165), and female
7% inches (85 4-100). Pyles
In the immature male from Kilosa there is a brilliant blue
upper labial streak, and a blue band across in front of the eyes
from side to side, a network of faint blue lines on the throat
and a darkish patch in front of the shoulder.
944 MR, A, LOVERIDGE: NOTES ON
There’ were thirteen large ovules (13x 8 mm.) in the ovary of
a specimen collected at Ndala on 15. xi. 21.
Amongst stomach contents were the remains of a scorpion, a
gvasshopper, and what appeared to be ants though they were much
-crushed.
AGAMA LIONOTUS Var. DODOMZ, var. n.
Known to the Wanyaturu as Khombele and the Wanyir amba
as Kinsuku.
Thirty-five specimenswere collected of the Dodomavar.at Kidai,
Kidete, Pwaga, Kidenge, Ikikuyu, Dodoma, Ndogwe, Gwao’s,
Pooma, Mdjengo’s, Mtali’s, and Izikisia, and what were believed
to be this form were seen at Mkindo River and Kilamatinde.
The local distribution of this species depends on rocks, for it
does not occur outside boulder-strewn or rocky--kopje country,
lt shows adaptability in that the Kilamatinde spockuens were
seen on the walls of a house.
The number of scale-rows at mid-body does not ‘appear to be
of much help in defining this variety.
2 Dodoma males 80, 89..............e0.. Gives an average of 84 scale-rows,
1 Taikisia | » 80 "5 as % 80 “4
7Gwao’s ,, 71, 72,74, 74, 74, 77,80 | ,, Sn ee olnaed #3,
1 Pooma”’ ',; © 70 Bae Hi baer iene a FONE ea:
4Mtali’s ,, 70,75, 76, 79 SC ee Oe chee
+ 13 Mtali’s females.70, 71, 72, 72, 73, 74,.74, ;
7, 76, 76, 76, 77, 79 Pept ere ay (Nae are
‘The localities .are arranged. from south.. to north, pall Th think,
though the series may be considered small, bear out my previous
contention that as you proceed south: from: the type locality of
‘A. Lonotus (Lake Rudolph) you get an increase in scale-rows,
T do not think this means .that the scales are smaller but that
the body girth is greater, in proportion to the greater oes
attained by the more southerly form.
The femoral pores of eight | males examined range from 10 to
13 with an average of 11.
I therefore feel. justified in proposing a varietal name for this
strikingly handsome form according to the description given on
page 142 of the P. Z. 8. 1920, from a specimen in my collection.
I now designate as types a male from Gwao’s, 10. x. 21, and a
female from Gwao’s, 3.x.22, which have been donated to the
British Museum.
The coloration of an immature male from Dodoma (11. vii. 21)
may. be of interest as it is so different from the adult :—“ Above
slaty-grey intermixed with brownish-olive or dull-coppery colour ;
small irregular patterns in dark brown on head, neck, and fore-
limbs; a semicircle of cream spots on parietal region with occipital
scale as apex of the semicircle having one cream spot anteriorly
in contact, and one on either side; coppery colour best developed
-on neck, where there are two slightly curving creamy stripes well
separated from the faint nuchal crest. Whole of lower surfaces
EAST AFRICAN LIZARDS. — 945
white, the throat only being mottled or streaked with greyish
reticulations.”
There were eight eggs (23 x 11 mm.) in the ovary of a Kidai
female and apparently quite ready for laying (18. ii. 23). Young
ones about two inches long were very numerous between Kidenge.
and Pwaga (ii. 23).
In the stomach of Giwao’s specimens was a quantity of fruit,’
in another the leaves of ‘“upilo.” Other indeterminate
vegetable matter in many others, also termites and ants of various
species, including, I believe, a stink-ant (Paltothyreus sp.). At
Izikisia I watched one of these lizards snapping up termites
after a rain-storm, which was causing them to emerge in large
numbers. ‘The stomach of an Ikikuyu specimen was crammed
with termites.
A very high percentage of these lizards are affected with a
parasitic nematode worm (Thelandros sp., 2 Q non det.).
Thad great difticulty in hunting these active lizards on the:
rocks until I offered a small reward to the delightful small boys
of the Wanyaturu tribe, who are experts with bows and arrows,
which they all appear to carry. They began shooting them,’
which I had presently to stop for lack of preservative. They gave
me to understand that they hunt and eat them in times of:
scarcity, and this 1s very possible, as these youngsters ae
ate the bodies of the rats as they were skinned.
AGAMA LIONOTUS var. MWANZ&, var. n.
The Wasakuma call this Agama “ kuli.” -
In Mwanza district forty-one specimens were collected of an.
entirely different Agama, but which has no specific characters by
which [can distinguish it from the foregoing. They are abundant
in Mwanza itself, but were not collected there, my series coming ,
from Mwadira, Shanwa, and Sagayo. For this colour-form I
propose the varietal name of mwanze, with Shanwa, Mwanza, as.
type locality. The g and Q types collected on 20,x.22 have,
been presented to the British Museum.
The longest neck-spines were Jess than half the diameter of tho
ear-opening, 7.e. 2mm.and 5 mm. Labials 10-11. Pre-anal
pores of the males ranged from 10 to 13 and averaged 11, this
being based on twenty-one males examined.
The number of scale-rows at mid-body according to ely
and sex are :—
1 Sagayo male 87 .............00605 gives an average of 87 scalerowe,
7 Mwadira males range from 71 to 90 ay 0 up Neca
165Shanwa ,, hh SOLES tre Wis, 76 mp
8 Sagayo females _,, » 710to 78 op ¢ 73 Ay
11 NO ep Pa » 70 to 85 Pera any (| a
which gives o ean B for the variety of 70 to 90 with an average
of 78.
The coloration of the ve male.—* Snout to just beyond mid-»
946 MR. A. LOVERIDGE: NOTES ON
body metallic purplish-pink ; fore-limbs from elbows to finger-
tips and whole of hind-limbs and tail a very rich metallic blue,
brighter below than above. The females are much more like
those of the Dodoma form, but are readily distinguishable by the
vermiculations on the back.” :
The greatest body-length attained by males in this series was
130 mm. in two specimens, both of which had regenerated tails ;
of the perfect males the largest was 131 inches (127+ 207). None
’ of the females collected were very large.
Females collected at Shanwa on 19, x. 22 had eggs developing
in the ovaries: (a) six measuring 20x10 mm., (0) eight eggs
17x10 mm,, (c) eight enlarged but still round 6 x 6 mm.
Iam almost inclined to say that this Agama is primarily a
vegetable feeder, and only secondarily insectiverous, Many of
the stomachs had nothing but vegetable matter in them, others
a mixture of vegetable and insect food ; of insects, ants appeared
to be the favourite fare, but it must be remembered that soft-
bodied insects such as diptera are so masticated that they are
vory difficult to find; a few beetle elytra were found and one bug.
I watched (both with and without glasses) a pair of these lizards
on several occasions come down from their rock and bite off the
sprouting green grass at its base and also the leaves of a small.
shrub. ‘the female always appeared bolder than the male, and
would venture away from the rock in pursuit of insects.
A Hawk (Melierax poliopterus Cab.), shot between Mwadira
and Shanwa on 19.x. 22, was found to have dined. off one of
these lizards, and a day or two later I saw a female Hawk of the
same species feeding its young in the eyrie with a full-grown
agama. Indeed, at Shanwa, unless the Kites eat these lizards
which simply swarm, there appears to be very little other food
for the large number of Kites which are ever circling about the
rocks,
The Agamas are also heavily infested with the nematodes
Strongyluris brevicaudata Miill., Uhelandros sp., and Stronyy-
luris, sp. nov.
AGAMA FLAvIcAUDA Werner.
Wern. Zool. Anz. xx. 1897, p. 264.
During November 1920 at Eldoret, Kenya Colony, I saw what
T am ‘almost certain were agamas of this species on a heap of
stones. ‘They were quite numerous, but I failed to capture any.
AGAMA ATRICOLLIS Smith,
Blgr. Cat. Liz. i, 1885, p, 358.
Upongo in Luguo (Kavirondo). - ee
Five males from Kilosa, Sanga, Kisumu, and Nairobi. This
lizard is thoroughly arboreal, most of my specimens being captured
on tree-trunks, ‘
iY
EAST AFRICAN LIZARDS. ° 947
The largest male measured 12 inches (165 + 240) ; this must be
something of a record for the species!
Preserved specimens give one a very poor idea of the colouring
during life. The following is taken from a Nairobi specimen
collected 14. viii, 20 :— Top of head coppery-green, sides of head
bright verdigris-green, edges of lower jaw the same. ‘Throat
light cobalt-blue, a few white spots below ear-drum on base of
jaws. A black blotch in front of each shoulder united by a dusky
pvtch across neck. Vertebral line from head to base of tail
occupied by a chrome stripe tending to green near the tail.
Anterior suyface of fore-arms cobalt-blue. Breast region dark
blue and dusky copper colour; belly lighter, with vermiculations.
Sides brownish, with black vermiculations and scattered yellow
scales. Under surface of all limbs and upper and lower surface
of tail brown.”
A Kilosa specimen, the large’one whose measurements have
just been given, had “Head bright cobalt-blue or coppery-green,
pvler scales intermixed with the darker on ‘the sides of face.
Dark blue on median line between eyes to about the occipital
scales. Tliroat bright blue in centre, remainder pale blue and
olive marbled with darker. Body olive, but heavily marbled
with black and speckled with blue. Tail pale olive ot base;
merging into blue about half its Jength. © Lower surfaces of
body, limbs, and tail pale sandy-or slightly rufous.” HP
This Agama consumes an enormous quantity of ants. I have
also found what were apparently ortliopterous eggs in its
stomach. | sis . TONNE. Si A, sl
I think that this Agama is the Mjombakdka of the Waswahili,
concerning which they believe that it will intercede for men
condemned at tlie day of judgment, when toads will come and
spit water upon them to alleviate their pain (Koch), . This ‘is
very interesting in view of the fact that N. African Agamas are
detested by good Mahomedans, who see in their bobbing heads a
caricature of the Moslem at prayer and think the lizard mocks
them. In this action of the Agama the Wakavirondo see a like-
ness to the motions of a woman’s head whilst grinding corn, and
consequently, when they see an Againa bobbing its head, enquire
if its mother is grinding to-day. Reinet?
GONURID S.
ZONURUS TROPIDOSTERNUM Cope.
Blgr. Cat. Liz. ii. 1885, p. 254.
One Zonure was caught running over papers on the table in a
tent at Msimba, Ilonga; another was found in the stomach of an
immature female Bare-faced Hawk (Polyborides typicus) at
Kipera, 5. viii. 22. Some two months later the former was
caught and eaten by a baby lemur which shared the same cage.
The Zonure fed readily on grasshoppers during its captivity. -
948 MR. ‘A. LOVERIDGE: NOTES ON
VARANIDA,
‘VVaranus ninoticus Linn.
Blgr, Cat. Liz. ii. 1885, p. 317.
The Ukerewe name for this Monitor is Kikwambo, the Ugalla
name is Mbulu, the Wasukuma call it Mbulu also.
_ Five specimens were collected at llonga, Kilosa, and Tindiga,
but also seen at Mkata River, Sagayo, Karungu Bay in
Tanganyika Territory, and Kisumu in Kenya Colony.
The largest female was 503 inches (519+761); monitors on
Lake Victoria attain an enormous size; a pair habitually basked
on a big rock south of Kisumu and were like small crocodiles.
One large female was killed by falling from a very high tree;
no branch fell with it, and the natives say that it is by no means
uncommon for them to fall asleep when basking on a branch and,
forgetting their position, move and fall.
In the stomach of this. specimen was a gecko (Z/. mabouia),
three large limicolarian shells, grasshopper legs, and. some cartila-
ginous lumps that looked like the pads of a cat's claws—if 80,
they had, been picked up from some skin thrown away. . Another
Kilosa specimen nearly 4 feet long had in its stomach 22 elephant
hawk-moth larve, 13 black coleopterous larve, and one cock-
chafer. Yet another contained crabs and a large quantity of
mud, A young male had two crickets and crab remains in its
stomach, None.had visible parasitic worms.
. Ticks (Aponomma exornatum Koch) were found about the
vent and feet of one specimen., Another monitor had them on
the fore-leg near armpit.
Varanus oceLzatus Riipp.-
Bler. Cat. Liz. 11. 1885, p. 308.
.. Kenge is the general Kiswahili name for monitors as Mbulu is
; oe the Wasukuma, who do not distinguish species. The Ugalla
have a special name for the Ry ed Monitor, which they call
Enumusa.
A male 48 inches long (557+ 657) was found crawling in the
road at Kilosa with bleeding mouth and fractured skull, It had
almost certainly fallen from a tall and solitary tree a few feet
away. Curiously enough a native had reported a ‘ pene”
being seen up this tree some months before.
A “single female 533 inches (597+ 763) from Tindiga had no
fewer than 35 eggs (60 x 40 mm.) in the oviducts, 13 of these
being in the left and 22 in the right. There were parasitic worms
(Physaloptera paradoxa v. Linst. ) in the stomach, and ticks
(Aponomma exornatwm. Koch) on the scales.
EAST AFRICAN LIZARDS. 949
AMPUISBANIDAE.
GrocALAMUS MopEstrus Gthr.
Blgr. Cat. Liz. ii. 1885, p. 453.
Two specimens which are, I believe, the only ones recorded
since Giinther described the species over forty years ago.
The type locality is Mpwapwa, and my collector obtained a male
from Ikikuyu, some 80 miles south of Mpwapwa, in sandy soil.
It measures 10? inches (240+ 34); 241 annuli on body ; a mid-body
annular ring consists of 36 segments and a caudal ring of 31.
The first Inbial on the right side appears to be incompletely
separated from the nasal. There are only three pre-anal pores,
one being obviously missing. The colouring above is plumbeous,
not brown, below pure white, somewhat transparent in life.
The other specimen is a young one, and was found in the
stomach of a Banded Mungoose (df. mungo colonus) at Usshora,
Mkalama Dist., 31.x.21, These records are extremely interesting,
extending as they do its known range to the west and south.
GEOCALAMUS AcuTuUS Sternf.
Sternf. Ergebn. Deutsch.-Zentr.-Afr. Exp. 1912, iv. 2, p. 209.
There is, in the British Museum, an Amphisbenid which I do
not hesitate to refer to this species. Collected at Samburu (near
Voi) Kenya Colony about 1897 by Betton. It has 209 annuli on
body and 22 on tail. It measures 230 and 29 min.
LAGCERTIDA,
Nucras Emini Bler.
Blgr. Ann. & Mag. Nat. Hist. (7) xix. 1907, p. 488.
The Wanyaturu know this lizard by the name of Kungulupe.
It is comparatively common in the dusty, sandy fields of stubble
which cover large areas in their district.
One which was captured alive at Mpanira had bolted down a
hole, obviously its own to judge by the tracks leading to it. The
hole was occupied in one portion by a giant cricket which had
probably excavated the burrow.
he Shanwa female darted across the path and into a very small
clump of grass surrounding a sprouting acacia. With the object.
‘of driving it out I set fire to the grass, which flared up and was
burnt in about two minutes. The lizard made no attempt to
escape and was picked up dead.
Forty-five specimens of a Nucras were collected at localities
intermediate between the type locality of WV. kilose (Kilosa,
Morogoro Dist.) and WV. emini (south shore of Lake Victoria), viz.
Tkikuyu, Mpanira-kwa-Sagoi, Kidenge, Suna, Pooma, Zinzerigi,
Mtali’s, Shanwa, Nyambita, Tambali, and Izikisia, representing
no fewer than four districts. ;
Proc. Zoou. Soc.—1923, No. LXII. 62
950 MR. A. LOVERIDGE: NOTES ON
The largest male measured 7% inches (62+ 120), and female 93
(75 +150).
Kxcopt in the matter of coloration and pattern and that the
keeling is less noticeable on the dorsal scales, they agree with the
description of V. kilose in all essentials. I had “only time to
exainine critically 25 of this long series, however,
Ventral plates in 28 to 32 rows; average of 25 specimens
30.
Dorsal scales 38 to 46 across mid-body ; average of 25 specimens
47-6.
- Femoral pores 9 to 12, with an average for the 25 specimens
of 11.
There are several individual variations. In one Pooma speci-
men the pre-frontals and frontal are rugose. The subocular is
between 4th and 5th upper labials in all cases except three
abnormalities, which are normal on one side. In two Pooma
specimens it is between 3rd and 4th, and between 6th and 7th in
‘one Shanwa specimen. In all 25 the normal number of 4 supra-
oculars were found. A Shanwa @ had 33 ventral scale-rows
with 40 scales across mid-body.
Miss Procter, though very busy at the time, kindly examined
the first 18 specimens which I sent home with a series of
N. kilos, and made the following memo. on them :—‘ The large
series which is before me points to intergrading between these
two species. The scales in emint forma typice ave granular but
keel-less. In the types of hilose they are strongly keeled. In the
present series the scales are tubercular (prominent), leading up
to the kilose state. The markings also vary. There are two
types—that of the typical kilose, and a speckled lighter brown
variety, ‘These points want going into thoroughly, Probably
kilose must rank asa subsp.” Unfortunately [ have been unable
to go over the whole series and critically compare thom with the
British Museum specimens, as I have had so much other work to
attend to whilst on leave.
Coloration of a male from Gwao’s was as follows:—‘ A white
lateral line from middle of ear-opening above fore-arm to hind-
‘limb; another commencing above ear-opening is broken up into
short dashes, and ends just above juncture of hind limb with
trunk; a third from outer border of parietals vanishes on tail ;
this one is bordered on either side by black specks; a fourth
(vertebral) line starting from occipital scale also vanishes on
tail; this one is bordered on either side by black specks. In all,
therefore, there are seven light lines and six rows of black spots.
General colour is sandy above, darker, almost olive, on crown of
head, Ventral and lower surfaces white.
A strikingly parti-coloured female from the same locality had
‘the head and forepart of the body to mid-body olive colour; there
‘are traces of only three of the light dorsal lines mentioned in the
description of the male; of these the lateral two are very white,
the interspace between them and above the other being black.
EAST AFRICAN LIZARDS. 951°
All markings vanish at mid-body, the posterior half being pinkish
shading to sandy-buff on the tail. Four other specimens have
the upper surface much speckled with black, approximately in ten
longitudinal rows. ‘Though showing niuch variation in colouring,
these lizards ave very different from WV, kilose and readily separated |
ab a glance.
A female (60+95) from Shanwa, 20. x. 22, had ovules rather
large ; another (67+r) from Ikikuyu, 10. ii. 23, held six ovules
measurmg 7X7 min. The testes of a male killed on .the same
day were enlarged.
Stomach contents of eight specimens examined were :—(1) cock-
roaches and termites, (2) a brown shield-bug, (3) grasshopper
and cricket, (4) spider, (5) cricket, many termites, and a spider,
(6), (7), (8) full of termites.
Nucras xinos.s Loveridge.
Move, PIZiS. 19225 p. 33.
Three specimens from Gwao’s and Mtali’s should be referred to
kilosee on colour grounds and possibly on the basis of keeled
dorsals. As pointed out under J. emini, there appears to be
intergrading in specimens collected mid-way between the type
localities of WV. emini and WV. kilose,and the latter is probably
only a subspecies of WV. eminz.
Since describing this lizard I have obtained eighteen further
specimens from the type locality. ‘I'welve were males, four females,
and two very young. ‘The disproportion in the sexes is probably
to be attributed to a greater agility on the part of the females
in eluding capture.
The largest perfect male measured 83 inches (65+ 147), whilst
another exceeded previous maximum measurements by being
67 mm. from snout to vent; the tail was reproduced in this
specimen. As the tail was missing in the type female, it
may be well to record the measurements of the three perfect
specimens in the present collection. (a) 50+117, (6) 504113,
(c) 43488.
A careful study of the scale characters of these specimens
enlarges our knowledge in several particulars, viz. :—
“ Ventral plates in 27 to 31 transverse series” should read 25
to 31. The average of these fifteen is 27.
“Dorsal scales 42 to 56 across mid-body” should read 40
to 56.
“Femoral pores 11 to 13” should now be 10 to 13, the
average of the fifteen specimens being 11:6.
In one the frontal and fronto-nasal were broadly in contact. :
Four of these specimens had grasshopper remains in their
stomachs. . 5
One specimen had a number of parasitic nematodes in the
stomach which were identified by Dr. Baylis as Physaloptera
abbreviata Rud.
62*
952 MR. A, LOVERIDGE: NOTES ON
Lacerta JAcKsoni Blgr.
Blgr. P. Z.S. 1899, p. 26, pl. x.
T take this opportunity of correcting an error on p. 147 of my
previous paper, when I hastily referred specimens of this lizard to
Lacerta vauereselli Tornier.
ALGIROIDES ALLENI Barbour.
Barb. Proc. New Eng. Zool. Club, iv. 1914, p. 97.
Hearing that Dr. J. Arthur was making an ascent of Mt. Kenya
this spring, I asked him to look out for tree-lizards, and he was
successful in obtaining three specimens of A. alleni on Feb. 4th
and 5th at 9970 feet.
LATASTIA JOUNSLONI Blgr.
Bigr. Ann, & Mag. Nat. Hist. (7) xix. 1907, p. 392.
Seven specimens from Kimamba, Tindiga, Tabora, Nyambita,
and Bukoba. Was also seen at Mkata Stn., Mkata River, and
all the way to Wami River along the foot-path. It shows a
great liking for sandy, sunny paths.
The coloration of breeding males and females is somewhat
different from that already described for this lizard in “ off”
plumage,” and was as follows :—
Male from Tindiga, 31. viii. 22.—‘‘ General colour reddish,
shading into nut-brown and pale fawn posteriorly and on tail.
The ‘blue spots on the sides’ are a very pale blue in one
specimen, and almost white in another. Upper and lower labials,
mental, lower sides of neck and belly (more particularly along the
edges) heavily blotched or patched with brilliant lemon-yellow.”
Female from Kimamba_ on 5, vill. 21.—‘* A yellow median
dorsal line with black edges commences at occipital and disappears.
on base of tail. A yellow line from the outer edge of the parietal
also vanishes on the tail. Another yellow line commencing on the
labials passes over fore and hind limbs and disappears on tail.
A series of yellow dashes, each surrounded by black, lie between
these two last-mentioned lines, the first dash commencing behind
eye, and the last ending above the hindlimbs. himbs slightly
spotted with black and yellow, but no such spots on the dorsal
region which is plain brown, Throat and underparts of body
pure white.”
I thought I had some very interesting variations to report on,
but Mr. Boulenger has described all such in his exhaustive treat-
ment of this species in the Monograph *, and leaves little to be
added.
‘Che number of femoral pores cited in tle Monograph for this
species is 13 to 16, a Nyambita specimen has 12, while the-
* Boulenger, ‘Monograph of the Lacertidw,’ vol. ii. 1921, p. 16.
BAST AFRICAN LIZARDS. 953 |
Kimamba female already mentioned has only 11. In two of
my specimens there are only four labials anterior to the sub-
ocular, which is mentioned as a matter of rare occurrence in the
Monograph.
Pairing takes place in J uly presumably, as large ovules ave to
be found in the ovaries early in August.
Two specimens were found in the stomach of a Heron (Ardea
melanocephala) at Sagayo, but as I did not see this lizard during
a fortnight’s stay at Sagayo, I have not recorded it from this
locality.
LATASTIA LONGICAUDATA Reuss.
Blgv. Cat. Liz, ii. 1887, p. 55.
Thirteen Long-tailed Lizards were taken at Pwaga, Itende,
Iguiwe, Dodoma, Mtali’s, and also many were seen near Rass,
One was also collected at Port Sudan on the voyage out.
A female from Dodoma exceeds my previous records for body-
size, being 1]? inches (100+196 mm.) over all. The series show
a good deal e variation; the occipital is very small and often
Broken up in these specimens.
Parasitic worms were found in the intestines of an Igulwe
specimen.
GASTROPHOLIS virrataA Fisch.
Blgr. Cat. Liz. 11. 1887, p. 7.
A single specimen taken alive at Kilosa on 15.1. 23, I found
it crossing the floor in a room in the Otto House; as ceiling,
walls, and floor were cement and the windows insect-proof, it
was rather a mystery how it gained admittance. ‘There were no
rains on at the time. In captivity it refused al) food for a week,
but drank a good deal of water; it then took two wasp grubs.
A month later it was allowed to escape by the boy in charge
whilst I was away. ‘I'he species is very rare.
HWREMIAS SPEKII Gthr.
Blgr. Cat. Liz. iii. 1887, p. 84.
Native names are Kungulupe in Kinyaturu and Kishagasi in
Kinyaramba.
One hundred and fifty-nine from Itende, Pwaga, Igulwe,
Gwao’s, Suna, Pooma, Mdjengo’s, Mtali’s, Tnaalbal de Ulugn, and
possibly Dodoma, It was common in 2 sort of “ sage-bush ”
scrub at Kidete; seen at Kidai.
This widely-distributed lizard is nowhere scarce where it occurs,
but some idea of its astonishing abundance at Mtali’s may be
gathered from the fact that on my arrival at this place at 11 a.m.
954 Mit. A. LOVERIDGE: NOTES ON
IT gave out that a small reward would -be paid for all lizards
brought in, and by 3 p.m. my boy had paid out for 150. On
learning this I stopped the offer, and in future was more careful
in askiny for specimens, as I had no wish to destroy such numbers
of a useful little creature.
The largest male measured 63 inches (524115), and female 63
(45+ 115).
A careful examination of this long series for Stejneger’s
Hremias sexteniata only revealed four aberrant specimens. ‘Two
of these had the subocular separated from the buceal margin by
the 5th and 6th labials, whilst two others had the subocular
separated on the one side by a Sth, or Sth and 6th labial, whilst
the other side was perfectly normal. The only two specimens taken
at Gwao’s—a male and female—were alike in having but 4 labials
anterior to the subocular on the right side and 5 on the left.
FE. sexteniata was probably founded on two abnormal individuals
‘which, in addition to the subocular being separated from the
mouth, had the central light dorsal line completely divided, making
six dorsal lines in all. ‘The abnormal speciinens referred to above
had only the anterior portion of the central line divided, as is
typical in 1’, spekii.
A female with enlarged ovules measuring 10 x 6 mm. was taken
at Dodoma on 20.xi.21. Many new-born young were seen on
the road from Mpanira to Pwaga (13-15. 11. 23).
Specimens from Suna and Tambali had their stomachs full of
termites.
Two were taken from the stomach of a Hawk (Melierax meta-
bates) at Ulugu, 11. xi. 21, and the pregnant female referred to
above was in the stomach of an Augur Buzzard (Luteo augur) at.
Dodoma, 20, xi. 21.
GuERRHOSAURID.
GrerriosAurus MAgor A. Dum.
Blgr. Cat. Liz. iii. 1887, p. 121.
Seen at Zanzibar, Mt. Kirui, Mkindo River, Kilosa, Ulugu, and
Iuguo. Most of these specimens could have been captured if
desired.
At Mt. Kirui they occur at a great altitude, and as elsewhere,
are associated with rocks, under which they retreat; one caught at
Kipera, however, had is refuge ina hollow tree- ‘ek unk, where it
remained whilst a fierce bush-fire burnt up the surrounding
herbage and blazed for fully five minutes against its retreat, einen
was full of smoke. It emerged none the worse when the fire had
.passed.
My native collector, who has caught several of these lizards,
‘states that he saw one dead at Luguo which had a bright red belly
and was not so coloured from putrefaction.
nr
EAST AFRICAN LIZARDS. 95
GERRMOSAURUS NIGROLINEATUS Hallow.
Blgr. Cat. Liz. iii. 1887, p. 122.
Three specimens available for the present paper, though others
were taken alive or seen at Frere Town, Dar es Salaam, Mbala,
Kilosa, Kidai, and Ikikuyu. This species is not nearly so abundant
at Kilosa as at Morogoro, for which it is diflicult to assign’ a
reason, as the type of country is similar.
In my last notes I assigned these lizards to G. flavigularis
flavigularis, following Sahmnilis ruling of its being the eastern
geographical race, ae he called the renee n G. f. nigrolineatus,
though I pointed out that many of my specimens confor med with
G@. nigrolineatus rather than @. flavigularis. Miss Procter demurs
to my veferring the three specimens under consideration to G.
flavigularis, xnd points out a striking difference not mentioned by
Schinidt. In G@. jflavigularis the laterals are smooth and the
dorsals only feebly keeled, but in G. nigrolineatus the laterals
are keeled and the slonsels so strongly keeled as to form almost
continuous ridges along the back.
A young male was taken on 5. vii. 21 which measured 103 inches
(90+174). Another male, total length 201 inches, had an excep-
tionally long tail 144 inches (145 £365). The tail is therefore
two and a half times the length of the head and body as against
two times as given in the Catalogue.
The points in which this specimen differs from “ flaviqularis”
of the Catalogue are :—Head-shields faintly keeled and pitted.
Froutonasals widely separated from frontal by broad suture of
the pre-frontals. Lateral scales quite distinctly keeled, Femoral
pores 16-15. I might add that there are 22 longitudinal and 58
transverse rows of dorsals. Ventrals in 8 longitudinal Lows.
The coloration of the living lizard is very handsome. General
colour: above nut-brown. A yellow vertebral line, bordered by
black lines, commences two scales behind the parietals and con-
tinues three- quarters of the way along the tail. A series of
black spots along the back between the vertebral and dorsal lines.
Sides seariet, alternate scales being brown and yellow so as, to
form vertical stripes. Under surface pure white.
For several days in succession I had seen this specimen disappear
into a burrow at the base of a bush; it had only come to occupy
the burrow quite recently, as I passed the spot four times daily.
Apparently it had cleaned out the burrow before occupying it,
as my attention was first attracted by the heap, and little slide,
of fresh earth at the entrance. Thinking that 16 might be laying
in the hole, I got my native collector +6 make a snare at the
éntrance of fio burrow which twice caught the occupant, but
it wriggled free. The laborious process of digging it out was.
therefore resorted to, and the burrow found to ‘he over a yard in
length with two smaller blind alleys opening off it.
Grasshopper's and many termites in stomach of an Ikikuyu
male.
956 MR.-A. LOVERIDGE! NOTES ON
‘SciNcIDa:
Masuia comorensis Peters:
Blgy. Cat, Liz. 111, 1887, p. 163.
A single male from Frere 'Town, Kenya Colony, 2. viii. 20,
Whilst collecting butterflies under the mango-trees I heard a
rustle and rush among the dead leaves some ten feet from the trunk
of a mango. Running to the tree I was just in time to put my
hand on a : handsomely- Soloured skink as it ran up the trunk.
It is with some misgivings I refer it to comorensis, as the head
seems broader anid the scalation slightly different from examples
in the British Museum collection.
Its total length was 81 inches (79+ 132 mm.).
The 6th upper labial “was longest and deepest on the right
side of head, and the 5th on the left.
It had 34 scales round mid-body, and agreed in every respect
save coloration with the description given in the Catalogue.
Above greenish-olive ; sides of endl nape, and back sprinkled
with green and black spots. Rims of ‘eyelid bright yellow. A
creamy stripe, black-edged above, below eye. Borders of ear-
opening yellow. An orange side-stripe commencing at ear
vanishes on base of tail ; sides below this pale bluish-grey, belly
gamboge-yellow. Trout bluish, a few black marblings on sides
of throat and base of tail.
MABUIA MACULILABRIS Gray.
Blgr. Cat, Liz, ii. 1887, p, 164.
Four females from Frere Town, Mkindo River, Kipera, and
Jumbe Sunguru’s, near Iilosa ; two were seen at Kidete.
In some specimens the toes of the adpressed hind-limb only
just reach the fingers, instead of reaching to, or nearly as far as,
the elbow.
The Kipera female measured 97 inches (82 +168), and had
seven white eggs in her ovary measuring 14x9inm, In her
stomach was a Small frog !
Masuli PLANIFRONS Peters.
Blgr, Cat. Liz. iii. 1887, p. 167.
Mabuia diesneri Sternfeld, 8.B, Ges. naturf. Berlin, p. 248,
1911.
¥our specimens from Ndogwe, Ulugu, Izikisia, and Tabora.
The Ndogwe individual was found in open Mbugwe country
with scattered thorn-bush. It bolted into a rat-hole at the root
of one of these, from which it was dug out. The Ulugu and
Izikisia specimens were seen on tree-trunks, and sought shelter
in fissures in. the bark or holes in the bole. ‘he Tabora
specimen was found freshly killed in a roadside shelter, Yet
another was seen at Luguo
EAST AFRICAN LIZARDS. 957
The type locality of MZ. planifrons is Taita, Kenya Colony, and
that of Jf. diesneri is Kibwezi, Kenya Colony, not far distant from
Taita. Jf. diesneri has also been recorded from ‘labora; my
Tabora skink agrees with the description of MZ. diesneri with three
exceptions: (1) the first supraocular is not in contact with the
large frontal; (2) the light stripe bordering upper edge of black
lateral band indistinct ; (3) no light patches on back and sides,
The other three specimens whilst agreeing in the main with
M. planifrons have some of the characters of Jf. diesneri, which
was founded on a single small individual. The colouring of the
two species as given in the published descriptions appears
identical, whilst the differences in scalation are not greater than
one meets with in variable skinks such as WZ. striata and M. varia.
I state the differences as given in the descriptions in tabular
form for convenience.
M. planifrons.
Scales in 28-29 rows.
Seales with 3-5 keels.
Frontal in contact with
and 3rd supraoculars.
Ear-opening with 2-3 project-
ing lobules anteriorly.
Adpressed hind-limb does not
reach the elbow.
Olive-brown abeve; two rows of
irregular black spots along
the back.
Light ocelli as in
ocellatus.
A black lateral band beginning
from the eye; on each side
of the back a broad lighter
longitudinal band.
and
Chalcides
M. diesnert.
Seales in 32 rows.
Scales with 3 keels.
Frontal in contact with Ist, 2nd,
and 3rd supraoculars.
Ear-opening with 4 projecting
lobules anteriorly.
Adpressed hind-limb reaches as
far as the elbow.
Brown above, with long rows of
little black spots.
Many feebly-marked light
patches on back and sides.
A deep black band runs along
sides of body, especially dark
on temples, bordered above
by light stripe.
Underparts yellowish - white,
sprinkled brown on sides of
throat. Borders of ear-
opening, labials, and rostral
orange-ycllow.
I tlrerefore propose uniting the two species by enlarging the
description of J. planifrons to read :—
Scales ti 28 to 32 rows.
projecting lobules anteriorly.
Frontal in contact with 2nd and 3rd
or lst, 2nd, and 3rd supraoculars.
Ear-opening with 2 to 4
The adpressed hind-limb may or
may not reach as far as the elbow.
Largest undamaged g measured 7} inches (65+127), largest
Q 12 inches (108+ 195).
The latter’s stomach contained a field
cockroach and what appeared to be the remains of a grasshopper.
958 MR. A. LOVERIDGE : NOTES ON
MaBuia VARIA Peters.
Blgr. Cat. Liz. iii. 1887, p. 202.
Mabuia isselii (Peters), Blgr. Cat. Liz, ii. 1887, p. 201.
Mabuia hildetrandui (Peters), Blgr. Cat. Liz, iii. 1887, p. 207.
Forty-three specimens from Mt. Kenya, Mbala, Mtali’s, and
Shanwa. Also seen at Kidai, Kidete, ixidenge, Itende, Ndogwe,
and Sagayo. It is almost invariably found in association with
rocks, though occasionally seen on, or beneath, logs and stones.
This large series was collected in the hope of throwing hight
on the relationships of varia, isselii, and hildelrandtii. I may
be considered rash in adding the latter to the already long
list. of synonyms of J. varia. ‘Tornier has indeed made isseliz a
synonym of varia in recording a specimen from this region ;
practically isseli’s ouly point of difference from varia is in its
possession of a single frontoparietal caused by the fusion of the
frontoparietals. ‘There jis only one such specimen in the series—
an adult male from Mbala, where it was taken near typical varia,
and what may be considered Aildebrandtii. It has 34 scale-rows.
The reason for zssedii only having 30 is due to the fact that so
few specimens have been collected that the range is not so well
known as in varia and hildebrandtii, in both of which it is 30-34.
Perhaps the feature distinguishing Adldebrandtu trom varia
more than any other is that “ the hind-limb reaches the axilla or
the shoulder of the fore-limb.”
In one Mtali 2 the hind-limb extends considerably beyond
the axilla, ina ¢ from Gwao’s and a @ from Mbala the axilla
ig just reached, ina ¢ from Mbala and many others it just falls
short ; and the variability of this feature, as shown by the series
now before me, demonstrates there is no taxonomic value to be
placed on this character. The amount of contact between the
anterior loreal and first Jabial also varies greatly.
The Mbala specimens have 34 scale-rows at mid-body.
If isselit and hildebrandtii are to be united with varia, the
description of varia would have to be enlarged in some respects,
and would then read :—
— Anterior loreal usually (rarely not) in contact with the 1st labial.
Frontonasal broader or as broad .as long. Frontonasal usually
(sometimes not) in contact with frontal. Hrontal a litile shorter or
as long as the frontoparietals and interparietal together, im contact
with And and 3rd (rarely 1st) supraoculars. Lrontoparietals
distinct or occasionally fused. Parietals usually meeting behind
the interparictal, though sometimes widely separated. Subocular
feebly or strongly narrowed inferiorly, Mar-opening with 2-5
lobules anteriorly, either short-pointed or lony lanceolate. 30-36
scales round mid-body. Ilind-limb reaching to the wrist or shoulder
(or beyond) of the adpressed fore-limb. Subdigital lamelle sharply
tricarinate, occasionally wiricarinate.
_ For the convenience of, anyone interested in the alleged
distinctive characters of the three forms, I have isolated those
characters where all three are not in agreement, and present
them in tabular form :—
EAST AFRICAN LIZARDS.
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‘960 MR. A, LOVERIDGE: NOTES ON
Two females from Mtali’s had each three well-advanced
embryos on 9.x.22, and two others from Shanwa, taken on
20.x.22, had in one case three young very near birth, and in
the other, three, in an early stage. It is curious that the
number of young produced should be so much less than those
recorded in my last paper. There were four embryos in a very
early stage in an Mbala skink on 27, 11. 23.
Sixteen stomachs were examined, and in every case were found
to contain termites, and in only one was any other food present
so far as I could see, and that appeared to be the limb of a
spider.
Minute nematodes in the stomach have been identified as
Physaloptera sp. indet. as 2 9 only.
Mapura opsti Wern.
Wern. Mitt. Nat. Mus, Hamb. xxx. 1913, p. 43.
Two specimens from Kisumu, Kenya Colony. About a mile
south of Port Florence, along the lake shore, these skinks are
quite common, but apparently localized and confined to the rocks
near the shore. Though structurally so similar to A. striata, they
are strikingly different in coloration, being by far the handsomest
Kast African skink I have yet met with. Unfortunately I
made no notes on the exact colour at the time, and nothing is
left of it in the preserved specimens. Speaking from memory
however, only the anterior portion of the body in the adults was
suffused with bright orange; the five longitudinal bands are
bright blue both in the adult and in the young.
In one specimen the parietals are in contact behind the
interparietal, in the other this is not the case. ‘The fronto-
parietals are not smaller than the interparietal as in JZ, striata.
There are five large sharply-pointed, and one rounded lobule,
anterior to the ear-opening. 36 scales round mid-body.
It may be worthy of note that several d/. striata were taken
at the same spot, but on tree-trunks, whilst J/. obsti is obviously
a vock-skink and is the most difficult to catch of any lizard. [
spent a whole day trying to get some, and finally got these two
speciinens from a local native.
MM. obsti was described from Kwa Mtoro, which lies between
Dodoma and Kondoa Irangi in Tanganyika Territory, so that this
record extends its known range considerably, I also saw it on
rocks a mile west of Tgulwe Stn. One was seen at Kidai, five
at Pwaga, and two at Itende, but they successfully evaded
strenuous attempts at capture.
Masuta srrira Petexs.
Blgr. Cat. Liz. iii. 1887, p. 204.
This familiar species is known to the Wanyaturu as Mwin-
I y
yausio, to the Wanyiramba as Tani, and the Swahili as Mjusi
salama.
EAST AFRICAN LIZARDS. 961
Sixteen specimens from Dar es Salaam, Mkata Stn.,
Kimamba Stn., Kilosa, Suna, Gwao’s, Mtali’s, Mkalama, Usshora,
and T'abora in ‘Tanganyika Territory, where it was also seen at
Uliya and Mbala. It was collected at Frere Town, Nairobi,
Kisumu, and Karungu Bay in Kenya Colony.
The largest male measured 9? inches (108 +136) and female
% inches (1094140), from Dar es Salaam and Kilosa re-
spectively. ‘I'wo specimens from Karungu Bay and Kisumu
had 37 mid-body scales, and many of these w
carinate :
Coloxation of a female taken at Nairobi on 12.viii.20.— Rich nut-
brown on upper parts turning to copper on tail, scales frequently
iridescent. Head-shields edged with black. Light stvaw-coloured
stripe commences above the eye, passes along the back
in the dorso-lateral region, and disappears on the base of the
tail. Upper labials and subocular pale straw-colour. Sides
thickly besprinkled with similarly-colored light spots inter-
mixed with black. Throat marbled with brown. Under parts
white. ;
A female taken at Kisumu, 18.xi.22, only measured 100+ 90,
yet had ten young im utero very near the birth. I should think
this is the maximum for this species. A female at Kilosa gave
birth to young on 22. ii. 23.
A lesser Black Stink-Ant (Paltothyreus tarsatus) came
running home with a crumb as large as itself, when a half-grown
skink ran to meet it, seized the crumb, and gave it a terrier-like
shake ; the ant let go and retreated and the lizard dropped the
crumb ; the ant hurried back, seized it, and made a dash for its
hole, the lizard giving chase but failing to catch it, Another
skink was seen to seize a large hawk-moth and dart with it into
a crevice in the wall. Examination of five stomachs gave the
following results: (1.) ants; (ii.) termites ; (ili.) cockroach and
two large wolf (Lycosid) spiders; (iv.) beetle and three bugs
(Lygeus sp.) ; (v.) beetle elytra and millipede rings of 4 mm.
diameter. ‘lhe feces of another held the remains of a green
bottlefly and a moth.
I have already drawn attention to the adaptability of this.
dominant species to all manner of habitats ; its activity is also
remarkable, as the following note made at Kilosa, 8.vi.21, will
show :—‘' A rustle in the roof-gutter and a plop in a bush four
Jeet from the gutter, and I was just in time to see a stri ped skink
recover its balance and run quickly down the stalks and stem of
a bush till it reached the ground, which it crossed with a rush to
the stump of a rubber-tree, up which it darted as if a first-class.
arboreal species.”
A Kavungu Bay specimen had a large grey cattle tick (? Rhipice-
phalus sinus Koch) attached to, and completely blocking up, the
ear-opening. Red acarine parasites were also abundant about.
anus, groin, axilla, and eas
Tlonga,
ere quinque-
962 MR, A, LOVERIDGE; NO'TES ON
MaBuia IRREGULARIS Lonnb.
Lénnb. Arkiv for Zool. Band 14, No. 12, 1922.
Lénnberg has recently described a skink from Soy, Kenya
Colony, as Mabuta (striata ? var.) irregularis n.*%, which is so very
similar to three of my specimens that I do not hesitate in referring
them to it, though the two immature ones are so dried up that
they sre of little use. ‘These three skinks were collected for me
on Mt. Kenya at an altitude between 10,000-11,600 feet by
Dr. J. Arthur. I think the species is quite distinct from
M., striata. he only differences between Linnberg’s description
and my adult skink are (1) parietals are in contact behind inter-
parietal, (2) a single pair of enlarged nuchals.
The largest agrees with AZ, irr equiaris i in differing from typical
M. striata in the following points, though JZ. str iata being such
a variable species, most of, if not all, these variations have been
recorded from time to time :—
(i.) Anterior loreal noé in contact with the first labial.
(ii.) Two pairs of supranasals.
(ul. ) Frontonasal (in its divided state) longer than broad,
(iv.) Frontal shorter than the frontoparietal and inter-
parietal together.
(v.) Pour supraciliaries, second largest.
(vi.) The supraocular is very well separated from lip.
Length of adult 6 inches (69 +83).
LydosoOMA SUNDEVALLI Smith.
Bler, Cat. Liz, ili, 1887, p. 307.
‘Ten specimens from Dar es Salanm, Mkata River, Kilosa, and
Mwanza in langanyika Territory; Nairobi, Kisumu, and Karungu
Bay in Kenya Colony.
The Mkata specimen was taken deep down in a termite heap.
At Kilosa quite a number were taken in snap-~back rat-traps ;
presumably insects went to the bread or cheese bait, and the skinks
followed for the insects. At Karungu Bay and Kisumu these
lizards were living beneath the boulders on the lake shore.
A female 93 inches (130+102) surpassed all my pacyipus
records and was the one taken from the termite heap.
A Nairobi male is much mottled on the under surface, like the
Morogoro specimens referred to in my last collection; it is in
striking contrast to all the other specimens collected this time,
which are pure white beneath.
Four developing ovules (11 x 10 mm.) were taken from a skink
at Kilosa on 1,ii.21 and four from the Mkata specimen on
30. viii. 21. A very young skink (35+ 25) was caught at Karungu
Bay on 16. xi. 22.
LyGosoMA FERRANDI Bler.
Blgr. Ann. del Mus, Civ. (2) xviii, 1898, p. 718.
Kilambamukoot is the Kinyaturu name and Kimilikuta the
Kinyiramba.
EAST AFRICAN LIZARDS.. 963
Ten specimens collected from Mbala, Ikikuyu, Mpanira-kwa-
Sagoi, Lukole, Kidenge, Itende, Kidete, Gwao’s, and Pooma,
and the remains of what Iam confident was a Ferrand’s Skink
was found in the stomach of a Serval (/. serval hinder) at Sagayo.
Most of my specimens were taken beneath logs.
The largest female measured 1% inches (95 +85).
Mbala specimens were coloured unusually; an adult freshly
killed was nut-brown above, each head-scale with a black mark,
and each dorsal seale likewise, those on three central rows
tending to form dorsal lines, which is achieved on 3rd row of
seales from vertebral and forms the top of a dark lateral band.
Lips whitish or yellowish, each upper and some of the lower
lubials bearing a blotch. Under surface of body lemon-yellow,
tail pure white, regenerated portion plumbeous; sides of lower
surface much speckled. yt .
The young specimens are strikingly similar to A. wahlbergi in
colouring except for the spotting on the head and the pinkish
tails, Mxamivation with a Jens, however, shows the markings
to be similar to that of the adult in reality.
Termites were found in the stomachs examined.
One skink was recovered from the stomach of a sand-snake
(P. biseriatus) at Ukikuyu.
ABLEPHARUS WAHLBERGIL Smith.
Blgv, Cat. Liz. iii. 1887, p. 350.
Twenty-five specimens from Dar es Salaam, Mkindo, Mbweni,
Kilosa, Mbala, and Mpanira-kwa-Sagoi. At the latter place the
species 1s very common amongst the Sweepings and vegetable
refuse bordering bush-paths. It is not: uncommon beneath fallen
tree-trunks, and one specimen was taken at the entrance of an
“earth,” down which it was fleeing. j
A. female measuring 4°. inches (48+68). A male with enlarged
testes was in breeding colours, undersides of limbs, tail, and
along both sides of the body being rich salmon-pink (Dar es
Salaam, 4. xii, 22).
The large female referred to above had six developing eggs’ in
ovary measuring 6 xX 3 mm.
MELANOSEPS ATER var. LONGICAUDA Tornier.
Torn. Zool. Jahrb. Syst. xiii. 1900, p. 602.
A single specimen at Mkata Station, where I found it beneath
the ground-sheet of my tent when striking camp. ‘I'he tent-was
pitched in a clearing in the bush, 24. viii. 21. ;
In referring to Melanoseps ater in the Catalogue, I see it
stated “ 3rd upper labial entering orbit,” whilst plate xxxvii.
fig. 16 in the same work does not clearly show this. . M
specimen differs from Tornier’s description in the following
points :—
18 scales round mid-body against 19 in Tornier’s. 130 scales
064 MR.,A. LOVERIDGE; NOTES ON
from chin to anal shield against Tornier’s 118-120. Mine
measured 82+ 30 and his 71441.
Coloration is uniform plumbeous above and below, whilst.
Tornier’s description reads ‘“ Black-brown scales, little light
borders,” and hig second specimen “ Ventral and back scales
black with white borders, back pale olive, two long rows of black
spots close together near middle line” (rough translation).
~The Mkata specimen is a female with two eggs measuring
9x4 mm.
Tornier’s types are from Kerogwe and Masai-land, roughly
from 100 to 200 miles farther north. I do not know of any
other record of this interesting worm-like skink.
Aconvrias MELEAGRIS Gray.
Blgr. Cat. Liz. iii, 1887, p. 427.
As no Acontias has been recorded from East Africa farther
north than Mozambique, I take this opportunity of drawing
attention to a series of this limbless South African skink which
were collected at Voi, Kenya Colony, by Capt. A. B. Percival,
and which are now in the Museum of the Kenya and Uganda
Natural History Society.
hey differ considerably in the matter of colouring, being
slaty-black above and pure white beneath in the half-grown ;
plumbeous above and slightly dusky beneath in the young. Not
agreeing with the colour deseription giver in the Catalogue.
Two specimens brought home for comparison measure 210+ 24
and 146+19 mm. respectively.
CHAMAELEONTIDSA.
CHAMELEON SENEGALENSIS Daud.
Bier. Cat. Liz. iii. 1887, p. 447.
A single female from Kabare, Bukoba, 26. xii. 22, measures
63 inches (92 + 66).
_ CuAMALEON PARVILOBUS Blgr.
Blgr. Cat. Liz. iii. 1887, p. 449.
wo males and a young one from Kilosa, Gwao’s, and Sagayo.
The last two found on bushes in the vicinity of rocky granite
kopjes.
he larger male’ measured 7 inches (95+ 80), the smaller
83-80; the distance between the end of the casque and com-
missure of mouth is 20 mm. and length of mouth 18mm. These
examples were a rich dark green in life. Stomach contents—a
grasshopper.
What appeared to be one of this species was found in the
stomach of a mungoose (7. g. lademanni) shot at Mbulu’s near
Gwao’s. ‘Che Kilosa Chameleon was killed by a young lemur
(G. panganensis) confined in the same large cage.
EAST AFRICAN LIZARDS. 965
CHAMELEON DILEPSIS DILEPSIS Leach,
Bigr. Cat. Liz. iii. 1887, p. 450.
Twenty-six specimens from Kilosa, Tkikuyu, Mt. Hundungula
Itende, Pwaga, Mbala, Mtali’s, and Tabora.
The largest female measured 14 inches (184 +172).
Mating takes place in the January and February (22. ii. 23)
rains. ‘The following note, taken direct from my diary, is of the
pairing of a female whose hind-limbs and part of tail appeared
partially paralysed, therefore what occurred may be somewhat
abnormal,
Kilosa, 19.i.21. There were some four chameleons in my
vivarium, when a native brought mea large female (1604182),
which was introduced into the case at 7.30 a.m.
?
7.45 a.m. I noticed she was on a branch facing a male; he was
raising his occipital flaps from time to time; both had
their throats vertically dilated. She leaned forward and
bit him gently about the neck and head. Pairing I imagine
took place shortly after.
8.15 a.m. As I was passing the case I noticed she was lying
almost on her back except the posterior paxt which was
horizontal. The male chameleon had his head entirely
underneath her or buried in the grass.
Her right fore-leg grasped his right hind-leg.
His right hind-leg grasped her body.
His left hind-leg gripped round her tail, which he kept
working to and fro, not moving himself,
His right fore-leg grasped her side.
His left fore-leg was in the grass.
8.21 a.m. She moved now and then, but except for the move-
ment of his leg, he was motionless.
8.25 a.m. While continuing to grasp his hind-leg she bites feebly
at him, turns slightly and grasps his back on the vertebral
line in the lumbar region with her left fore-leg.
8.27 a.m. Quiescent. Colours strikingly different. The male a
very pale grey-green with broad whitish vertical bands on
sides, ‘The female bright green, One of her hind-legs,
which I suppose is partially paralysed, is doubled back in
a most uncomfortable-looking position.
8.32 a.m. Anterior part of female distinctly on her back; right
fore-leg of female grasping dorsal ridge of male now in
the middle of his back. The male making movements.
8.37 a.m. Lines like leaf-veins appear on sides of male, who is.
inflated and taking deep breaths.
8.40 a.m. Commenced to disengage.
8.41 a.m. Separated, followed by the female excreting.
At this point I had to leave for the office, but was told that.
shortly after I had left the female commenced to follow the male-
about. On my return at 2.10 p.m. courting was in progress,.
Proc. Zoou. Soc.—1923, No. LXIII. 63
966 MR. A. LOVERIDGE: NOTES ON
both clinging to the gauze of the cage. The female gently biting
the male on cheek, the male responded by raising his occipital
flaps, inflating throat enormously, showing skin between scales.
She then followed suit.
Kilosa, 29.i.21. During the past ten days the female has been
following the male about and sometimes vice vers. Paivi ‘ing has
doubtless taken place, though [ have not witnessed it until to- day,
when the female was again lying on her back in the grass.
Kilosa, 1.11.21. The female was apparently dying, and so I
gave chloroform. Subsequent post-mortem showed that the
stomach contained plenty of fresh insect remains, but the intes-
tines were choked with a very hard lump of matter, which on
being broken up was found to be mainly composed of chitinous
parts of insects.
Kilosa, 22.v.21. One of my chameleons on a branch in the
window dropped an egg on the floor between 2 and 2.20 p.m.
This egg measured 12x 9 mm. and, being soft, did not break.
Kilosa, 25.v.21. The chameleon mentioned in my last note
died egg-bound, there being plenty of fresh food in stomach.
There were 46 eggs in the ovary measuring 14 x 9 mm.
Kilosa, iv.-6. v.22. No less than three captive chameleons died
when full of eggs, the highest number being 64. Why they
refuse to lay in captivity when provided with a bush and soil
I do not know.
I found a young chameleon (45+44) on a bush at Tindiga,
14, he Zalle
Two Ikikuyu males were found to have been feeding on ter-
mites. A chameleon was found hanging by one hind-foot from
a branch as if dead. It had disgorged three grasshoppers, one of
which was still kicking feebly, together with a lot of blood and
mucus. It would appear that the chameleon had _ greedily
swallowed his prey without crunching it, and had been killed by
the grasshopper kicking and injuring the stomach with its spiny
lees. However, on taking up the apparently defunct reptile, it
walked off in a dignified manner and ascended its branch.
Mr. Crowther, 5 whom I sent six of these chameleons, writes
me that they were not very keen on beetles and flies, but would
run for woodlice, were very fond of slugs but not much of snails,
whilst mealworms they despised. Their attitude towards flies
may be due to the fact that they were forcibly fed on these
dluring the voyage home.
As elready related elsewhere, chameleons have a very real
enemy in siafu, which ascend the bushes at night. ‘The chameleon,
awakened by the biting little fiends, throws itself from the branch
in an effort to escape, when it is set upon by the driver-ants below,
which do not leave it till all the flesh is picked off its bones.
I lost quite a number in the garden in this way.
It is most extraordinary to what extent a chameleon can hold
on when it so desires. One was released on the balcony, and it
sought freedom by departing on the telephone wire. I saw it
EAST AFRICAN LIZARDS. “967
just before dark clinging in a high wind to the swaying wire at
a distance of 80 feet from the balcony and 20 feet from the
ground. The following morning it finished the remaining
10 feet and descended the iron pole. I might say that I mea-
‘sured the ‘90 feet very carefully lest I should exaggerate in my
enthusiasm for the performance. :
Nematodes from one specimen have been determined doubt-
fully as Physaloptera varani Parona, as only females were found.
CHAMELEON BITENIATUS Fisch.
Blgr. Cat. Liz. iii. 1887, p. 452.
A single female from Kabare, 8.i.23. Measured ie 60mm.
-and held twelve eggs measuring 8mm. in diameter.
CHAMALEON HOEHNELIL Steind.
Steind. Sitz. Ak. Wien, 1891, p. 307, pl. i. fig. 1.
‘Two males from Kabete, Kenya Colony.
Tornier has made hoehnelit a var. of biteniatus*, and Miss
Procter, after examining the two specimens above, writes of
hoehneli :—‘ This appears to me to be a variety of Ch. biteniatus
Fisch. In C. biteniatus there is a strong sexual dimorphism of
the casque, which in the 2 2 is low, andinthed ¢ strongly raised
and crested. ‘The two Kabete males do not differ in any essential
from males from the Aberdare Mtns., K.C., in the British Museum
collection. ‘The gular crest may also be low, or very greatly
developed ; likewise the dorsal, which is always arranged with
every 2ndor 3rd spine the longest. One specimen of Ch. hoehnelii
in the British Museum collection has every 3rd spine remarkably
stoutly developed.”
CHAMELEON JACKSONIL var. VAUERESCECH Tornier.
Torn. Zool. Jahrb. Syst. xix. 1903, p. 176.
Love. P. Z.S. 1920, p. 163.
In my previous note on this three-horned chameleon I re-
marked that only four specimens were available of a large series
collected, many of which remained at Nairobi. Shortly after
I returned to Hast Africa, Miss Procter wrote me that a missing
jar of reptiles which had been sent to the British Museum during
the war had been found in another department. Amongst other
things it contained ten‘of these specimens, to which I referred and
wanted. Three 2 9 were collected dn 27.iii.15 and received
the same register number, viz. 3253. One of these (100 mm.
long) and the male 3534 have already figured in my notes.
Miss Procter points out that in only half of the 9 Q referred to
in iny tabulated list are the preorbital horns as long as the
rostral, and the additional individuals now available contain four
* Tornier, Thier. Deutsch. O.-Afr. Rept. 1896, p. 61.
63*
968 MR. A, LOVERIDGE: NOTES ON
more 9 9 which present an interesting link between forma
typica and var. vauerescece, and nullify my contention that there
is little foundation for saying that the preorbitals are much
shorter than the rostral horn in 9 2, but I think still show it is
impossible to rely on their relative lengths as a guide to sex as
Meek has suggested.
Miss Procter has given me the note she made on the specimens
when they turned up, and [ reproduce this below, together with
the table which she drew up :—
Register Number .......... 3809. 3534. 3543. 2545. 3253. 3253. 3253. 3687.
Sexe eee Sean r ese SING) Dewees 2 g eh eh
Length of head and body. 113 91 81 96 103 100 94 94
Length of rostralhorn .... 25 18 14 17 11 11 6 3
Length of preorbital Represented by mi-
TWAT podoaseod onbapboodponse, CACC) Teens Mh Teh alt 6-6 nute spines 6 mm.
(1 only) long in both speci-
mens as In forma
typica,
“Taking the species as a whole, the rostral horn is the one to
develop first, and is in all cases stouter than the prorbitals.
The prxorbitals were evidently first developed in the g. In
forma typica the g has the three horns of equal length; these
are represented in the Q by three short sharp spines (usually not
more than 1 mm. long). In these Nairobi specimens the horns
develop in the 9 ina highly variable degree. The rostral always
more or less developed; when the rostral is short the preorbitals
are absent as in forma typica; when the rostral horn is long the
preorbitals may be short, medium, or equal in length to the rostral.
In one ¢ of var. vauerescecee the preorbitals are actually longer,
though still more slender, than the rostral, in another they are
shorter asin the 2. Var. vauerescecee therefore differs from forma
typica in the presence of a horn, or horns, of variable degree of
development in the female.”
CHAMELEON MELLERI Gray,
Blgr. Cat. Liz. iii, 1887, p. 472.
Three specimens, ‘Two from Kilosa on 8, vi. & 8.ix. 22, of which
one was on a low cocoanut-palm., ‘he third was a half-grown
individual from Kisanga, taken 24. 11, 23.
As I know of no description of the coloration in life, it may
be as well to give that of one of these females. Snout a rich
dark green; a vertical band of pale yellow from buccal border
below eye extends upwards to postecular region, but not on to
crest. A semicircular patch of pale yellow round forearm
anteriorly. Four vertical bands of a rich dark green on body, first
partly covered by posterior ends of occipital lobes, the second about
mid-body bifurcates before reaching the dorsal crest and each
arm of the fork again bifurcates on the crest, the third and
fourth are similar to the second; the inter-band areas pale yellow
EAST AFRICAN LIZARDS. 969
or yellowish-green. ‘en similar dark green bands on tail with
yellowish interspaces much mottled with green. ‘The tail, which
is generally kept coiled, has a very peculiar appearance, as the
bands coincide, giving the appearance of spokes in a wheel.
This description might be summarised by saying “General
colour a rich dark green, with intermediate more or less complete
vertical bands of yellow from subocular region to tip of tail.”
When angry C. mellert gapes widely, ‘The interior of the
mouth is dark blue-black, corners of mouth blue, inner edges of
lips pink. ‘Two soft lobes of a lung-like appearance on inside of
jaws near the front. The dark green bands become almost black,
light yellow becomes vivid yellow, and the whole of head and
body becomes sprinkled with black blotches, a particularly large
one appearing just above the junction of the forearm.
The coloration of the Kisanga specimen in life was as follows :—
General colour the palest of greens, almost white. A black spot
below the occipital flap. Four vertical black bands on body, the
first cominencing just behind the forearm. ‘Ten similar bands on
the tail, each of the fourteen bands with a greenish-white spoti
dorsally. Limbs with three or more bands sometimes very in-
distinct. Though kept under observation for three months it
never became a rich green (Q 136+153).
At Mbala on 26.ii.23 five eggs measuring 22x17 mm. were
found beneath a stone on a path. ‘There may have been more, as
they had been disturbed by natives when I found them. The
embryos within measured 23-+36 mm. and the hind-limb mea-
sured 16 mm. There were eight distinct vertical bars on the body
and traces of more than eight on the tail.
One specimen on being killed was found to have the intestines
choked by four calculi composed of insect remains.
a
y
i nay
t 4
)
ON SOME NEW OR LITTLE-KNOWN ACARI. 971
47. On some New or Little-known Species of Acari.
By Sranuey Hirst, F.Z.8.
(Submitted for publication by permission of the Trustees of the British Museum.)
[Received September 10, 1923: Read November 6, 1923.]
(Text-figures 1-24.)
TRACHYTES ? AUSTENI, sp. n. (Text-fig. 1.)
Deutonymph. Body about twice as long as wide. Dorsal shield
undivided, covering the entire dorsal surface. It is furnished
with a slight median longitudinal ridge and a lateral ridge on
each side, these three ridges uniting at the anterior end of the
dorsum. Tube of peritreme reaching forwards on to the dorsal
surface above the coxa of the first leg, and then bending back
again to form a short loop. There are a number of small round
platelets, each bearing a hair situated laterally on the soft
integument between dorsum and venter. A pair of similar
platelets is also present on the little triangular cone in which the
body terminates anteriorly, ‘The ventral plates are not quite
like those of 7’. swmatrensis Zacher; the large anterior one
(called sterni-metasterni-genitale by Zacher) has the lateral
projection on each side opposite the interval between the second
and third cox more angular than depicted by Zacher; whilst
the anal plate is larger than in 7’. sumatrensis, almost touching
the large anterior plate, and sometimes quite in contact with the
coxal (metapodial), the latter being very: well-developed. All
the ventral plates are ornamented with minute spots, each
composed of a group of punctations. Anterior ventral plate:
with five pairs of very short hairs. Anal plate with seven pairs
of hairs, including two pairs situated near the anal aperture.
Posteriorly there are several of the minute circular platelets,
each carrying a short hair.
Length (including palpi) °48 mm.; width :22 mm.
Host: A large fly (Pantophthalmus tabaninus Thunb.); Brazil.
L&LAPS UGANDANUS, sp. n.
Q. A large and very strongly chitinized species. Sternal plate
very like that of Z. muricola Traigirdh and L. giganteus Berlese ;
faint reticular markings are present on its surface. Glenito-
ventral plate widely expanded behind the cox as in L. muricola,
etc.; apparently there are no linear markings on it. Lateral
platelet near the last coxw much larger than is usually the case in
the genus Lelaps and curiously shaped, ending in a sharp, almost
spiniform point anteriorly; its surface is ornamented with a
rather irregular network of linear markings. Paired hairs on
anal plate slightly longer and also more slender than the unpaired
972 MR, SPANLEY HIRST ON
posterior hair. Hairs on venter numerous, most of them being
short, but the lateral and posterior ones are somewhat longer ;
there are three pairs of quite long and fine hairs to the side of
and behind the posterior end of the genito-ventral plate. Coxa
Text-figure 1.
TERZI—~
i}
Trachytes austeni, sp.n. Deutonymph. Ventral view.
of first leg with a short outer spine and a long inner seta, neither
being stout. The spine or thorn on the second and third coxx
is not very thick ; whilst that of the fourth coxa is short and
quite slender.
NEW OR LITTLE-KNOWN ACARI 973
Measurements. Length of body (not including capitulum)
1820p; its width 1290. Length of sternal piate 440; its
greatest width 420 p, its least width 380. Length of genito-
ventral plate (including the operculum) about 760; its greatest
width 590 ». Length of lateral platelet 195 w; its width 142 p.
ITab. Off a rodent, Bugwe, Bumungi, Uganda. Collected by
W.N. van Someren (20. iv. 1922).
LIPONYSSUS GORDONENSIS, sp. n. (Text-figs. 2 & 3.)
Q. A single dorsal shield is present, being shaped as shown in
text-figure 2. On the surface of this shield there are a number
of very short fine hairs. In the anterior half these hairs are
grouped together, but posteriorly they are fewer in number,
apparently always being paired; the paired hairs at the extreme
Text-figure 2.
c
Liponyssus gordonensis, sp. n., 2. Dorsal view.
posterior end of the dorsal shield are much longer than the
others. Hairs on softer uncovered part of dorsum not very long,
but much longer than those on the dorsal shield (except the
terminal pair on the latter), and they are curved as in LZ. saurarum,
ete. Sternal plate trapezoidal and only furnished with two pairs
974 . MR, SEANLEY HIRST ON
of hairs; its posterior margin is not thickened. Genito-ventral
plate narrow and its posterior end pointed. Anal plate pear-
shaped and not so narrow as that of LZ. sawrarwm. Hairs on
venter not very numerous, and much shorter and straighter than
those on the dorsum. Peritreme extending as far forwards as the:
coxa of the second leg. Coxe of legs without spurs. Legs.
Text-figure 3.
Liponyssus gordonensis, sp.n., 2. Wentral view.
* ) 2
furnished with numerous hairs, which are mostly slender, those
on. the ventral surface being very fine. There is a pair of some-
what thicker hairs or set on the dorsal surface of the femur of
the first leg. Chelicera normal for the genus, its fingers not.
heing provided with teeth.
_ Length (including palpi) ‘68-95 mm.; width -35--58 mm.
NEW OR LITTLE-KNOWN ACARI. 975:
Host: A skink (Mabuia quinquestriata Lichtenstein) ; numerous.
specimens collected by the author on this host at Khartoum
(Feb. 1923).
Lironyssus SYLVIARUM Can. & Fanz.
Miss E. Knight, of the Lister Institute, has sent me a number
of specimens of LZ. sylviarwm found on hens at Bletsoe, Bedford-
shire. ‘The mites occur round the vents of the birds, and seem to:
multiply very rapidly. ‘They form colonies, and do not leave the
birds during the daytime, as is the habit of the common red
mite of fowls (Dermanyssus galline Redi). The tail-feathers,,
particularly the fluffy portion near the base, become covered with
great numbers of mites. Later on, the parts of the skin attacked
by the mites become raw, and a hard scab appears afterwards.
Although previously found on Pigeons in this country, this is the
first case of this mite becoming a pest of poultry here. In the
United States a number of cases of L. sylviarum attacking hens.
are known, and it is regarded as a very injurious species, egg
production of the infested birds being greatly reduced.
DERMANYsSUS (ALLODERMANYSSUS) SANGUINEUS Hirst.
This species was described from specimens found on Laitus
rattus and other rodents in Egypt. It has also been recorded by
Ewing as occurring in the United States (Proc, U.S. Nat. Mus.
1922, vol. 62, art. 13, p.25). During February 1923, 1 collected
a number of specimens of this mite from Rattus alewandrinus
grey-bellied form) and from J/us musculus gentilis at Khartoum,
Sudan.
Ruunowyssus (NroNYssoIDES) NUCIFRAGA, sp. n. (‘Text-figs.
4 & 5.)
Q. Two large dorsal shields are present, covering practically
the entire upper surface of the body instead of only a single
anterior dorsal shield as in most species of Mhinonyssus.
Apparently only one or two pairs of exceedingly short and
inconspicuous hairs are visible on the dorsal shields; there are,
however, also some very minute and inconspicuous circles on the
shields ; these minute circles no doubt represent the sockets of
the hairs present in other forms. Hairs on venter also mostly
obsolete or absent ; when present they are exceedingly short and
inconspicuous. Sternal plate indistinct. There is a clear space,
ventrally which probably represents the genito-ventral plate; it.
is wide and apparently rounded off posteriorly. Anal plate
distinct and pear-shaped, bearing three very short hairs.
Peritreme very short. Coxe without spines or hairs, only minute
circles being present. A few very short spinules occur on the.
proximal segments of the legs, and some longer fine hairs on their
976 _ MR, STANLEY HIRST ON
tarsi. There is also a fine hair on the dorsal surface of the
penultimate segment of the anterior legs, and a fairly long and
fine but rather stiff hair, which has its basal ring rather strong
and conspicuous, is placed dorsally near the distal end of the
tarsi.
3. Genital pore anterior in position. Anal plate apparently
narrower than in the female sex.
Measurements. @: length (including palpi) -66-70 mm.;
width ‘39-40 mm. 6 : length (including palpi) °66 mm.; width
*30 mm.
Host: Nucifraga caryocatactes; three female specimens
collected by Herr ‘Tischler at Heilsberg (5.ix. 1913) and a male
from Ulmenhorst (12. x.1911); Thienemann Coll.
Text-figure 4.
Rhinonyssus (Neonyssoides) nucifrage, sp. n., 2. Worsal view.
Norr.—Owing to the presence of two dorsal shields instead
of a single anterior one, as is usually the case in Lhinonyssus, it
is necessary to place this species ina new subgenus— NVeonyssordes.
Rhinonyssus (Neonyssoides) nucifrage resembles Neonyssus
intermedius Hirst in having two dorsal shields and also most of
the hairs replaced by minute circles, differing from the Jatter in
the shape of the ventral plates, presence of distinct spinules on
the legs, etc.
NEW OR LITTLE-KNOWN ACARI.
Text-figure 5.
Rhinonyssus (Neonyssoides) nucifraga, sp.n., 9. Ventral view.
Key to the species of Ancystropus present in the
British Museum Collection.
haeeii of legs obsolete or absent .........sccceeee00, 2.
977
Pulvillus of legs well-developed...........::::ss0eee0 8, (Subgenus Heristaspis
(Sternal plate much wider than long; the two rows
‘ of dentiform processes on posterior surface of first
leg strongly developed ............:.ceceeceereceees Ae zelebort Kolenati.
‘ | Sternal plate only slightly wider than long; the
two rows of processes on posterior surface of first
L leg much weaker 0.0.0... eeeieeereee A. @thiopicus, sp. n.P
Posterior spurs of coxse obsolete. Hypostome short, :
rather stout, the barbules unusually large ......... A, macroglossi, sp. n.
2
3.
eee spurs of cox better developed. Hypo-
Kxolenati.)
stome shaped otherwise ........... .....000e 4.
Posterior spur of first coxa long..........0cceeeeeee As calcaratus, sp. ni.
4.
Posterior spur of first coxa much weaker ....,.......... A. lateralis Kolenati.
‘978 MR. STANLEY HIRST ON
Ancystropus ZELEBORI Kolenati. (Text-fig. 6.)
Ancystropus zeleborit Kolenati, Parasiten der Chiropteren,
1856, p. 25.
Q. Dorsal shield not very long, and its posterior end is wider
and blunter than in A. lateralis, ete. Peritreme very short, being
restricted to the dorsal surface. There is a very strongly
Text-figure 6.
\-
4 4
1
-* ‘i
Je y ;
ty } \ {
7G) ) \ ' f f
& \ t
\ i
\ t
=a
TERZID Wy Kk
5 /
7
4 “ al ae ii
PRE aL Berard ER ON lo - ¥
\ " a q /
Ancystropus zelebori Kolenati, 2. Ventral view.
chitinized structure at the base of the capitulum ventrally which
seems identical with the jugular plaque, but it is coalesced
laterally with the capitulum instead of being free; it has
distinct and rather wide auricule posteriorly much as in some
ticks. Sternal plate situated far forwards, being placed just
behind the base of the capitulum ; it is shorter than in the other
known species of Ancystropus, being much wider than long.
NEW OR LITTLE-KNOWN ACARI. 979
Process on dorsal surface of capitulwm well-developed and conical.
Hypostome rather long and slender. First leg greatly enlarged ;
its claws very large; the pulvillus very much reduced in size, being
obsolete. Lateral sete of this leg shorter than in A. lateralis ;
there are two rows of strong backwardly-directed denticles on the
posterior and postero-ventral surfaces of the first leg, also a few
projections on the anterior (inner) surface, and one or two rather
strong ventral tooth-like projections. First coxa with the spur
on its posterior margin poorly developed; second and third cox
each with a slight lobe-like spur posteriorly, these spurs being
weaker than in A. lateralis and A. calcaratus.
Length of body (not including capitulum) °73 mm.; its width
“D1 mm.
Host: Kolenati states that either Rhinopoma microphyllum or
else Pieropus cegyptiacus is the host of this species of mite. I
have only seen a single example, ex Kolenati’s collection. —
ANCYSTROPUS ZTHIOPICUS, sp.n.? (Text-figs. 7 & 10a & c.)
Q. Apparently the dorsum is not divided into two portions by
a transverse line in this species. Dorsal shield (scutum) shaped
as shown in text-fig. 7 (the dotted line). Hairs on anterior
margin of dorsum very short. The pair of hdirs present just
behind the posterior end of the scutum in A. lateralis is missing
in A. ethiopicus. Sternal shield larger than in A. zelebori
Kolenati, and its shape is quite different. Process on dorsal
surface of capitulum conical.
The strongly chitinized internal portion of the jugular plate
varies considerably in shape in specimens of this mite from
different localities; the posterior margin of this internal part
is strongly concave in the examples from Damba Island, Lake
Victoria, the posterior angles (auricule) being large and strongly
produced ; whereas in specimens from the Gambia the posterior
margin is almost straight, the auricule being practically absent ;
the typical specimen from Zanzibar is rather intermediate in this
respect, the posterior margin of this part of the plaque being
somewhat concave with well-developed auricule. Mypostome
slender and fairly long. Palp rather short and fairly stout.
First leg much enlarged as in A. zelebori, but the backwardly-
directed little chitinous processes on the posterior side of this
limb are weaker than in that species; those of the lower row are
sometimes distinct and dentiform, being especially well-developed
in examples from Damba Island, Lake Victoria; but those of the
upper row are quite weak; one or two of the processes on the
anterior surface of this leg are large, however. Claws of first leg
apparently rather smaller than in A. zelebori; the pulvillus
obsolete. Posterior spur of first, coxa obsolete; second and third
cox each with a distinct lobe-like posterior spur; a weak
pointed posterior spur may also be present on the fourth coxa.
Hairs on dorsal surface of legs shorter and finer than in A. lateralis
and also smoother, the accessory hairlets being obsolete.
980 MR, STANLEY HIRST ON
Length of body (not including capitulum) ‘94 mm. ; its width
°65 mm.
Host: A fruit-bat (Zpomophorus minor) from Zanzibar; a
single specimen (the type) from this host. Also specimens
probably referable to the same species of mite from the following
hosts and localities :—AJicropteropus pusillus, Gambia (Dr. P.
Rendall). From a fruit-bat, Damba Island, Lake Victoria
(Dr. G. D. H. Carpenter).
Text-figure 7.
a AY
oh BS
Ancystropus ethiopicus, sp.n.? 2. Ventral view.
Norr.—It is possible that Kolenati’s A. mulleri from Pteropus:
vulgaris (= Pteropus niger?) is allied to the species described
above under the name 4d. ethiopicus or perhaps even identical
with it. The bat, Pteropus vulgaris, however, only occurs in the.
Mascarenes, Réunion, and Mauritius,
NEW OR LITTLE-KNOWN ACARI. 981
ANCYSTROPUS (Merisrasris) MACROGLOSSI, sp. n. (Text-figs. 8,
9, & 100.)
2. Dorsal surface not divided into two divisions, but there isa,
short transverse line situated just behind the pointed posterior
end of the scutam; the latter is shaped very much like that of
A. lateralis. Number and distribution of hairs on dorsal surface
Text-figure 8.
Ancystropus (Meristaspis) macroglossi, sp. n., 9. Ventral view.
exactly the same asin A. laterulis. Jugular plaque well-developed
and transversely elongated, being even wider than in A. lateralis.
Sternal plate smooth and shaped as shown in text-fig.8; there are
- three pairs of hairs on its surface and also the usual two pairs of
minute punctations. Genital operculum situated just behind the
sternal plate, its anterior margin overlapping the hinder margin
Proc. Zoot. Soc.—1923, No. LXIV. 64
982 MR. STANLEY HIRST ON
of the latter. Genito-ventral plate very similar to that of
A. lateralis but wider, and furnished with a pair of hairs as in
that species. Posteriorly there is also a somewhat triangular
smooth area of the integument. Hairs on venter not numerous.
There is a pair of hairs or sete on each side of the posterior end
of the body as in 4A. lateralis, but they are longer than in that
species. Process on dorsal surface of capitulum stout and conical.
Hypostome rather short, comparatively thick, and with the
terminal barbules exceptionally well-developed. First (proximal)
Text-figure 9.
THRE)
Ancystropus (Meristaspis) macroglossi, sp. n., 9. Palpi and hypostome,
greatly enlarged.
segment of palp with the outer angle projecting and almost tooth-
like. First leg thickened much as in A. lateralis, and furnished
with rather similar lateral sete and a pair of paddle-shaped
terminal setee as in that species. The chitinous processes present
on the posterior surface of the first leg in A. lateralis are obsolete
in this new species, and those usually present on the inner surface
are also obsolete or absent; the anterior edge of the ventral
surface of each of the segments of this leg is concave, however,
NEW OR LITTLE-KNOWN ACARI, 983.
and produced laterally so as to be almost dentate. Claw of first
leg small ; the pulvillus present. Some of thé hairs on the dorsal
surface of the legs are long. Hairs on ventral surface of legs
very few in number. First coxa apparently without any
posterior spur; the lobe-like spurs on the posterior margins of
Text-figure 10.
a. Jugular plate of Ancystropus ethiopicus (specimen from Zanzibar).
b. Jugular plate of Ancystropus macroglossi.
ce. Jugular plate of Ancystropus ethiopicus (specimen from Damba Island,
Lake Victoria).
the second and third cox are very slight, being almost imper-
ceptible. oa
Length of body (not including capitulum) -63 mm.; its width
-D4 mm.
Host: Macroglossus minimus; Gilolo.
We have only a single specimen of this new mite in the
collection.
ANCYSTROPUS (MERISTASPIS) CALCARATUS, sp. n. (Text-fig. 11.)
@. The whole dorsal surface is rather strongly chitinized, and
it is difficult therefore to see the outline of the scutum itself.
There are a number of shallow pits on the surface of the scutum.
Chetotaxy of dorsum much as in A. lateralis, but the hairs seem
shorter. Process on dorsal surface of capitulum rounded instead
of conical as in A. lateralis, etc. Jugular plaque strongly
chitinized and elongated transversely, being much wider than
long. ternal plate almost flask-shaped, for it is roughly oval in
shape, with a short anterior neck-like portion: It is smooth,
being without markings, except for two pairs of minute punc-
tations, and there are three pairs of hairs on it. Gentto-ventral
plate very short, rounded off posteriorly and bearing a pair of
hairs. Hairs on vente very sparse as in A. lateralis, and although
‘quite short they are longer than in that species. In unmounted
specimens there is sometimes a slight but noticeable swelling on
each side of the venter just before the posterior end of the body.
Paired spines or hairs at posterior end of body minute and
inconspicuous, being much smaller than in P. lateralis. First leg
enlarged; the lateral sete are very like those of A. lateralis, and
there is a términal pair of paddle-shaped sete as in that species.
64*
984 MR. STANLEY HIRST ON —
Denticles on postero-ventral surface of this leg also very like
those of 4. lateralis, but stronger, the anterior margin of the
ventral surface of the segments of this leg being strongly concave.
There is a stout spinule ventrally on each side of the distal end
of the first tarsus, close to the origin of the claws. Claws of first
leg not very large; the pulvillus present. Spur on coxa of first
leg much longer than in the other known species of the genus.
Second and third cox each with a lobe-like posterior spur.
Text-figure 11.
Ancystropus (Meristaspis) calcaratus, sp... 2. Ventral view.
There are a number of rather long hairs on the dorsal surface of
the legs; a pair on the femur of the first leg is the longest, the
outer hair being the longer of the two.
Length of body (not including capitulum) °70—about °94 mm.;
its width -53--6] mm.
Host: A large flying-fox (Pteropus sp.); Rook Island, August.
1913 (4. S. dAfeek), ex Hon. N. C. Rothschild’s coll.
NEW OR LITTLE-KNOWN ACARI. 985
Ancystropus (MERIsTASPIS) LATERALIS Kolenati. (Text-figs.
12 & 13.) .
Pteropus lateralis Kolenati, Pavasiten der Chiropteren, 1856,
p. 29.
Meristaspis lateralis Kolenati, Sitzb. K. Ak. Wien, 1858, xxxiii.
p. 84.
2. Dorsum divided into a large anterior and a_ shorter
posterior portion by a well-defined transverse line, situated just
above and behind the last pair of legs. Dorsal shield ill-defined,
Text-figure 12.
A ©
ey wae A
Ancystropus (Meristaspis) lateralis Kolenati, 2. Ventral view.
but shaped as shown in text-fig. 12, the posterior end being
rather sharply pointed. This shield reaches backwards as far as
the transverse line. Hairs on dorsum short and few in number;
there are several hairs near the anterior and antero-lateral
margin, viz. a median pair and three hairs on each side a little
further back, also a hair on each side opposite the gap between
the third and fourth legs. On each side of the pointed posterior
end of the scutum there is a short hair. There is also a pair of
986 |, MR. STANLEY HIRST ON
very short hairs a little behind the middle of the posterior part:
of the dorsum. Apparently there are no hairs on the scutum
itself, only minute punctations. Process on dorsal surface of
capitulum conical, pointed, and backwardly directed.
.. Jugular plaque well-developed, being strongly chitinized, much
wider than long, and practically rectangular. Sternal plate
shaped as shown in figure ; it is very like that of some species of
Spinturniz, the anterior end being narrowed ; this plate has a
narrow marginal strip of paler and weaker chitin ; the surface is
smooth, being without any markings, except two pairs of minute
Text-figure 13.
fe)
\
d]
Ancystropus (Meristaspis) lateralis Kolenati, g. Ventral view.
punctations; there are three pairs of marginal hairs. Hairs on
venter very few in number; there is a pair on the hinder margin
of the minute genito-ventral plate. Behind this plate on the rest of
the venter there are only four or five pairs of very short hairs, and
they are well separated from one another. Part of the venter is
very smooth, being devoid of the usual linear markings, and
perhaps this area represents an obsolete plate. On each side at
the posterior end of the body there is a pair of characteristic
rather stout stiff sete, Hypostome slender, the terminal barbules
NEW OR LITTLE-KNOWN ACARI. ‘987
fairly well-developed. First pair’ of legs greatly enlarged, much
as in A. zelebori; the lateral sete of this leg much better
developed than in that species, being rather stout and projecting
sideways and slightly forwards; on each side of the distal end of
this leg there is a paddle-shaped seta, its tip being flattened and
truncated. At. the base of each of the lateral sete there is a
curious projection ending both proximally and distally in an
angular process or tooth, partly formed or at least accentuated
by the concave distal margin of the ventral surface of the
segment. Similar projections may occur also on the inner
surface of the first leg. The upper lateral row of denticles
present in A. zelebori seems to be absent in the present species
and also in A. calcaratus, sp.n. Claws of first leg not very
large; pulvillus present. Coxe 1-3 each with a lobe-like spur
on the posterior margin. A pair of very long hairs is present
dorsally on the femur of the first leg and another shorter pair on
the patella; similar but shorter hairs are also present on the
same segments of the second and third legs; the trochanter of
the third leg also has a rather long dorsal hair. A rather long
dorsal hair is present also on the trochanter, femur, patella, and |
tibia of the fourth leg. All these long hairs have very minute
accessory spinules or hairlets, causing them to have a slightly
roughened appearance. Lateral hairs of legs stiff, plain, and not.
very long.
3. Scutwm covering the entire dorsal surface. Jugular plaque
with the posterior margin sumewhat concave. Sternal plate very
like that of the female, but with the usual anterior genital pore.
The pair of hairs at the distal end of the tarsus of the first leg
are not paddle-shaped, their tips being finely pointed. Spurs on
coxee not so well-developed as in the female.
Q. Length of body (not including capitulum) -70—"79 mm. ;
its width 53-56 mm.
3. Length of body (not including capitulum) *50 mm.
Host: A fruit-bat (Rousettus egyptiacus). Ihave examined the
cotype deposited in the Brit. Mus. Collection by Kolenati; also
specimens from Rehoboth and Jaffa, Palestine, Jan. 1912 and
Feb. 1920 (7. Aharoni), and examples from Cyprus, collected by
Miss D. M. A. Bate. "|
PERIGLISCHRUS INTERRUPTUS Kolenati. (Text-figs. 14 & 15.)
Pieroptus interruptus Kolenati, Die Parasiten der Chiropteren,
1856, p. 29; LPteroptus hipposideros Kolenati, ¢. c. p. 29;
Periglischrus interruptus Kolenati, Sitzb. K. Ak. Wiss. Wien,
1858, xxxiii. p. 80; Periglischrus hipposideros Kolenati, t. c.
p. 82; Periglischrus glutinimargo Kolenati?, t.c. p. 80.
@. A small species with the posterior end peculiar in shape,
being wider than the rest of the body and flattened dorso-
ventrally, forming a kind of rim. The weak sternal plate
apparently has no hairs on its surface, but there are three pairs
‘988 - MR. STANLEY HIRST ON
of minute circles on it near the lateral margins, and sometimes
also two additional pairs of rather indistinct circles situated
nearer the middle of the plate. Genital operculum very small and
situated between the third coxe. Genito-ventral plate very small,
narrow and wedge-shaped in outline, the posterior end being
pointed. Peritrveme very short. Hairs on venter few in number
and exceedingly minute and inconspicuous. There is a pair just
behind the posterior end of the genito-ventral plate and also
three pairs at the posterior end of the venter. here are also a
Text-figure 14.
Periglischrus interruptus Kolenati, 2. Ventral view.
a. End of tarsus of leg of same showing pulvillus.
number of minute circles which do not seem to be provided with
hairs. Coxe of legs without spurs; they are rather curiously
shaped (see text-fig. 14), the outer posterior angle of the third
coxa being strongly chitinized and projecting somewhat, whilst
the same part of the fourth coxa forms a long narrow strip.
Numerous hairs and sete of moderate length are present on the
dorsal surface of the legs, including a noticeably long hair on the
femur and another on the patella of the first leg. Ventral
surface of legs almost hairless, except the tarsi which bear a
number of hairs. Pulvillus of legs peculiar in shape; there isa
\
NEW OR LITLLE-KNOWN ACARI. ‘989
well-developed additional lobe or expansion on each side of the
basal support of the claws, besides the usual terminal lobes
(text-fig. 14a).
3. Sternal plate shaped as shown in figure; there are two
pairs of minute circles situated near the lateral margins
posteriorly. Hairs on venter either short or replaced by minute
circles. Coxe without spurs. The two modified sete placed
near the distal end of the penultimate segment of the palp ar
quite long. i
Text-figure 15.
Periglischrus interruptus Kolenati, g. Ventral view.
Hosts and localities. Kolenati gives Rhinolophus clivosus
[= R, blasii] as the host of Periglischrus interruptus in_his
*Parasiten der Chiropteren,’ but later in Sitzb. Ak. Wiss. Wien
gives R. euryale, allotting a new name—P. glutinimargo—to a
mite of this genus from Egyptian specimens of A, clivosus. In
addition to Kolenati’s cotypes of P. interruptus and P. hippo-
sideros, the British Museum possesses examples from hinolophus
990 . MR, STANLEY HIRST oN
euryale, Grotto of Pietralbello, Pontelecchia, Corsica (collected by
S. Hirst, 17. iii. 1922), and from the same host at Ajaccio:
(S. Hirst, 5. iii, 1922).
_ PaRATETRANYCHUS INDICUS, sp. n. (Text-fig. 16.)
‘g. Palp with the terminal finger (A in text-fig, 16a) stout,
but rather short, being shorter than the stiff rod-like sete
situated near it. Dorsal sensory finger (B in text-fig. 16 a) club-
shaped and akout as long as the terminal finger. When seen
from the side, the ventral part of the claws of the anterior legs
seems to be a strong curved claw-like process, but if examined
earefully, it is seen to be subdivided into several parallel teeth,
which are rather stout in the case of the first leg, those of the
second leg being weaker. Ventral part of claw of posterior legs.
Text-figure 16.
Paratetranychus indicus, sp. n.
a. Palp of male.. 6 & c. End of collar trachea showing variation in shape. d. Palp:
of female. e. Claw of posterior leg of male. jf. Penis. g & h. Claws of
anterior legs of male.
divided into six quite fine hairs as in all the legs of the female
sex; the dorsal claw-like part is longer than in the anterior legs.
The hard chitinized part of the penis isshort and strongly curved,
its end being shaped rather like that of 7’. telarius (= 7. althew
of Hanstein, Zacher, and some other recent authors), one of the
barbs being quite weak.
9. Body elongate-oval, the body+the head-plate being about
twice the greatest width of the former. Terminal finger of palp
(labelled A in figure d) of female short and very wide. Collar
trachea normally ending in a single elongated terminal cell
(text-fig. 160); in one specimen, however, the end consists of
two cells lying side by side, one being more elongated than
the other (text-fig. 16c). Claws of Jegs of female similar to
the posterior ones of the male, consisting of a long slender
NEW OR LITTLE-KNOWN ACARI. 991
ae part and a ventral projecting portion split into six fine
airs.
Measurements. 3 : length (including palpi) 33mm. ; greatest
width ‘147mm. 9: length (including palpi) 51-54 mm.;
greatest width :235 mm.
Host-plant: Sorghum, India. Mr. Y. Ramachandra Rao, of
the Coimbatore Agricultural College, informs me that this mite is
sometimes a serious pest of Sorghum or cholam (Andropogon
sorghum). It is found in large colonies under the surface of the
leaves and increases rapidly in numbers. The attacked portions
turn bright red as if attacked byrust. The mite is greyish green
in colour when alive. :
TEPRANYCHUS CRATHGI Hirst.
I. Zacher asserts (Mitt. Biol. Reichs. f. Land-u. Forst. Berlin,
Heft 21, Dec. 1921, p. 91) that the species of Red Spider described
by myself under the name Vetranychus crategi is a synonym of
his V'etranychus viennensis. ‘Vhe description of 7. crategi was
published in the Proc. Zool. Soc. July 1920, whereas that of
T. wennensis was published in Zeitschr. angew. Kntom. Berlin,
vii. No. 1, September 1920. 16 will be seen that 7. crategi has
the priority and that 7’, viennensis is the synonym. It is true
that Zacher had circulated an earlier typewritten document
which does not bear the impression or stamp of any Society or
publisher, nor even the name of the place at which it was issued.
Tt bears the date “17. Mai 1920” and is entitled “ Vorliiutige
Diagnosen einiger neuer Spinnmilbenarten.” There are several
corrections in pencil in the copy in my possession. It is quite
evident that this document does not constitute publication and
that the three new species mentioned in it are not valid ; in fact,
it would seem that Zacher realises that this is so himself, for he
redescribes all three species as new at « later date in Zeitschr.
angew. Entom. Berlin, as mentioned above.
Anycuus Latus Can. & Fanz.? (Text-figs. 17 & 18.)
3. Collar trachee of a simple type (see text-fig. 17 d), ending
in a single long cell. Hairs or sete on dorsum twenty-six in
number (not including the finer hairs near the anus) ; these dorsal
hairs are stiff, being rod-like and slightly pectinated. Penis
shaped as shown in figure, being strongly curved and claw-like.
Palp with the termina! sensory finger represented by a conical
spine, which is only about half the size of the two usual stiff red-
like sete present close to it; dorsal sensory finger short and
slender.
Tarsi of legs apparently without any claw, but the usual tenent
hairs, which are four in number in the male of this species, are
present, being situated on ashort peduncle. Besides the ordinary
rather long and slightly-feathered hairs on the legs, there are a
992, MR. STANLEY HIRST ON
few modified and sensory hairs. Dorsally near the apex of the
tarsi there is a curved hair or seta with the end enlarged, being
flattened and pectinated. ‘There appears to be only one of these
hairs on the anterior tarsi, but there are two on the third tarsus.
There are several short, stiff curved rod-like sensory hairs
(sinneskolben) at the distal end of the anterior tarsi; four of
these sensory sete are also present on the penultimate segment
of the first leg, two dorso-lateral ones posteriorly and another
anteriorly, and one ventral in position. Besides the short distal
sensory set# on the second tarsus, there is also one rather near
‘the proximal end; whilst there are two on the penultimate
segment of this leg. Tarsus and also the penultimate segment of
the posterior legs each usually with a short, stiff sensory seta near
the proximal end, but that on the penultimate segment of the
fourth leg may be absent.
Text-figure 17,
Anychus latus Can. & Fanz.?
a. Penis. 6. Palp of female. c. Palp of male. d. End of collar trachea.
e&f. Penis. g. End of a leg.
2. Terminal sensory finger of palp fairly well-developed, being
peg-shaped and almost as long as the longer of the two rod-like
Setie situated close to it. Dorsal sensory finger not very long, but
slender and practically cylindrical.
3. Length of body (including palpi) °375 mm.; its greatest
width -192 mm.
2. Length of body (including palpi) -437 mm.; its greatest
width °304 mm.
Plant-hosts: Lebbek-trees at Wadi Halfa ; specimens collected
by 8S. Hirst (25.1.1923). Also specimens from a leguminous
tree or shrub at Makwar, near Sennar, Biue Nile, Sudan
(9.11. 1923), collected by S. Hirst.
Norre.—The finer structural details are not all given in
Canestrini’s and Berlese’s figures of A. latus, so that I have some
doubt in referring my specimens from North Africa to this
NEW OR LITYLE-KNOWN ACARI. 993
species, but I think that this identification will prove to be
correct. It is possibly also the same as the mite mentioned by
F. C. Willeocks under the name Bryobia sp. (Lebbek Red Spider).
{ lee Highley-det
yy, Sy
Anychus latus Can. & Fanz., Sg. Dorsal view.
in bis “Survey of the More Important Economic Insects and
Mites of Egypt,” Bulletin No. 1, Sultanic Agricultural Society,
p. 266 (1922).
094 “MR. STANLEY HIRST ON
, DEMODEX SCIURINUS, sp. n. (Text-fig. 19.)
- Variety of D. melesinus ?
2. Body about five times as long as the greatest width of the
cephalothorax. Abdomen longer than the combined length of
capitulum and cephalothorax. Capitulum not much wider than
long. Striations on dorsal surface apparently all longitudinal in
direction, the convoluted pattern present in D. melesinus
Text-figure 19.
Demodex sciurinus, sp.n., &. Dorsal view.
apparently being absent. Spinules on capitulum very minute as
in D. folliculorwm and D, melesinus, but not so short as in the
latter. Distal part of capitulum sharply angular and salient
laterally.
_, Measurements. Total length of body 165 w; length of cephalo-
thorax and capitulum 67,4; of abdomen 98. Greatest width
NEW OR LITTLE-KNOWN ACARI. 995
of cephalothorax 33 4; of abdomen 29 »; of capitulum (at base)
22 4; length of capitulum 18 yj.
Host: Brown Squirrel (Sciwrus vulgaris). A specimen of this
mite was collected from a squirrel received from Inverness,
21.iv. 1922.
"TARSONEMUS TRANSLUCENS Green.
This mite has already been recorded as occurring on tea in
‘Ceylon and India, and also described by myself from specimens
found on Capsicum in Trinidad (7. W. Urich Coll.). Professor
G. 8. Kulkarni has brought to the Museum specimens found on
potatoes at Poona. He states (Agric. Journ. India, vol. xvii.
part 1, Jan. 1922, pp. 51-54) that this mite is the cause of the
very serious ‘‘ Jurda” disease of Chilli (Capsicum) and also of
the “‘ Zambera” disease of potatoes in India.
'TARSONEMELLA, Subgen. nov.
Closely allied to Tarsonemus, but the legs of the first pair
stouter than the others and without pulvillus. Also the pseudo-
stigma normally present between the first two pairs of legs in
Tarsonemus is absent. L*ourth leg of female modified in a
similar way to that of the typical Z'arsonemus, the terminal
segments being attenuated and ending in a long hair.
Tarsonemus (TARSONEMELLA) AFRICANUS, sp. n. (Text-figs.
20 & 21.)
2. Shape rather elongate-oval, the body being much longer
than wide. Tarsus of first leg with a well-developed but not
very large claw, which is slender, curved, and sharply pointed.
Dorso-laterally near the distal end of this tarsus there is a slender,
striated cylindrical seta of fair length, and near the middle there
are two similar but smaller sete, one of which is very slightly
clavate. ‘lwo or three stiff hairs with broken-off ends like those
present on the same segment in Pseudotarsoncemoides spiri-
tarsus, Sp.n.,also occur on the first tarsus. T'arsus of the second
leg with a striated seta dorsally of about the same size as the
longest one on the first tarsus, and also a shorter and stouter seta
on its posterior side. ‘Tarsus of second and third leg with a
pulvillus and paired claws. Hairs on body short, especially the
ventral ones; owing to the fact that the specimens of this mite
are mounted in balsam, it is difficult to be certain of the exact
number and distribution of these hairs.
Length (including capitulum) 200 p.
Host: A hymenopterous insect (Agaon aystrum Waterston);
from Koforidua, Gold Coast (4.iv. 1921).
996 MR, STANLEY HIRST ON
Text-figuire 20.
TERII~—
Tarsonemus (Tarsonemella) africanus, sp. n., ¢. Dorsal and ventral views.
Text-figure 21.
Tarsonemus (Tarsonemella) africanus, sy}. 0., 9. ‘Terminal segments of
first leg greatly enlarged.
NEW OR LITTLE-KNOWN ACARI. 997
PSEUDOTARSONEMOIDES SPINITARSUS, sp. n. (Text-figs. 22 & 23.)
2. The usual two little scars (representing accessory stigmata ?)
are minute and oval in shape, being situated widely apart from
one another not far from the lateral margin of the conical
anterior prolongation of the cephalothorax. Pseudostigmata
Text-figure 22.
Pseudotarsonemoides spinitarsus, sp. D., 2.
oval in shape and furnished with very fine short hairs. There
is a hair on the dorsum behind the pseudostigma. <A transverse
row of four fine hairs is present on the dorsal surface of the first
abdominal segment. Second abdominal segment apparently
Proc. Zoou. Soc.—1923, No. LXV. 65
998 MR. STANLEY HIRST ON
without any hairs dorsally. Third abdominal segment with a
pair of dorsal hairs. Fourth abdominal segment with two pairs
of hairs, the inner ones being much shorter than the outer,
exactly as in P. eccoptogasteris. The paired lateral epimeral
structures lying between the first two legs are not joined to the
central linear (longitudinal) chitinous structure (Vitzthum’s
figure of the ventral surface of P. eccoptogasteris shows these
lateral epimera joined to the longitudinal central chitinous
structure). First leg slightly enlarged; its claw well-developed,
but not very large and moving against a short, stout curved
spinule. ‘There are a few fine hairs of ordinary type, including a
long fine hair near the middle of the dorsal surface, on the first
Text-fi gure 23,
Pseudotarsonemoides spinitarsus, sp. n., 2.
a. Terminal segments of first leg.
6. Tarsus of second leg,
tarsus. ‘Three of the hairs are blunt, having the broken-off
appearance characteristic of certain hairs also present in
Tyroglyphid mites. There also seem to be three sensory setie
(sinneskolben) on the first tarsus; the one nearest the distal end
is cylindrical, and projects almost at a right angle to the segment ;
the other two sensory set lie parallel to the tarsus, one of them
being stout and club-shaped, the other slender and difficult
to see. ‘Tarsus of second leg with a slender sensory seta
(sinneskolbe) similar to that present on the first tarsus; there is
also @ strong dorso-lateral spinule on the second tarsus (see text+
fig. 23 dD).
Length of body (including capitulum) 267 »; its width 135 p.
NEW OR LITTLE-KNOWN ACARI. 999
Host: The Klm-bark Beetle (Scolytus destructor); a single
female specimen found on the larva of this beetle at Kew,
19. x. 1922 (2. C. Fisher).
Nore.—This species seems to differ from Vitzthum’s P. eccopto-
gasteris in having a stout spinule on the dorsal surface of the
second tarsus.
LisrkOPUOROIDES, gen. nov.
Body flattened dorso-ventrally instead of laterally, as is the
ease in the genus Listrophorus, etc. All legs with a sucker or
pulvillus, that of the first leg being minute and that of the other
legs sinall also. ‘Terminal segments of first and second legs
grooved ventrally, the sides of the groove being provided with
little knobs and processes for gripping the hair of the host.
LisTROPHOROIDES ZTHIOPICUS, sp. n. (Text-fig. 24 a.)
3. Body moderately elongated. Capitulum shaped rather
like that of a louse, being roughly triangular but with the lateral
angles salient ; it is short and wide. Posterior end of body in this
sex sometimes expanded and almost lobe-shaped as shown in
text-fig. 24; in other specimens the sides of the posterior end of
the body are straighter (but this is perhaps due to distortion
through shrinkage). There are three pairs of hairs on the
posterior end of the body, but they are inconspicuous, two pairs
being very short and fine, and the remaining pair not very long,
Anal suckers not distinct. Tarsi of anterior legs short and
modified as described above; the tarsi of the posterior legs are
rather long and fairly slender, but the other segments of these
limbs are rather stout. Posterior margin of third coxa furnished
with two blunt spurs, the inner one being fairly large.
Length (including capitulum) 450 »; width 187 p.
Host: Cricetomys gambianus ; Accra (Scott-Macfie Coll.).
ListrorHorus BOTH#, sp. n. (Text-fig. 24 6, c, d.)
dg. Body narrow and elongated, being about four times as long
as wide. Capitulum also elongated and rather narrow; its front
margin shaped as shown in text-fig. 24c, being only slightly
prominent in the middle. Posterior end of abdomen shaped as
shown in figure, terminating in a pair of very delicate lobe-
shaped lamine. Just in front of these lobe-like structures there
is a more strongly chitinized curved line, which is probably the
real posterior margin of the body. <A pair of long hairs spring
from the posterior end ‘of the body. Anal suckers distinct but
small, and rounded in outline. ‘Tarsus of first and second leg
furnished with two hairs with blunt ends, one of them being
rather long.
2. Body also elongated as in the male. Unfortunately our
female specimens are not well mounted, and therefore cannot be.
described.
65*
1000 ON SOME NEW OR LI'TTLE-KNOWN ACARI.
Text-figure 24.
op Highley dd
a. Listropacroides ethiopicus, zen. & sp... g. Ventral view.
b. Listrophorus bothe, sp. n., d. Dorsal view.
c. Ventral view of anterior extremity of Listrophorus bothe, 6.
d, Veutral view of posterior extremity of Listrophorus bothe, 6.
3. Length (including capitulum but not long posterior hairs)
380 wp. Greatest width 93 p.
2. Length (including capitulum) 430-440 p.
Tost: Gerbille, ut Bothaville, Orange Free State (G. A. UW.
Tedford).
CAPARINIA ''RIPILIs Michael.
‘Tie presence of » system of tracheal tubes in sarcoptid mites of
the genus Otodectes, namely in Otodectes cynotis var. cati and var.
furonis, has already been pointed out by the author (Journ.
Quekett Micr. Club, vol. xiv, 1921, pp. 229 & 230, text-fig. 1).
‘Tracheal tubes also occur in Caparinia tripilis, and ave very
similar in appearance to those present in Otodectes cynotis.
A parently Otodectes and Caparinia ave the only members of the
family Sarcoptidie possessing a respiratory system of tracheal
tubes ; bat further investigations on this subject are necessary.
THE GENERATIVE ORGANS IN THE MALE CIIMPANZERL. 1001
45. On the Pelvic Muscles and Generative Organs in the
Male Chimpanzee. By Crannus F. Sonsraa, M.D.,
I.Z.8., Anatomist to the Society and Demonstrator of
Anatomy, Universiiy College.
[Received October 1, 1923: Read November 6, 1923. ]
(Text-figures 58-64.)
The present paper is intended to fill a gap in the literature on
the anatomy of the Chimpanzee, for, although there ave descrip-
tions of parts of the male generative organs and the arrangement
of the muscles and pelvic viscera, no full account of these parts
exists. They have been described fully in the female by Lart-
schneider (4) and myself (9). The material on which this account
is based consisted of the pelvis of an adult male animal, which
lived in the Society’s Gardens for nine years, and a young male,
about nine years old, which I received soon after it arrived in
London.
The Male Generative Organs.
Keith (2) states that “The external generative organs are of
surprisingly small development; but it must be kept in mind
that they have been studied on animals, for the greater part
immature, or, if adult, upon animals contracted by long immer-
sion in alcohol.” In the two fresh specimens examined by me
the penis was certainly small, but the scrotum with its contained
testicles was large.
The penis is tapering, and it is frequently seen in the erect or
semi-erect condition. Duvernoy (1) points out that it is often
erected when the animal is drinking, particularly if a little wine
is added to the fluid. After death the penis may be quite con-
cealed by the prepuce. ‘he anterior extremity has a slight
collar-like rim, with a small process underneath. This is the
sole representative of the glans penis. In the Orang, according
to Milne-Edwards (6), there is no rim ; and Duvernoy figures an
expanded, vertically cleft glans of dark colour in the Gorilla.
Kohlbriigge (3) figures a globular expansion of the end of the
penis in the Gibbons. I also observed a well-marked glans
penis. The meatus urinarius is vertical and there is a fossa
navicularis. ‘he prepuce is thick in both Chimpanzees and
Gibbons.
The corpora cavernosa axe cylindrical and of small, but uniform. ‘
diameter. ' They are not expanded close to their origins as in
Man. They are bound down to the lower borders of the pubic
rami, and are covered on three sides by the ischio-cavernosus
1002 DR. C. F. SONNTAG ON THE PELVIC MUSCLES
muscles. The corpus spongiosum has a small bulb attached to
the triangular ligament. Its diameter is greater than that of
either corpus cavernosum. ‘The corpora cavernosa soon come
together. At first they are separated by a thick septum formed
from their contiguous walls, but the septum soon thins out and
disappears. ‘Ihe doxsum has a groove for vessels and nerves, but
the corpus spongiosum is not lodged in a ventral groove. ‘These
conditions are seen in text-fig. 58 M-H. In the Orang the
conditions are very similar to the above for the septum between
the corpora cavernosa. soon disappears; and the core of erectile
tissue is smaller and the fibrous tunic is relatively thicker than in
the Chimpanzee and Man.
The penis contains a small bone, first described by Crisp*. It
Text-figure 58.
E Ga
Cross-sections through the prostate and urethra (A-C) and through the elements of
the penis (H-II). These sections are arranged in order from the base of the
prostate to the anterior part of the penis; D: the orilices of the ejaculatory
ducts in the urethra.
is a little more than half an inch long, fusiform in shape, with
rounded extremities. Crisp figures a rather different shape,
whilst Milne-Edwards (6) described and figured the bone as ex-
panded towards one end in the Orang. ‘The penis is fixed to the
front of the pubis by two suspensory ligaments, between which
vessels and nerves run to the dorsum.
The serotwm is capacious, and varies in size from time to time.
The skin is darkly pigmented, and has crisp hairs. ‘The median
raphé is marked on the surface as a white line, which runs on to
the penis. The Gibbon has not got a prominent scrotum, no
raphé is present, and the root of the penis lies nearly between the
ischial callosities.
Lhe perinewm is long in the male, but short in the female.
* Proc. Zool. Soc. 1865, p. 48.
AND GENERATIVE ORGANS IN THE MALE CHIMPANZEE. 1003
The testicle (text-figs. 59 and 60) is relatively larger than in
Man. In an animal 47 inches long from the crown to the heels,
the testis is 2°3 inches long, 1°7 inches wide, and 1 inch thick.
The line of attachment of the tunica vaginalis (L.T.V.) runs
obliquely across the epididymis (EPID). he upper pole is full
and rounded, and a well-marked recess (S.H.R.) lies between 1t
and the overhanging globus major (G.Ma.) of the epididymis.
The lower pole is not so massive as the upper pole. The epididymis
is a long, narrow body, whose ends are folded inwards towards
one another. The flexures constitute the globus major (G.Ma.)
and globus minor (G.Mi.). The middle part is straight. On
section the testis is seen to have a thick tunica albuginea (‘T.AL.)
with blood-vessels, which are very tortuous. Trabecule pass
inwards between the tunica albuginea and the corpus Highmori-
anum (C.H.), which is a large mass of connective tissue with
seminal ducts and large blood-vessels. The corpus is relatively
Text-figure 59.
CMA.€
2: Sian
ANTERIOR VIEW MESIAL VIEW
LATERAL VIEW
External appearances of the testis. Letters in text.
larger than in Man. ‘The trabecule bound well-marked loculi, in
which the brownish-yellow seminiferous tubes lie. The trabeculee
also form guides for blood-vessels. The elements of the duct
apparatus are as in Man. Thus there appear in succession the
seminiferous tubules, vasa recta, rete testis in the corpus High-
morianum, vasa efferentia, coni vasculosi, and the canal of the
epididymis. ‘he epididymis is loosely bound to the testis. At
the upper pole of the testis a hydatid occurs.
The testis of the Gibbon (text-fig. 60 B) presents a totally
different picture from that in the Chimpanzee. <A deep fissure
divides the organ into two. No corpus Highmovianum is seen,
and the surface appears divided by many small sulci into
numerous lobules, which are granular. No marked trabecule are
present in the sulei between the lobules. ‘The internal fissure
corresponds to an external fissure.
1004 DR. C. F. SONNTAG ON THE PELVIC MUSCLES
The sac of the tunica vaginalis is completely shut off from the
abdominal cavity in the Chimpanzee and Man. In the Gibbon
and Orang the orifice remains open, but a ridge prevents the
testis passing to the abdomen in the latter (Milne-EKdwards: 6).
The general characters of the testis, particularly the size of the
corpus Highmorianum, are identical in the Chimpanzee and
Orang. Duvernoy (1) points out that the testis of the Gorilla is
small relatively to the size of the animal; his illustration shows
that the epididymis has the same appearance as, but is relatively
larger than in the Chimpanzee,
Retzius (8) has made observations on the spermatozoa of the
Apes, and finds that those of the Gorilla resemble the human
spermatozoa most closely.
The vas deferens is gradually formed by a narrowing of the
canal of the epididymis; and it has the whip-cord consistence
Text-figure 60.
Transverse sections through the testis of the Chimpanzee (A) and Gibbon (B).
The halves of the cut testis are folded outwards. Letters in text.
which is usual. It lies in the posterior part of the thick sper-
matic cord. Its relationsin the inguinal canalareasin Man. It
runs on the deep surface of the pelvic wall along a line corre-
sponding to the diameter of the acetabular cavity. Then it passes
between the bladder and the immense seminal vesicle of its own
side. At the level of the middle of the bladder the vasa deferentia
come together and run parallel to one another to the prostate.
They exhibit fusiform ampullary dilatations ; then they narrow
and pass into the prostate, where they unite into the common
ejaculatory ducts with the ducts of the seminal vesicles. In Man
and the Orang they receive the seminal ducts before they enter
the prostate.
The seminal vesicles (text-fig. 61, V.S.) are two immense, pyri-
form bodies, whose narrow ends are directed downwards and
AND GENERATIVE ORGANS IN THE MALE CHIMPANZEE. 1005
inwards towards the prostate gland. They conceal the back of
the bladder and extend upwards along the rectum (text-fig. 62, R.).
Each consists of a long, coiled tube with no saccular dilatations.
The same arrangement is present in the Orang and in the
Cebidee, whereas Man has many glands and a short tube.
Leuckart (5) pointed out that the length of the tube is inversely
proportional to the importance of the diverticula, so the vesiculze
are important secreting organs in Man. The gland has few
ducts but an enormous number of lobules in J/andrillus (10)
and some other Cercopithecide. The tubes in the Chimpanzee
contain milky fluid. ‘Thus the vesicles show the following
variations :—
1. Tubes small, lobules numerous—Homo, Cercopithecide.
2. ‘Tubes long, no lobules—Simia, Anthropopithecus, Cebus.
Text-figure 61.
als
The internal generative organs of the Chimpanzee. B: bulb of urethra; CAP:
prostatic capsule laid back; C.L.A: cut edge of levator ani; C.S: corpus
spongiosum; T.J: tuber ischii; T.L: triangular ligament; U: urethra.
Other letters in text.
The prostate (text-fig. 61, P.G.) is of moderate size, and pyra-
midal in shape with a rounded apex. In a Chimpanzee whose
bodily length is 47 inches, the prostate is ‘8 inch long and
-55 inch wide across the base. It has a_ well-marked capsule
derived from the pelvic fascia, and its relations to the neighbour-
ing structures in the pelvis are asin Man. A furrow represents
1006. DR. G. F, SONNTAG ON THE PELVIC MUSCLES
a trace of the original division of the gland into two parts, and
there is a small pyramidal lobe (P.L.). The tissue is dense.
When sections are made through it (text-fig. 58 A-D) it is seen
how the urethra is not surrounded by prostatic glandular tissue,
‘but is ante-prostatic; and Milne-Edwards showed how the same
arrangement is present in the Orang-Outan. The ducts of the
vesiculze seminales enter the prostate above and lateral to the two
vasa deferentia. The vasa and seminal ducts unite to form two
tubes, which open side by side into the urethra at the proximal
end of a broad ridge (text-fig. 58 D.). The mouths of the ducts
are thickened.
Cowper's Glands (text-fig. 61, C.G.):—-These glands, which are
Text-figure 62.
The rectum (R.) and its relations to the seminal vesicles (V.S.), hemorrhoidal artery
(H.A.), and pudendal artery (P.A.); CO: coceyx; C.G: Cowper’s glands ;
G.S.8S.L: great sciatic ligament; T.I. and T.L. as in text-fig. 61.
the size of large peas, are situated as in Man. Well-marked
duets open into the urethra. The glands are not covered by
fibres of the bulbo-cavernosus muscles. They are relatively larger
in the Chimpanzee and Orang than in Man. The orifices of the
ducts are very minute.
The bladder has well-marked lateral ligaments, and the urachus
ig small, but the anterior ligament is well marked. ‘Lhe anterior
wall of the bladder is closely bound down to the pubis below.
The mucous membrane is thrown into fine longitudinal folds,
quite unlike the greatly corrugated interior which was seen in a
female animal.
AND GENERA TIVE ORGANS IN THE MALE CHIMPANZEE. 1007
Muscles of the Perineum and Pelvis (text-figs. 63 and 64).
When the skin is removed from the perineum the subjacent
fascize are seen to consist of an imperfect superficial fatty stratum
and a deep fibrous layer, which is not connected to the triangular
ligament.
The sphincter ani externus (text-fig. 63) is a powerful muscle,
and there is continuity between it and the levator ani. 1t con-
sists of several parts, most of which form collars round the gut.
The superficial part (8.A.8.) arises in front from the triangular
Text-figure 63.
The pelvic muscles. A.C.R.: ano-coccygeal raphé; ©.P.P: central point
of perineum. Other letters in text.
ligament in the central point of the perineum ; they sweep round
the anus, but they have no fixed posterior attachment. Two long
muscle bands (8.A.R.) arise from the inner aspects of the posterior
parts of the ischial tuberosities and meet behind the anus; some
of the fibres are continuous with those of the superficial part of
the sphincter. ‘The complete band appears like a sling. It is not
present in all Chimpanzees, and it may correspond to the retractor
ani of lower Mammals. The third part of the muscle (S.A.T.) is
composed of fibres arising from the central point of the perineum
and sweeping round the gut; it forms a thick collar above and
1008 DR. C. F. SONNTAG ON THE PELVIC MUSCLES
behind the superficial part. ‘The deep part of the muscle (S.A.D.)
runs round the gut from the central point of the perineum in
front to the ano-coccygeal raphé behind ; its deep aspect is in
contact with, and receives fibres from, the levator ani.
Transverse Perineal Afuscles:—In the two male Chimpanzees,
as in the female animal already deseribed by me, there is no trace
of the superficial or deep transverse perineal muscles.
The bulbo-cavernosus (text-fig. 63, B.C.) arises from the median
ventral raphé on the bulb and corpus spongiosum, and the fibres
run ventro-dorsally. The posterior fibres completely encircle the
bulb, are continuous with the fibres of the opposite muscle, and
Text-figure 64.
CS.
(XE
The levator ani. ©.C: corpus cavernosum; C.S: corpus spongiosum; T.L.U :
upwardly curved base of the triangular ligament. Other letters in text.
are inserted into the triangular ligament. ‘The two muscles are
separate distal to the bulb, and gradually leave the ventral snr-
face of the corpus spongiosum; they are inserted by small slips
into the fascia on the dorsum of the corpora cavernosa.
The ischio-cavernosus (text-fig. 63, I-C.) wvises from the inner
aspect of the lower end of the ischial ramus, the inner aspect
of the tuber ischii and the great sacro-sciatic ligament. It is
inserted into the under surface of the crus penis and the outer
and dorsal aspects of the corpus cavernosum.
When the penile muscles, which are in contact, are removed,
AND GENERATIVE ORGANS IN THE MALE CHIMPANZEE. 1009
the crura, bulb, and triangular ligament are exposed. The liga-
ment is strong and composed of two layers, which meet behind
and are turned up over the posterior part of the bulbo-cavernosus
muscles. The terminal part of the pudendal artery turns up over
the recurved base (text-fig. 65, P.A.). The bulb of the corpus
spongiosum is attached to the perineal surface of the ligament.
When the superficial layer is removed ‘the two large Cowper’s
glands are seen lying in pockets at the sides of and above the
bulb. The glands shine through the superficial part of the
ligament.
The compressor wrethre encircles the urethra between the layers
of the triangular ligament.
Between the anal canal and the obturator internus there lies on
each side a narrow, deep ischio-rectal fossa. And the pudendal
artery, after emerging from under cover of the great sacro-sciatic
ligament, passes along under the fascia at the meeting of the outer
and inner walls of the space. The vessel turns up over the trian-
gular ligament and ends by supplying the crura, bulb, Cowper's
glands, and the skin of the perineum (text-fig. 63, : A. ). Within
the space it gives off the inferior hemorrhoidal artery s the
sphincter and Tevator ani (text-fig. 63, I.H.A.).
‘Che ischio-rectal fossa has the same boundaries as in Man, and
the fasciz covering its walls are likewise similar. Alcock’s canal
is, however, poorer.
The levator ani (text-figs. 63, 64, L.A.) is covered on both sur-
faces by fascie as in Man. ‘The inferior fascia is delicate and
continuous with the equally delicate fascia over the obturator
internus. The muscle arises from the back of the body of the
pubis, the margin and inner wall of the true pelvis and the place
where the ischial spine lies in Man. he fibres pass downwards
and inwards to be inserted into the central point of the perineum,
the sides of the anal canal, the sphincter ani externus, the ano-
coceygeal fascia, and the lower sacral and coccygeal vertebri.
The anterior two-thirds of the muscle ave thick and powerful, but
the posterior third is thin.
The coceygeus has similar attachments and relations to those
in Man,
Conclusions.
A. The external generative organs in Man differ from those of
the Chimpanzee in the following particulars :—
1. The glans penis is well marked.
2. There is no os penis.
3. The corpus spongiosum lies in a groove on the ventral
rue of the fused corpora cavernosa.
= The septum between the corpora cavernosa persists longer.
. The connective-tissue walls of the corpora cavernosa are
aoe so thick and they have more erectile tissue.
Man also differs from the Orang in the above features. The
chief difference between him and the Gorilla lies in the absence
of the.os penis.
1010 DR. GC. F. SONNTAG ON 'THE PELVIC MUSCLES
B. The internal generative organs in Man differ from those in
the Chimpanzee in the following respects :—
1. The testis is relatively smaller and the corpus Highmori-
anum is smaller. The epididymis has a different form.
2. ‘The vesicula seminales consist of a short tubular and a
large glandular part, whereas the reverse is the case in the
Chimpanzee. Moreover, according to Milne-Ndwards, the
Chimpanzee has the same characters as the Orang.
3. Cowper’s glands are relatively smaller.
C. Man and the Chimpanzee agree in the closure of the com-
munication which originally existed between the tunica vaginalis
and the peritoneal cavity. In the Orang and Gibbon this remains
open.
D. The urethra is ante-prostatic in the Chimpanzee and Orang,
whereas it descends through the prostate in Man.
EK. The orifices of the ejaculatory ducts are close together in the
Chimpanzee, and they appear rounded. In Man they are sepa-
vated slits. In AMandrillus (10) there is a common orifice for the
two ducts. Milne-Edwards (6) points out that the arrangement
in the Orang is as described above in the Chimpanzee.
I’, The spermatozoa of the Chimpanzee do not resemble those
of Man, but those of the Gorilla do according to Retzius (8).
G. The interior of the bladder and rectum is as in the female
alveady described by me (9).
H. The muscles of the perineum and pelvic floor are built on
the same plan as in Man, but the Chimpanzee has a retractor ani
and no transverse perineal muscles are present. ‘The ischial spine
varies, so there are slight modifications in the origin of the
levator ani.
It is, therefore, evident that Man differs in many ways from
the Chimpanzee and Orang in the characters of the male repro-
ductive organs. He agrees with the Gorilla in the presence of a
glans penis, the characters of the vesicula seminales, and in his
spermatozoa.
Bibliography.
1. Duvernoy, G. L.—‘‘ Les Caractéres anatomiques des grands
Singes.” Arch. Mus. Hist. Nat. Paris, 1855-56, pp. 1-248.
2. Keren, A.—‘* An Introduction to the Study of the ecilono
poid Apes.” Natural Science, vol. ix., 1856.
3. Koutsrican, J. HW. F.-—* Versuch einer Anatomie des Genus
Hylobates.” Max Weber's Zool. Ergebnisse einer Reise in
Niederland, ete. 1890.
4, LarrscHNeiEr, J.—‘‘ Die Steissbeinmuskeln des Menschen
und ihre Beziehungen zum JZ, levator ani und zur Becken-
fascie.” Denkschr, K. Ak. Wiss. Wien, 1895, pp. 95-136.
10.
AND GENERATIVE ORGANS IN THE MALE CHIMPANZEE. 1011
. Leuvcxarer.——Arts. ‘ Vesicula Seminales” and “ Vesicula
Prostatica” in Todd’s Cyclopedia of Anatomy and Phy-
siology, Vol. iv.
. Mitne-Epwarps, M. A.——‘‘ Observations sur Deux Orang-
Outans Adultes.” Nouv. Archiv du Mus. d’Hist. Nat.
1895, pp. 57-82.
. Oupemans.—“‘ Die accessor. Geschlectsdriisen d. Stiugethiere.”
1892.
. Rerzius, G.——“‘ Ueber die Form der Spermien bei den Anthro-
poden Affen.” Biolog. Untersuchungen, Jena, 1910.
Sonnrac, C. F.——“ On the Anatomy, Physiology, and Pathology
of the Chimpanzee.” P.Z.S. 1923, pp. 323-429.
Sonntag, C. F.—‘‘On the Anatomy of the Drill.” P. Z. 8S.
1922, pp. 429-454.
EN
Ne
i
NOTES ON EAST AFRICAN INSECTS. 1013
phe 49. Notes on Hast African Insects collected 1915-1922.
By Arrvaur Lovenriper, F.8.8., C.M.Z.8.
[Received October 20, 1923: Read November 6, 1923.]
Tt was my intention on returning to England after my first
term in East Africa (1915-1919) to publish such field-notes as
were of general interest as each order of insects was worked out,
over twenty thousand insects having been collected.
~The difficulty of getting these determinations performed has,
however, prevented The carrying out of my original plan, and as
these notes are accumulating, I have made sundry extracts from
my journal in the hope that entomologists may find amongst these
casual jottings something of interest.
The present miscellany i is divided into two parts, (A) beings a
list of Hymenoptera and their Dipterous mimics taken at Kilosa,
and (B) more general observations on Hymenoptera, Coleoptera,
Lepidoptera, and Orthoptera. Next year, when more of the
material will have been identified, it may ke possible to publish
a second instalment covering the same and other orders.
Localities may be grouped as follows :—
1. Kenya Conony (late British East Africa),
Frere ‘own, on mainland opposite Mombasa Island,
Mbunyli, north of Taveta and west of Kilimanjaro.
Nairobi, 327 miles west of Mombasa.
Ngong, some six miles from Nairobi.
2. TANGANYIKA Ternrrory (late German East Africa).
Dutumi, in Morogoro District.
Handeni, in Pangani District.
Izikisia, 40 km. N.K. of Tabora in Tabora District.
Kahe, in Moshi District.
Kilosa, 80 km. W. of Morogoro in Morogoro District.
Longido West, z.e. west District of Mt. Longido, Arusha.
Myombo, 75 km. W. of Morogoro in Morogoro District.
Mkomasi, 30 km. N.W. of Lushoto in Usambara District.
Morogoro, on Central Railway, 2100 km. west of Dar es
Salaam.
Moshi, on Tanga Railway in Moshi District.
Ngari Mtoni, six miles west of Arusha, Arusha District.
Ngeri-ngeri Ree south of Ngeri-ngeri Station, Cent. Rly.
Singida, 120 km. north of Kilamatinds i in Dodoma District.
Tambali, in Tabora District.
Proc. ZOOL. Soc.—192 3, No. xy s, HS ee «666
1014 - MR. A. LOVERIDGE: NOLS ON
3. Portucurse Hasr Arnica.
Lumbo is the railhead on mainland 3 miles from Mozambique.
My thanks are due to Professor Poulton for getting most of
the identifications made (the work has been particularly difficult
owing to rough handling of a store-box in the post, resulting in
many of the specimens being badly damaged and in some eases
detached from their labels), for proof-reading and his many
kindnesses and ready help; to W. C. Crawley, The Rev. F. D.
Morice, Dr. J. Waterston (Hymenoptera); Major K. K. Austen,
D.S.0., E. Brunetti, F. W. Edwards (Diptera); G. J. Arrow,
K. G. Blair (Coleoptera); B. Uvarov (Orthoptera); Dr. F. A.
Dixey, Prof. E. B. Poulton, W. H. T. Tams (Lepidoptera); A. S.
Hirst (Arachnida), for making the determinations; and to my
chief, Mr. C. F. M. Swynnerton, for the interest he has always
shown in entomological pursuits, particularly those connected
with mimicry and warning coloration.
The material referred to in the following pages is preserved in
the Hope Department of the Oxford University Museum, where
it may be studied by any naturalist.
A. FIyMENOPrERA AND THEIR DirrErous Mimics.
With one exception all the models and mimics mentioned below
were taken within an area of 300 square yards on the Otto Planta-
tion, Kilosa, Tanganyika Territory, and with six exceptions
(mimics 4, 6, 8, 11, 38, & 40) all were taken on the mosquito gauze
protecting the verandahs of one or other of two houses not more
than two hundred yards apart and in the same clearing. ‘This
rather adds to the interest of the specimens, and it will be under-
stood that unless otherwise stated all the specimens were taken
on this gauze.
The actual specimens referred to in the notes were received for
every number except, perhaps, No. 34, which is a little uncertain.
1. Xytocopa rnconstans Sm., 2. 31.1i1.21. Only a single
example of this Carpenter-bee taken, a very perfect model to (4).
The timber of both these houses is infested with the borings of
(2) and (3), which I consider form an association in the matter
of markings or colour from which (4) is likely to benefit.
2. Xyiocopa carrra L., 9. 13.iv.22. Very common.
3. XyLocoPpa NicRiITA F., 9. 24.1v.22. Very common,
4. HyprrecuIA BIFASCIATA Griinb., 9 (Laphrine: Asilide).
13.iv.22. This Dipterous mimic was taken at 8 a.m. resting in
bright sunshine on the verandah-rail just four feet below the
nesting-holes of X. nigrita F. When approached it flew with a
faint buzz to the rain-gutter on the roof and right alongside the
nesting-holes: netted.
5. PSAMMOCHARES sp., probably venans Kohl (Pompilide:
Fossoria). 6.vi.21. Very common, as also several similarly-
‘coloured but larger species, which can be seen flying about the
paths and open clearings on any sunshiny day.
EAST AFRICAN INSECTS. 1015
_ 6. OrEcrocERrA praBotus Wied. (Tachinide). 17.iv.22. When
walking down a steep path at 1.45 p.m., I noticed this Dipterous
insect fly up suddenly—the flight was short and rapid—and,
valighting again upon the path, it ran to and fro with jerky move-
ments of the abdomen till it reached a stone, upon which it rested.
‘The movements were almost identical with those of (5) and its
allies. The fly must be very scarce here, as I have not met with
it before during twelve months’ residence at Kilosa.
7. SYNAGRIS CARTNATA Sauss., var. ALBONOTATA Beq., 9
(Eumenide: Diploptera). 1.iv.22.. Very common, asare many
other similarly-coloured wasps, blue being their common livery as
yellow and black is for the vespids at home. JLthynchiwm luctwosum
pees is one of these ; it also enters into the colour association of
No.
8. BromMopaiLa CAFFRA Macq. (Ortalide). 9.11.21. On a
leaf beside the path, almost at the same spot as (6). I do not
regard this fly as a direct mimic of (7), but it must derive advan-
tage from adopting the livery of the wasps, which are so common.
On 17.11.17 I took a number of these flies on flowers and herbage
at Morogoro, and noted at the time their slowness of movement.
This sluggishness was so pronounced that it was not a difficult
matter to catch them by hand *.
9. Kumenes MAXILLosA de G., 9 (Humenide). 9. iv. 22,
10. EUMENES DYSCHYROIDES Gribodo, 9. 29.1. 21.
11. Puysocernana sp., 9 (Conopide). 25. 111.22. I think this
is a perfectly wonderful mimic of (9) and (10), but the semi-
transparent wings are more like those of other hornets taken
here whose abdomens, however, are normal-shaped. ‘This fly was
resting on the blossom of a Zinnia at 5.30 p.m., and buzzed
loudly when caught by the fingers; it was extremely sluggish.
‘In my diary I have just come across the following note under
date 3.v.17, Morogoro. “'Took a fly from a spider’s web; it is
the very image of the long-waisted wasp. I was deceived by its
appearance at first, but the spider was serenely tackling it.”
Whether this is the same species I cannot say, as the fly was sent
home for identification and cannot be traced at the moment.
It is either in the Nairobi or British Museum (Nat. Hist.).
12. PsAmMMocnHARES sp. A. (Pompilide: Fossoria). 1. v.22.
Scarce, at Jeast in the houses; there is a bee here very similarly
-coloured which I] have taken several times.
* The fly ejects vellow liquid from its mouth when handled, and was refused by
‘Dr. G. A. K. Marshall's baboons and Cercopithecus (Trans. Ent. Soc. Lond. 1902,
p. 531). Bromophila caffra is figured by Dr. Marshall on plate xxiii. fig.27, as one
-of a S. Rhodesian group of insects with “ black bodies, blue wings, and red or yellow
. heads” (figs. 20-27). See also Dr. G. D. H. + Carpenter i in Proc. Ent. Soc. 1918, p. ¢,
-and Trans. Ent. Soc, 1921, p. 72. \
66*
1016 MR. A. LOVERIDGE: NOTES ON
13. Necrtromy1a MACULIPENNIS Lw. (Stratiomyide). 2.1. 22.
This fly was buzzing up and down the white ceiling of a lighted
oom, when it was approached by an exceptionally large gecko:
(Hemidactylus mabouia), who followed its movements for some
time, but hesitated to take it.
5”)
close resemblance to the model, During the five months since
2.1.22 five specimens were taken on the gauze, it being about
equally common with the model in this situation. There is a bee
here very similarly coloured.,
13a. 6.iv.22. Another example with closed wings, showing its.
14, APIS MELLIFERA ADANSONI Latr. 22.111.22. This bee is
a plague, existing in countless thousands in the plantation and
often making its combs in the houses (see p. 1019).
15. Enisranopres quinquemacutatus I'., 9 (Mristaline: Syr-
phide)., 2.1.22. Common but not exactly plentiful; buzzes like:
the bee both in fight and when caught; it causes quite a pricking
of the fingers with its feet when held. On 31. iii. 22 I caught one
of these flies and offered it to my monkey (Cercopithecus albigu-
laris rufilatus), who would have nothing to do with it though he:
eyed it closely. I then proffered it to a Fennec Fox (Octocyon vir-
gatus), who was watching me and was within two feet of the
monkey at the time. He promptly snapped it out of my fingers
and crunched it up. I then released the monkey, who went, off
hunting grasshoppers, and was still busy feeding when I went to
bring him in half-an-hour later, so it was not a question of |
satiety *.
16. Mrcacutne sp., ? uNGULATA Smith, 2 (Apide : Anthophila).
-6.vi.21. A common wild bee. Another bee, Vomia vulpina
Gerst., would act a as model with (16), as also with (14), and (39) ;
and all four species form together a compound colour association
into which still other bees probably enter.
17. Simormes crassrpEs F., @ (Hristaline). 21.1i1,22. A mimic
of (16).
18. Psammocuares sp. B, 1.iv.22. Common.
19 & 19a. PLAGIOSTENOPYERINA SUBMETALLICA Jw. (Steno-
pterine: Ortalide). 11.&15.v.22. In houses even commoner
than the alleged model; when walking slowly up the gauze with
closed wings they look very waspish.
20. IcantA AMBIGUA Gribodo (Vespide: Diploptera). 1.iv. 22.
This wasp builds small nests on the verandah, where the little
colonies of from two to five individuals are quite a common
spectacle.
21, OpyNerus TRopIcALIs Sauss., @ (Vespide). 1. iv. 22.
A closely-related species with similar habits.
’ * Compare Dr. Carpenter’s experiments on monkeys with Hristalis tenax L,
(Trans, Ent. Soc. 1921, pp. 25, 31, 33, 72, 100). ;
LEAST AFRICAN INSECTS. . 1017
22. Dacus rncronaLis Walk. (Trypetide). 1.iv.22. On taking
this fly, I went and captured examples of the models from close
by. ‘'Lhis wonderful mimic is apparently commoner than the
model, but not so in reality I believe, the explanation being that
on enter ing the house it immediately flies to the gauze to escape,
while the wasps, knowing their way about, make straight for
their nests. Accompanying (22) were two other somewhat smaller
but very similar species, together with the following.
22a. DAcUs BREVISTYLUS Bezzi. 21.iii1.22. A smaller species,
and possibly forming a protective association with (22).
23. Cernceris sp.. 2 (Sphegide : Fossoria). 1l.v.22. Not
so common as (20) or (21). The very similar /caria distiyma
Gerst. was also taken.
24, Dacus sp. nr. LOUNSBURYI Coq., 9. 13.iv.22. The only
Specimen taken. .
25. Paniscus opacuxus Thoms., very near the British ¢estucews
Holmgr. (Ophionine : Ichneumonide). 2.1.22. A common ich-
neumon fly.
26. A species of Pimprinm (Ichneumonide) not in Brit. Mus.
9.iv.22. By no means abundant. 2h
27. Puaryura sp. (Mycetophilide). 11. v.22. The only speci-
men of this Dipteron I have taken, though I have seen but not,
. captured others.
28. A species of Microprzip& (Diptera). 2.1.22. Common.
29, Puucticus ELONGATUS F., Q (Stratiomyide). 2.1.22. Very
common. It will bo observed that the model (25) rnd Dipterous
mimics (28) and (29) were taken on the saine day, and not only so
but at the same time, on the same gauzed window within a radius
of two feet. I have subsequently seen the model and mimic
together on several occasions.
29 @. SryLocastEr sp. (Conopide). 1l.v.22. Only a single
specimen taken of this very ichneumon-like Dipteron.
30. Pison sp., probably xanrnopus Brullé (Sphegide: Fossoria).
15.iv.22. Common. i
31. Nomia sp., probably sameas unnamed species from Entebbe
in Brit. Mus. (Andrenide : Anthophila). 27.111.22. Common.
Another bee of the genus Halictus probably enters the same
association.
32. Sarcus sp. (Stratiomyide) and 32 a. Probably Saraus sp.
20. iii. 22 & 9. iv. 22. Dipterous mimics of (30) and cme with
closed wings they are very bee-like.
33 and 330. Humerus sp. (Syrphidm). 3.1.22 & 6.1.22. Bee
mimics. (33a) has its wings set in the attitude of rest seen in
life.
34. Number missing. A large example of No. 31 bears the
following date, but it is rather small as a model for (35). 24, i. 22,
Bee.
1018 MR, A. LOVERIDGE: NOTES ON
35. Microvon sp. (Syrphide). 17.iv.22. A very bee-like fly
when seen walking up the gauze; the illusion is accentuated by
the faint buzz it gives when caught.
36. Iscniopon scUTELLARE TF., 9 = Syrpuus meyprius Wied..
(Syrphine). 1.v.22. This hover-fly is very like its English
relative in appearance, and does not resemble closely any parti-
eular wasp which 1 have taken locally.
37 and 37a. Rnyncntum sp., prob. RADIALE Sauss. (Eumenide =
Diploptera). 20. ii1.22 & 14.1,22. A common wasp.
38. AncyLorriyNncuus Latr. (= XipHocerus Lw.) sp., $ (Dasy-
pogonins: Asilide). 31.ii1.21, Rare: a pair was taken wm
cop. in the open. <A very beautiful general mimic, whose capture:
recalled exciting moments with my first Asilus crabronifornus L.
in Cornwall. 5
39. MreaacuiLe piticrus Mor., Q@ (Apide : Anthophila).
7.1.22. A very common form of bee; there are many species.
with similar markings, either with the transverse bars similarly
‘coloured or picked out in blue or chrome.
40, SisyrNopytEs sp., prob. new, ? (Dasypogoninze). 26.vili.21.
This is the only specimen of the above series not taken at Kilosa,
but at Mkata River some twenty-five miles away. The model
or members of the group of models are almost sure to be found at
Mkata, where I did practically no collecting. This fly, so far as
my memory serves, was captured because it alighted on my coat
sleeve.
B. Genera OBSERVATIONS ON INSECTS.
HYMENOPTERA.
The actual specimens referred to in the following notes on
Hymenoptera were only received for Nos. 41 and 49.
41. A species of Brrityuipm, now regarded as_ Fossorial
Hymenoptera (Proc, Ent. Soc, Lond, 1922, p. xxvi), This small
Hymenopteron, so ant-like in habit and in the way it runsabout,
bites (or stings?) most painfully. The usual place to find them
was on the back of one’s neck inside the collar, where they had
probably got by dropping from the thatched roof. Owing to
their numbers they constituted quite a pest at Jumbe Mbulu’s.
(Singida, Dodoma Dist., 11. x. 21.)
42. Curysipip®. (Actual specimens not available, and it is
unsafe to give names of others.) Some Chrysids, four in number, :
on my window had evidently just emerged from their host cell,
for one of, their number still had the paper cap of its cell adhering
to its feet. (Morogoro, 29. ii. 17.)
43. Xynocora carrra L, My native collector having left one
of these bees in my butterfly net I received a sting. The pain
was very sharp but soon subsided, and no swelling resulted.
(Morogoro, 10.1. 17.)
EAST AFRICAN INSECTS.» 1019
‘44, XyLocora TorRIDA Westw., 9. When resting in their
holes these bees effectually close the opening of the hole with the
dorsal surface of their abdomen pressed closely against it,
yet the tip also so placed as to leave the sting operative. As a
result; I have twice been stung—the first time when leaning
against a pole in a hut, and to-day when removing a rubber-
tree pole which was infested with these carpenter - bees.
I inadvertently placed my hand over one of the entrances, and
was promptly stung on the tip of the second finger of the left.
hand. The pain was intense for a quarter of an hour—fully as.
bad as that of a scorpion sting. Blue was applied for ten minutes.
and the pain passed off. By 4 p.m. there was only a stiffness and
soreness. The following morning the top section of the finger
was much swollen, ‘bam and very tender, and so remained for
three days. (Kilosa, 17-20. 1. 22.)
45. MeLipona Braunsi Kohl (Apide: Anthophila). This bee,
which for my own convenience I have’ christened ‘‘ the thirsty
bee,” was very much in evidence at Lembeni and Dutumi.and in
certain spots at Morogoro and Iilosa. It settles in crowds upon
any exposed surface, such as face and arms, to drink the per-
spiration. It causes much annoyance by hovering in front of one’s
eyeballs, and I noticed they congregated on the lower eyelids of
the transport mules to drink their tears during the campaign,
and caused the animals considerable suffering or at least dis-
comfort. I have often brushed them off as one would a fly
without ever receiving a sting. (Morogoro, 24.1. 18.)
46. APIS MELLIFERA ADANSOoNI Latr. The East African Wild
Honey-Bee is one of the greatest pests in the country. In
Nairobi they do not cause much trouble, but in Tanganyika
Territory they were a great nuisance during the cam paign..
The battle of Tanga was a classical instance; in this fight, firing
through the trees annoyed the bees, who attacked and routed
both combatants *.
My first introduction to the aggressiveness of the insect
occurred near Moshi when crossing a clear, fust-flowing stream..
With legs crossed on my mule’s neck, the water being up to the
saddle-flaps, I was almost unseated by the sudden start given by
the animal, which ducked her head under watery. The next
moment something stung me on the back of the hand which for
the moment I imagined to bea tsetse, an insect with which I was
unacquainted at that time and which we had been told to look
out for in the vicinity of these rivers. Glancing down, however,
I saw it was a wild bee, clouds of which were coming round us,.
and many of my companions were already stung. The horses,
without requiring any .urging, started splashing and galloping
* inven Lettow in his book on ‘this campaign Weeves that the bees were enrazed te
the firing of his machine-gun company, which had to retire no less than the enemy >
and that the widespread rumour that a trap had been laid.was entirely fictitious. °
1020 MR, .Ae LOVERIDGE: NOTES ON
through the water and scrambling up a very steep bank. Nearly
all, the men were waving their helmets wildly about: their
heads, which was probably a mistake, as most of the stings
received. proved to be upon the head. I myself received between
fifteen and twenty—two of these were on my arms, one on a
finger, the rest were about the ears and on the head.
We stampeded for a quarter-of-a-mile before pulling up in
answer to the blowing of a whistle—it was a good thing that
none of the enemy was about to take advantage of our disorder.
As weformed up into squadrons oné of the ofticers rode down the
line asking for me, as he knew that I carried a pair of ento-
mological forceps and he wanted a sting removed from just below
the eye. One man, who had dropped his helmet and gone back to
fetch it, claimed to have a hundred stings; sixty were removed
by a friend of his that day, and I took out eleven the following
morning. The stings were much more severe than those of an
English wasp, and the unpleasant effects lasted from 24 to
48 hours. The person of a hundred stings had the site of each
marked with a purple spot. One of mine had a dark clot of
blood at the site of puncture, and there was a swelling for
six inches around this; the other stings I received caused scarcely
any swelling. The nest must have been dependent from one of
the branches overhanging the river, and { imagine that someone
had brushed against it. (Nr. Moshi, 13. iii. 16.)
At Handeni an Indian was stung to death. The unfortunate
man had entered a deserted banda in which was a bee’s nest, and
the swarm had attacked him; it is surmised that he tripped
and fell, for when found_he was lying unconscious by the roadside
and expired shortly afterwards. I was told that.a couple of mules
were stung to death near the same spot. When at Morogoro,
a monkey was brought to me that had been stung to death by
bees. At the time of attack it was chained to a pole, and though
several persons made gallant attempts to tescue it they failed.
On removing its skin there was scarcely a square inch of its body
unmarked by a sting, the surface of the flesh being covered with
blood. :
_ A swarm entered my monkey’s box, which was fixed on a
tall pole; the animal became excited and received two stings, on
eyebrow and neck respectively. It was rescued by my boy, who
received three stings; three other natives who were passing also
got stung. At 5 p.m.a Kavirondo dug out the pole and, balancing
it on his shoulder, walked through the camp with it; on nearing
the incinerator the bees got excited and, buzzing about his face,
caused his nerve at last to give way, so that he dropped both pole
and box and fled incontinently. It so happened that the box
containing the irate bees fell in the vicinity of the native latrine,
and the rapid exit of boys from the place in various states of
attire caused much merriment amongst their fellows. A sting
which: I received upon my cheek took three days to subside,
(Morogoro, 9, ii, 18.) é Vi i
EAST AFRICAN INSECTS.. 1021
A galvanised iron building in which I took up residence for a
time had a cupboard in one corner, in former times the place
being used as a cotton-mill. This cupboard had two pipe-like
apertures, and seemed to the local bees to be an altogether
desirable nesting-site. On 9.1.17 they swarmed, numbers of
them buzzing up and down the window pane by my table. Others
persistently buzzed round my nose—not that that organ is in the
least flower-like: one of these bees being struck at, stung the
offending forearm some three inches above the wrist. The ‘sting
was not left: behind, and the pain, sharp enough at first, subsided in
fifteen minutes ; the arm swelled considerably, and this swelling
did not entirely disappear till the 13th, 7. e. four days later.
On January 26th they were again unusually bad. Nearly a
thousand must have invaded my quarters during the day; they
came like a cloud, and at one time there must have been fully
five hundred on the window panes; the noise created reminded
one of the room of a wireless operator. I just managed to carry
on with my work, though the angry humming of some round nose
and mouth was trying to the nerves. On February 5th I killed
747 bees on the window pane by quietly crushing them one by
one with-a little metal pill-box. One’ hears of swarms every-
where. Whilst seated writing near the window, I was un-
expectedly stung upon the eyelid by a bee which flew in at the’
window and straight to my eye without any provocation whatever
on my part; the poor creature could not extricate its sting, and
J had to pull it off forcibly, after it and my evelid had fluttered
up and down for a few seconds. It is very usual for these bees
to attack the eyes, and is not a matter of chance.
Another time I noticed some boys pulling a hamali cart in the
road outside, when a sudden commotion arose. One native dived
beneath the.cart and tried to conceal his head beneath his arms,
another stripped off his kanzu and tried to wrap his head in it
and ran away; the bees were left in the road in possession of the
cart, which probably contained some sweet stuff. A similar
instance came to my knowledge at Kilosa, where the cart contained
goor; the bees held up all the traffic in one of the main ORS
pinlle they looted the cart. i
Twice I dislodged the offenders from the cupboard fore!
mentioned by means of smoke. On a third occasion I was telling
a friend of my troubles, when he said ‘ Why do you not clear
them out?” I remarked that I had not the necessary materials
for making a smoky fire. He made answer: “ What do you want
with a fire, a stick is all that is necessary ?” J fetched him this,
and then, seeing that he was in earnest, took refuge inside
my mosquito net. The cupboard, I might add, was nine feet
high with a floor-space of about four feet square. Presently
I ered the stick striking about in the cupboard, and he called
out that the bees were all gone, in proof of which he poked out
a large piece of comb on the end of the stick. It was perfectly
true; he had knocked down the nest in broad daylight; and‘ the
1022 MR. A. LOVERIDGE: NOTES ON
bees had fled out of the pipe-hole like a cloud of smoke and he:
had not received a single sting. He said that to the best of his.
knowledge he had never been stung in his life, but a few days.
later this. was no longer the cave, for he donned a cap in which
was one of his homeless bees, and he was stung on the head.
Within the next few days he received several more stings.
The bees are dependent on water, and if they get into a room
from which there is no escape they soon die. They are very fond
of sisal flowers, and clouds may be observed round the tops of the:
poles when the flowers are in bloom. The blossoms of the rubber-
trees are another favourite source of nectar, but the resultant
honey: which I have tasted is bitter and unpalatable to the
Kuropean, though the natives devour it readily, A. strange thing
that I noticed at Morogoro was their liking for commercial rubber.
Some boxes of this stood in the sun just outside my quarters, and
the heat melted the rubber which Jeaked out from the damaged
boxes; the bees might be seen crawling over this at all hours of
the day. Urine also seems to have a great attraction for them.
When the rubber blossoms are in bloom the presence of the vast
numbers of bees attracted to the plantations causes numbers of
Bee-eaters (Jlerops apiaster and nubicus) to congregate.
They have few enemies [imagine. When swarming in the office
roof numbers of them got into the web of a spider, which after
wrapping them up very effectually in silk, cut them loose, so that
they were constantly landing on my table. A robber-fly settled
on my wife’s arm one day with a bee in its grasp; a leg of the
bee had got entangled in her sleeve, and some movement caused
the insect to fly off; but returning, and failing after several
attempts to disentangle its prey, it settled down and sucked the
juices in situ. (Kilosa, 18.1.22.) A mantis was seen to take one
from the window one day, as is mentioned elsewhere.
' At IGlosa on 14.ii.21.a comb measuring some six inches.
in length and three-and-a-half across was formed in the store.
This was the work of four days, the swarm being small—-a large
swarm will make an astonishing quantity of comb in a short time.
One swarm, which had been established for fully a week in the-
office roof and had a huge comb with quantities of honey, suddenly
left at 10 a.m. in the morning without any apparent reason ;
I watched them go, and then examined the comb and found but
two bees left.
On 12. ii, 22 I recorded that there had been four swarms in
the office and three in my store-room during the past month.
This swarming generally takes place during the earlier part of
the year, synchronising with the greater rains and the abundance
of food furnished by the rubber blossoms, which doubtless assists
multiplication and the necessity for swarming.
. Mr. C. F. M. Swynnerton devised a most effective way ,of
dealing with them, A small quantity of cyanide was crushed and
dissolved in hot water and squirted over the swarm with a syringe
after dark, with the result that they fell like a shower of rain
EAST AFRICAN INSECTS. 1023
and only a few individuals would ever recover. For greater:
effectiveness the swarm would be enclosed as much as possible
with matting supported by poles.
47. BeLonocAstER enrisrus Fabr. (Vespide : Diploptera)..
These wasps, whose paper nests are to be found pendent from the
verandah ceiling of most Hast African houses, were originally
inhabitants of caves, I suppese. One day I came upon a huge
overhanging rock—almost a cavern—from whose roof were
suspended at least thirty large nests of this species; the half-dozen
workers hanging to each assumed threatening attitudes in.
characteristic fashion at my approach. On my going still nearer,
one flew off and stung meon the chin—a thing they rarely do in
houses, where they are more used to human beings. The pain
was not very bad, though the resultant swelling caused several
kind enquiries as to whether I had toothache. (Mt. Longido,
13. ii. 16.)
In climbing a Flamboyant Tree to reach a dove’s nest I had
the misfortune to disturb the wasps on a nest below me. _ The-
first intimation of this was the receiving of a sting on the calf of my
lee, six inches below the knee. I drove the wasp off, but in doing
so received another sting on my first finger. The sting on the
leg was bleeding, and looked as if’a piece the size of a pin’s.
head had been bitten out. Putting on my puttees I went off for a
four hours’ walk, feeling scarcely any inconvenience. On removing’
the puttees at one o'clock I noticed that the leg had been trying
to swell under its wrapping. By 2 p.m. I could scarcely walk,
and the limb was very swollen. At 6 p.m. the poisoned leg
measured 142 inches as against the 13% inches circumference of
the other. Next morning it was 15 inches. I had no idea that
a, wasp’s sting could have such bad effects. The finger swelled a
little, but this subsided the same day ; it was stiff and a. little:
sore on waking the following morning, but had evidently had
5
a much smaller dose of venom. (Morogoro, 15. vi. 17.)
48. Synacris mstuans I’, subsp. RuFA Stadelmann (Humenide)..
T came across the nest of this species for the first time to-day. It
consisted of two mud-cells, roughly 35x20 mm. in length and
breadth. These were side by side on the under surface of a leaf
and attached to the mid-rib. The leaf measured 700 mm. in
length, but by reason of the weight attached, hung vertically.
The leer cell was already sealed ; the wasp was resting in the
upper one with its head outwards, completely closing the entrance ;
this was at 4.15 in the afternoon, the weather: being dull and
eloudy. (Kilosa, 25. 111. 2)
49. PsAMMOCIIARUS VENANS Kohl, @ (Pompilide: Fossoria)..
My wife, who was preceding me down a rough path, suddenly
jumped from her cycle and zalled me to sce “such a large spider:
being stung by a wasp.” : As I reached the spot the spider was
standing on its anterior legs; it might have had six applied to the.
1024 MR.-A. LOVERIDGE: NOTES ON
ground, but my impression was four, the remainder being in the
air and the ventral surface of its abdomen so tilted as to be visible
to me standing above. ‘The ventral surface of this spider (Ctenus
sp., 2: Clubionide) is a very bright scarlet with a black basal
patch, ornamented by two very white spots which, I take it, serve
as eye-spots, ‘lhe spider was a female [ believe, as I aim familiar
with the species, having taken one only half the size of this
specimen the previous week.
The wasp, which had flown off when first disturbed, had now
returned, and was running hither and thither between the
f=)
stationary cycle wheels looking for the spider; when close to its
victim the latter bolted with great swiftness across the road,
absolutely abandoning its “ warning” or ‘terrifying attitude.”
Swift as it was, the wasp was swifter. I could not say if it ran
or flew along the surface of the road—probably a combination of
both; on overtaking the spider it pounced upon it with the
greatest ferocity, curved its abdomen under, and stung vigorously ;
they rolled over in the road, and as I had nothing but a pen-
knife wherewith to capture them, suntorbinntely they were
somewhat damaged. (Kilosa, 28.11. 22.)
50. DoLicnomuUTILLA GUINEENS!S Fab.,f. auratA Bischoff(Mutil-
lide). Rolled in my blankets under a bush, and reading by
candle-light, I felt an insect crawling up my leg. Supposing it to
be an ant I attempted to brush it off, when I felt a sharp sting,
whose eflects travelled quickly up my Jeg to the hip. The pain
lasted 48 hours and was like a bad nettle sting, leaving a red and
lumpy rash. (Handeni, 25. vi. 16.)
At Morogoro and Kilosa these wingless females were constantly
found in the house... At 4.15 p.m. one having crawled up my
leg, got caught between my stocking and slipper. It stung
me on the upper surface of the foot about 13 inches behind the
little toe. Vor ten minutes the pain was frantic—far worse than
a bee sting. An hour afterwards all pain had disappeared, but
treatment had been adopted by placing a small crystal of
poe permanganate on the site of the sting. (Iulosa,
7. vi. 1.)
Ifurnroeyna (Ants).
The actual specimens referred to were only received for
Nos. 52 and 53.
51. ? Paurormyrevs rarsatus IF. or Mecaponera rarens F,
A: flight of termites had. just taken place after heavy down-
pours of rain, and the termites were busy discarding. their
wings, when I noticed one of these ants carrying off a termite ;
another ant was carrying a spider, whilst a third was laboriously.
dragging along a stick-insect many times its own bulk. eee
West, 30.1. 16.)
Passed a dozen o more companies of the lar ge black Benentae
ants which wander across the road in a fusiform body, quite’
_ EAST AFRICAN INSECTS. 1025.
unlike the well-ordered columns of the siafu. Neither do they
attack anyone who disturbs them, like the siafu, but, instead, run
hither and thither, making a hissing noise not unlike that which
asnake might make when rustling away among dead leaves. Per-
haps it is an imitation which serves their purpose. (Kerogwe-
Handeni Rd., 6. vii. 16.) Later experience of these ants makes
me wonder why they were in fusiform formation when I made
the note, for generally the hosts are in column formation, unless
I am confusing two species.
52. Patroriyreus tarsatus F. I have already described the
combat of this species with siafu, and have recorded elsewhere
how they attack the eyes of tortoises, causing the poor beasts to.
die *, Water-tortoises, however, when in their tank readily seize
and eat Paltothyreus, as did my Wennec Fox (Otocyon virgatus).
of his own accord when one of these ants emerged from a burrow
near him. (Kilosa, 20. ii. 22, 25. ii. 22 & 10. iv. 22.)
I was throwing out some of the peppery seeds of a paupau fruit,.
when I noticed a sudden activity about. the spot where they
were falling, and to my surprise saw fully fifteen stink-ants.
carrying them off. Whether the ants had already been on
the spot or had emerged from their holes on scenting the paupau
I cannot say. They soon found the paupau in the tortoise
enclosure and commenced carrying off the remaining seeds, so
they are not wholly carnivorous in their diet. (Kilosa, 15.iv. 22.)
03. Myrmecoruites of PaurorHyrREus TARsATUS. A number:
of these ants were drying their pupe and larve on the path
after heavy rain; on my appearance they started hurrying
away, and I was astonished to see scores of Lepisma-like insects.
(probably Ctenolepisma sp.) of all ages hurrying after them
and running in and out amongst them, also a small beetle
(Onthophagus pugionaius Boh. : Copride) in fewer but consider-
able numbers followed the ants in company with the Lepisma ;
when touched, these beetles shammed-death. Mr. Arrow, who.
kindly named the Orthophagus, said that it was a common species,
and suggested that its presence with the ants might be accidental,.
but there was no doubt about the existence of a true association.
(Kilosa, 16. ii, 22.)
D4, CREMASTOGASTER CASTANEA Sm., var. TRICOLOR Gerst, A pin.
with an atom of food on its point was lying on a ledge, when I
noticed five of these ants take hold of it and carry it to the
edge of the ledge, One ant appeared to be supporting most of
the weight, having gripped the pin near the point. I therefore
chased off the other four ants and timed the fifth, which supported
the whole weight of the pin for fifteen minutes, when I had to leave.
(Kilosa, 30. xii. 21.)
These ants, which are a perfect pest in a food cupboard, being
fond of meats as well as sweet things, lately started eating.
i Proc. Mnt..Soe. Lond, 1922,.p, xli.
1026 MR, A. LOVERIDGE:: NOTES ON
away a cork to get at the raspberry syrup which the bottle
contained. 1t was not a question of eating the surface of the
cork which might have been in contact with the syrup, they
ate right into the heart of the cork and then downwards.
(Kilosa, 15. iv. 22.)
On sitting down to breakfast I noticed one a these ants
walking round and round the stem of a vase. I commenced to
count how often it did the round, and after 39 times, found it
took two minutes to do 29 rounds; at the end of five minutes it
seemed to be going as strong as ever, so again timed it, and found
it did a little more than 143 rounds per minute. At the end of
ten minutes, as we were timing it again, it began to vary the
tour by turning about, and only did 94 rounds to the minute,
Tt had, however, already done some 150 rounds, and, falling back
into its old stride, continued for another five minutes, making
‘in all a quarter-of-an-hour since we began to time it; during
this time it had accomplished approximately 200 rounds of a
‘circle 4} inches in circumference, giving us an idea of its accom-
‘plishments when in column, of 96 yards per hour or more probably
100 yards per hour.
The ant then came down on the table-cloth, but soon returned
to the groove and recommenced its absurd parade, which it was
still doing when I returned to lunch four hours later. It had,
of course, come in on the fresh-cut flowers and was far from
home and friends, but why should it exhibit so little intelligence
as to wear itself out in this manner? Was it the scent of its
own feet that hurried it along, thinking it was on a regular run,
-or was it the reflection of itself in the. highly-polished concave
groove that made it fancy it was accompanied by others? I
cannot say, but that it was not a foolish individual is proved by
‘the fact that whenever flowers are brought in with this species
upon them the same thing happens, sometimes as many as four
ants taking part in the procession, frequently going in opposite
directions and moving out of each other’s way when they meet.
-(Kilosa, 18. v. 22.)
5. CARDIOCONDYLA EMERYI For, An ant which causes a great
deal of inconvenience to householders I call the ‘Sugar-ant” from
its fondness for that commodity in particular, and sweet things
in general. In my present quarters they swarm on the bed,
table, and cupboard and many of them die in the jam; they cross
-one’s paper when writing, and have to be brushed off one’s
neck, but never retaliate. (Morogoro, 14. xii. 16.)
It is interesting to note that the sugar-ants which formed
long columns to and from my cupboard are now absent; only a
few individuals are to be seen each day. (Morogoro, 20. iii. 17.)
56, PuagioLeris (ANOPLOLEPIS) CUSTODIENS Sm. This species
is very abundant both at Kilosa and Morogoro, where it will be
‘found running about in incredible numbers on paths and open
ground and sometimes swarming on tree-trunks, They attack
EASY AFRICAN INSECTS. 1027
‘any small insect which they can overpower, and to-day I found
a small toad (Bufo reqularis) being actually dragged along by
-a host of these ants, which so obscured their burden that until I
had picked it up and shaken them off T could not tell what it
was. ‘hese ants cause the natives considerable annoyance by
biting their bare legs. (Morogoro, 12. ii. 17.)
57. Gicopmyiia smArAcpina F. This long-legged, somewhat
transparent-bodied, yellow ant fastens leaves together to form its
nest, and should one brush against the branch bearing the nest,
the occupants rush out and stand quivering all over it, whilst
-others hurry off with open jaws to seek the disturber of their
peace and soon cover all the foliage; when they get upon a
person they bite savagely, more often on the neck than elsewhere.
‘Their favourite tree is the mango, upon whose fallen fruits they
feed; they also eat paupau; I have never seen them carrying
insects. hey nest in Javanese silk-cotton trees, which bear
leaves that seem to be suitable for their purpose. (Kailosa,
2k We Pa)
58. Dorytus (ANomMA), probably nigricans Illig. The siafu,
to whom I have referred elsewhere *, have few enemies. One
morning I shot one of those timid creatures, an elephant shrew
(Petrodromus nigriseta), beside a column of ants, which were
-crossing a bush-path at 6a.m. To my surprise, on opening
the stomach of the shrew, I found it to contain a number of
these pests. (Morogoro, 12. vi. 17.) :
59. Dorytus uELyotus L. On no fewer than three occasions
my attention was drawn to amphisbenids (Monopeltis colobura),
which had been attacked underground by these fierce ants and
so tortured that they came to the surface, on one occasion at
9.15 p.m., when the sun beat fiercely upon the scorching sand—
conditions which these burrowing lizards detest. ‘This species
-of ant will attack any freshly-killed body from underneath, but
‘appears to hate the light, and never exposes itself. (Lumbo,
eels)
60. Camponotus (ORTHONOTOMYRMEX) sERIcEUS F. One of
these greyish ants, which are very common in the house and seem
to be much addicted to jam, was carrying a spider along the floor.
I picked up another carrying a small chrysid. (Kilosa, 1. xii. 20.)
61. Camponotus sp. I believe it was one of the many species
of this genus which disturbed me when lying down beneath a
giant tree to sleep, rolled up in my blanket; they invaded me
in hundreds and were about half an inch in length. Unfortu-
nately, when turning over, I imprisoned one between my cheek
and pillow, and received a severe nip. (Ngeri-ngeri, 24. viii. 16.)
I noticed an ant in my tent to-day which appeared peculiar,
-and on examination it proved to have the head and thorax of
'* Proc. Ent. Soc. Lond. 1922, p. xxxiis
1028 MR, A, LOVERIDGE: NOYES ON
another ant of the same species attached to its antenne by the
jaws of the dead head. Presumably the wearer of this ornament
had bitten his comrade in half during some argument. (Moro-
. goro, 30. vi. 17.)
COLEOPTERA.
‘The actual specimen referred to was only received for No. 65.
62. CiciNDrLA BREVICOLLIS Wied. A number of these beetles.
were collected on sandy ground and paths by the river-bank ; they
harmonized so well with the ground that they were very
difficult to see. ‘Chey were also extremely active, taking to wing
on the slightest provocation. (Morogoro, 14. xii. 16.)
63. A tiger-beetle apparently of the same species, flying in front
of me, kept settling motionless on the path. I, thinking it wasa
tsetse, wondered what it was doing there, seeing we were in a
cultivated area of some extent, so jumped off my cycle and then
saw that it wasa beetle, I stalked it, and shot it with a garter!
(Kilosa, 1. vii. 21.)
64. Anruia srriatopunctTata Guér., ¢. This large Carabid
Was seen running acrcss the are of light in front of my tent
about dusk. In attempting to pick it up, [ thrice felt a coolness
upon my hand—the fine spraying of a liquid. Just then my boy
arrived with a killing-bottle, and as he knelt to seize the beetle,
it discharged this secretion full in his eyes, which were at least
eighteen inches away, I believe, The pain caused was agonizing,
and though bathed immediately, his eyes became very bloodshot
and swollen, and so remained for three days, when the discomfort
of the protruding eyes had practically subsided, though they
were not normal for some time afterwards. The odour was
very powerful, flying to the eyes like formalin*. The beetle
is quite common in the district, (Izikisia, Tabora Dist.,
Alen exay 109)
65. Poritnra LiavuLaTaA Ohs., 5 (Rutelide), Among a series of
these beetles collected by Salimu during the past week was a
Rutelid whose right meso-leg was missing—had never grown. On
the opposite side the middle leg had three feet with one base—
that is to say, the beetle had five legs on the one side and only
two on the other. (Uluguru Mtns., 1. iii. 1921.)
66. Myapnris ocuLAta var. tricoton Gerst. (Mylabride: THete-
romera). In my noteon the ovipositing of this beetle which was
kindly communicated by Prof. Poulton, he drew attention to the-
typescript being indistinct as regards the fraction of an inch
expressing the length of the eggs (Proc. Ent. Soc. Lond. 1921,
p- xcii). This should read one-eighth of an inch, and not one-third,
. * Compare Dr. Carpenter’s experience with Anthia fornasinii Bert. (Proc. Ent.
Soc. Lond. 1918, p. c) and his experiment on a monkey with A. striatopunctata
(rans. Ent. Soc. 1921, p. 10); also Dr. Marshall’s experiments on monkeys with
beetles of this genus (Trans. Ent. Soc. Lond. 1902, pp. 610, 611).
BAST AFRICAN INSECTS. 1029
67. MyLasris picinctA Berto]. In this connection 1 came
across a note to the effect that this red-and-black beetle was
excavating a burrow on a native path through the mealie crops.
(Morogoro, 9. vii. 17.)
68. Mynabris sp. A number of Alylabris beetles with
searlet-inarked elytra were feeding upon some Cucumis plants
on May 29th and again to-day. On both occasions there were
about eight to each plant and none on the surrounding herbage.
The inter esting thing about them was the way the males (which
were much smaller than the females) followed the females up and
down the stem or leaf as the case might be, stroking their elytra
and abdomens with their antenne. Both antenne of the male
would be raised simultaneously, and with these he stroked the
female most caressingly. The larger beetle fed unconcernedly or
walked about, followed closely by her devoted attendant. (Ngong,
Nairobi, 3. vi. 15.)
LEPIDOPTERA.
The specimens referred to were received for Nos. 70, 75, Go.
77, 78, 80, and probably 73, 74, and 79.
ane never gets tired of the wonderful butterfly fauna of Kast
Africa ; and between Mt. Kenia and Mozambique at one time or
another, the writer has seen extraordinary assemblages of them
in almost inconceivable numbers and variety, but nothing ever
surpassed the display on 17. iii. 16, when they were feasting on
the dead transport animals left stranded on the wayside by the
tide of war.
On leaving Moshi for Kale we passed through miles and iiles
of rubber, coffee, and other plantations. It was a glorious
though somewhat steamy day after heavy rain. Never in ny
life have I seen such thousands of butterflies, all apparently
freshly emerged, as they were in the pink of condition. There
was one patch ef Papilio demodocus about eighteen inches in
diameter and containing about one hundred closely-packed insects
feasting on the intestines of a bullock. On a raw buckskin there
was a big crowd of blues, and every puddle in the road was
surrounded by numbers of them. (Kahe, 17. iii. 16.)
69. HuRYTELA HIARBAS Drury. Came upon a bush to which
« number of Cetoniids were clinging; one of these was on the
lower side of a horizontal branch, and immediately above him
was the Nymphaline /#. hiarbas, which was applying its proboscis
to a spot of moisture on the twig immediately above the beetle’s
head ; each time it protruded its proboscis to do so, the beetle
relaxed its grip with one of its pro-legs and waved it in the
direction of the butterty ; replacing the leg, it repeated the action
with the opposite leg; the object of this was to drive the butterfly
away from the exuding sap. Returning later I found no fewer
than three beetle- -butterfly groups, and concluded that the
Proc. Zoou. Soc.— 1923, No LXVII. 67
L030 Mh, A, LOVERIDGE: NOTES ON
beetles scarify the twigs to obtain the sap, and resented the
butterflies partaking of the fruit of their labours, Later in the
afternoon a larger Cetoniid was seen in a tree going through the
same performance with a Oharaxes. (Karura Forest, Nairobi,
21, vii. 19.)
70. Teracotus casra Gerst., Q (Pierine). Took a butterfly
to-day with one pair of wings fully developed; the other pair
still in the chrysalis. The butterfly was vainly flapping its fore-
wings. , (Kilosa, 13. xii. 20.)
71, CaropsILiaA FLORELLA Fabry. (Pierinw). Besides the migra-
tion of this butterfly noticed on 3.v.15, another migration
lasting several days was witnessed in 1919. Ju both cases the
buttertlies were going in a north-westerly direction. In the
second migration the wind was strongly in their favour, but if 1
recollect aright, adverse on one somewhat cold day *. (Nairobi,
v. 19.)
72. Parinio pemobocus Esper. Watched this Swallowtail
curving its abdomen under the leaves of a lime-tree for purposes
of oviposition. It did not seem easily satisfied in the selection
of a leaf, and the eggs were laid singly. (Kilosa, 28. xi. 20.)
Took a very young larva which has a wonderful resemblance
to a bird-dropping. (Kilosa, 29. xi. 20.)
Very common 77 cop. just now. (Kailosa, 18. xii. 20.)
Trom later observations I have come to the conclusion that
the species breeds all the year round, though principally in the
rains. Larve were taken at Durban on 21. ‘xii, 14.
73. NEPHELE PENEUS PENEUS Cram. When the rubbei-trees
are in bloom hundreds of these hawk-moths are to be seen
flashing about the tree-tops, where they are difficult to obtain.
Later in the season they come to the paupau flowers, and the
only way to net them is by cutting off all the leaves of the
paupau-tree. They are not nocturnal, strictly speaking, but
crepuscular, becoming active again about 5 aim. (Kailosa,
31.v.22.) Elying with them was the much rarer Vephele bipartita
Butl.
74, Acrias Mimos# Boisd. (Saturniidie). Discovered the cocoon
of a Queen Moth, which is one of the prettiest things in the way of
cocoons. It was spun on the lichen-covered twigs of a fir-like
tree. It is of a very silvery appearance, and if a section be cut
out, the texture will be found so strong that it is impossible to
tear. There are small breathing-heles at the top end, just beneath
the opening from which the moth emerges. At the lower end
are a few more holes, possibly for the sake of a current of air or
to carry off any moisture. (Makindu, 31. vii. 16.)
* Compare the records of migration of this species in Proc. Kut. Soc. Lond. 1921,
p. Xx. :
HAST AFRICAN INSEC'S. 10381
Of four live cocoons received yesterday from Natal a male
emerged at 2 p.m. this afternoon. By 3 p.m. it seemed to have
finished developing the wings, whose monstrous tailsare 3 inches
in length. At 4 p.m. it opened the wings and rested with them
flat—the resting moth had a most striking appearance. (Nairobi,
23. vill. 19.)
At 9 p.m. I heard a crackling noise from one of the remaining
cocoons, from which I had cut a small section the better to
observe the pupa within. The latter was swollen, making the
abdominal rings very noticeable. I watehed the moth climb
from the funnel and withdraw its abdomen very suddenly ; after
recovering from its exhaustion, it adjusted its position on the
outside of the cocoon. With the exception of the mauve costa,
the wings appeared to be entirely orange-yellow at this stage; a
pale green blush began to suffuse the wings, commencing at their
basal parts. ‘The green spread and deepened in tone until it had
replaced the greater part of the yellow. On emergence the tails
were very short—a quarter of an inch perhaps—and were the
last part of the wings to develop. At 11.45 p.n. I was awakoned
by the moth flying round the room, and on turning up the light
found a second specimen had emerged and was nearly dry.
Both were males, (Nairobi, 27. viii. 19.)
75. RiropoeasrriA virneA Plotz (Arctiine). When walking
up the path at 4 p.m. I disturbed a hunting party of the [lelmeted
Shrikes (Prionops talacoma), which flew into a large tree over-
head and recommenced their researches there. Immediately
afterwards a fine moth in emerged condition dropped down on
the path, almost at my feet. I poked it with a twig, but, as
it did not move but shammed death, tried to pick it up by one
of its vivid red legs, whereupon it began to exude a bright
yellow fluid from both ‘‘ shoulders,” this fluid coming out as a
froth with a marked rotary motion. I passed-the twig beneath
the moth, hoping it would clamber up it; to my surprise it
seized the twig with its hind-legs only, and, as I raised the stick
in the air, it picked up two lumps of quartz from the path, the
larger lump measuring 10x15x6mm. As I carried the moth
along with me, it carried the quartz for fully a minute before
dropping it, and for yet another minute it remained supporting its
own weight by its hind-legs. The pillars of froth stood up well
from its shoulders, being about 3 mm. in diameter and 8 or
10mm. long. Suddenly the froth dissolved into drops, which
fell upon my hand, and the moth with rapid darting flight made
off. The fluid smelt to me like the secretions of the cloacal
glands of a British Grass-snake, which is also used as a defence *.
(Kilosa, 8. vi. 21.)
* Dr. Carpenter has observed the same method of defence in an allied species,
R. leucoptera Hmpsn., and has recorded the acrid odour and taste of the secretion
(Proc. Ent. Soc. Lond. 1918, p. xevii).
67*
1032 Mk. A. LOVERIDGL : NOLES ON
76. MaxeraA mMarcHatt Boisd. (Ophiderine : Noctuide).
Whilst in the garden my wife’s attention was drawn to some-
thing moving amongst the dead leaves. A rolled-up leaf
19x8 mm. was making progress by a series of little jumps.
Placed on a tablecloth, it journeyed first in one direction and
then in another quite aimlessly. On opening the leaf a flattish
moth larva was found. (Kilosa ,12. iv. 22.)
On opening the leaf I found the maggot-like larva had pupated.
(Kilosa, 24. iv. 22.)
On going to the box in which I had placed the chrysalis,
I was sorry to find the small brown moth had emerged some time,
as it was already dry; the time taken for development was thus
extremely short. (Iilosa, 5. v. 22.)
77. ANAPHD RETICULATA Walk. (Notodontidie). Last November
T found the cocoon of a wild gregarious silk-moth. ‘The cocoon is
a khaki-coloured, strongly-woven structure 190 mm. (74 inches)
in length and 110 min, (4} inches) in diameter at the broadest
part. Through its axis passes a branch of the food-plant, whose
twigs and leaves are incorporated with the mass of silk. At the
upper end are two chimneys or funnels protruding 20 mm. (# in.)
from the surface of the pear-shaped cocoon. ‘Through these the
moths emerge at dusk and for two hours afterwards. I never
recollect seeing any emerge after 9 p.m., though doubtless they
occasionally do so.
The business-like hurry with which the moths eimerge, as if
accustomed to do so all their lives, is quite amusing. The head
of each moth as it emerges is protected by a little helmet of
shamrock-shape, each of whose “ leaves” is deeply concave, ‘This
helmet is, of course, part of the head-piece of the chrysalis, and
serves to protect the eyes and delicate head of the moth as
it pushes its way out of the funnel. As soon as the head
was clear of the funnel, up went a pro-leg and gave a push
to the mask, which fell to the ground; quite a heap of these
were found each morning almost immediately beneath the
funnels.
Moving hurriedly and crabwise the insect then travels from the
funnel to the under side of the cocoon, where it hangs whilst its
crumpled and draggled wings open out, develop, and take shape,
the brown lines on the creamy ground becoming clearly defined.
The wings were then raised above the back in the position of
those of a butterfly; from time to time they were shivered,
apparently to test their muscles. After hardening, they are
brought to the usual position of a moth at rest, lying one across
the other upon the back, forming approximately a triangle. If
disturbed at all the moth discharges a pinkish excretion, and
this fluid leaves with such force that I should think it carries a
distance of quite nine inches from the moth.
Theoretically this is the middle of the rainy season, but there
have been no rains recently; nevertheless three of the moths
BAST AFRICAN. INSECTS. 1033
emerged this evening. J will give the rest of the dates of
emergence serially, with the numbers of moths which emerged
from the cocoon each day. (Morogoro, 18. iii. 18):—.
MomchveliQthy-es.ce: 9 March 25th oe)... 40
eA DOG HAW wees 133 ay Ob eee 56
aN Shaan 10 Bat AAT AAR eer 22,
AE ROTC ea: 13 Bud tosh eran 8 9
A TCA R errs 16 BRU AO tne aha 10
Ren ED AN eee 25 Ran EO AA Gun wis Sede ny 4
After this not more than ones or twos emerged at long
intervals and often crippled, the total not exceeding ten. If we
assume there were ten, it gives the astonishing total of 240 moths
emerging from this one cocoon. (Morogoro, 30. iii. 18.)
Found a second cocoon on a small shrub and, assuming the
moths had emerged, put it awayinatrunk. (Kilosa, 30. xii. 20.)
On opening the trunk I found some 14 or 16 had recently
emerged. (IGlosa, 10. iii. 21.) From this date the following
emergences took place : -— :
Marche thee. ease. 4. Marcela lithe ene 1
bis ale Sl Waa] ras tee eee 2 ce iaetal cca hye eaesec: 2
Pim maleS GU er apeacccs 2 ea MOG gates nil
Saul dhe Nel avhs Same eeaiates 2 ee Teed 0) sl naar tan 1
Hs eel So git, Seana ead 2 BR Ki ey 5 aa 1
This cocoon therefore only produced one-seventh the number
of the former, which was quite twice its size. ‘The total number
emerging was 32. (Kilosa, 24. iii. 21.)
Thinking a record of the incidence of sex among those that
emerged would be more interesting than the bare totals, on
receiving a nest from the Tabora District brought back by one
of my native collectors, the results were tabulated as follows :—
December 26th ....... ks POR
A Dik eames Dn, 6 dS Qk?
hs ASIEN Sp ogsee or 3 6 & @
a DANEEL, cscs gc 2, 5 &?
Janay aa Oblhyy snes ee ben.
sda pial Othe cen ies my Oe eo 6 Oo
“ UD We Se eel ey SIS S ELL VS Eo
Le id Oth salt i, @€2O2 O06
is Sth eae ee 2.99
af Weed Magadaoodes Lee
a IARI ee eds 3, 2 2 Y (one was a cripple)
= USthi se ee. 1.2
es LG Ghose ae ane 1, 92
Total emerged 47, composed of 2 not sexed, 19 males, and
27 females. As one would expect, males preponderate in the
first days of emerging and females very markedly towards the
1034 MR. A. LOVERIDGE: NOTES ON
end. ‘There is a curious gap between December 29th and
January 9th, when none emerged ; this was not to be accounted
for by the weather, which was almost uniformly hot and dry
throughout, though I believe one or two showers did occur.
(Kilosa, 19. i. 22.)
78. An AmypriA and another Tineid too worn to be identified ;
also Hphestia cuutella Walk. (Phycitine ;Pyralide). Writing from
an ex-enemy cotton warehouse where I was camped for a few
days :—‘‘ There are some hundreds of clothes-moths hatching out,
in this building every day, presumably having bred in the large
stock of cotton which the Germans have housed here for two
years past. Iwas very interested in seeing a dozen males
assembling on the outside of my mosquito not and moving to and
fro in ceaseless agitation over a square inch of net—not till then
did I notice the female on the inside of the net. Killed the
males. Next morning there were some more, so I killed the
female and drove off the males, but they returned to the spot
and continued to move over it for probably, half-an-hour after she
had gone.” (Morogoro, 19. iii. 17.)
79. Dreara (Jana) sp. (Hupterotide), The large and hairy
caterpillars of this moth are from three to four inches long, and
their clothing of fine hair over an inch in length makes them
look inviting to stroke. ‘The white or greyish hair, however,
conceals sharp rufous-coloured spines arranged around the body
segments. If you attempt to pick up this caterpillar the spines
are driven into your flesh, where they remain, being very brittle.
Fortunately they are not barbed, so they can be drawn ont,
though it is difficult to do so as they snap on account of the
afore-mentioned brittleness. In endeavouring to pick up one of
these larvee to-day T utilized a leaf of a rubber-tree to protect
my fingers, and was astonished to find the spines driven through
the leaf and into my finger. Unless poison was removed in
passing through the leaf, [ do not think they are poisonous,
as only mechanical irritation was felt. The leaf bristled with
them, and a piece of paper proved to be no better protection.
When interfered with, the caterpillar half curves its body, which
has the effect of bringing the spines into prominence and making
them more rigid, I think. (Kilosa, 25. 111. 22.)
One of the caterpillars cast its skin yesterday and the other
to-day, though collected independently. (Kuilosa, 7. iv. 22.)
80. PRockssIONARY CATERPILLARS. I came across a proper cater-
pillar procession for the first time to-day. A clump of the
caterpillars, owing to their long, wavy, white hair which almost
obscured their dark green skins, bad the appearance of a large
luxuriant patch of American blight. The clump was about four
feet from the ground on the bark of a tree, and reaching to the
ground was a single line of caterpillars; another line led up from
the clump. Each member of the procession had its head closely
applied to the “ tail” of its predeeessor, and they moved forward
BAST AFRICAN INSECTS. 1035
very slowly, after the fashion of London taxicabs closing up in a
press. The caterpillars were 14 inches in Jength. (Handeni,
97. vi. 16.)
Some six miles from Kilosa along the railway line for Kigoma
is Jumbe Sunguru’s village. Here close to the line is a mighty
wild fig-tree, big as an English elm, and as I was passing it to-day
J saw that every bit of this great tree as far as its topmost twigs
was smothered in grey hairy caterpillars, each bearing on its
back two spots— tufts of black hairs in reality. The caterpillars
seemed to be moving slowly up the tree with long silken threads
in great profusion as if to guide them, and as I walked around
the tree surveying this wonderful sight from different points, I
came upen a small column travelling with haste across the path.
Following this line back, I found they were descending from a
smaller tree—no larger than a crab-apple, and of much the same
appearance—which still had plenty of leaves, so why they should
leave a Jand of plenty and go on this pilgrimage to a tree already
much depleted of foliage and overstocked with caterpillars, I
could not understand.
T could not see the caterpillars feeding on the leaves, but a
gentle rain of droppings kept falling, and the surrounding ground
was already well-covered, whilst unsightly masses of web and
silken thread loaded with excrement depended from the larger
branches. It was difficult to see what the main body on the bole
of the tree would get to eat to-day, but for the time being they
had something else to distract their attention. Several minute
black ichneumon flies were hovering over them, and when one
of these approached a. caterpillar, the latter would jerk its head
and forepart of the body most violently backwards, and the little
fly would move off. How it could get close enough to oviposit
an egg in the skin of such a hairy caterpillar was the problem
confronting the insect, but doubtless it would eventually succeed
in doing so. (Kilosa, 30. vi.21.)
On August 6th I revisited the tree and collected four pup
which were concealed beneath a mat of webbing. Two were of
one species and two of another.
‘T'wo pup were green, but the shells left after emergence were
transparent and colourless except for their bright red “ tails.”
They produced ochre-coloured moths with a reddish spot on each
fore-wing—WVygmia crocosticta Hmpsn., or a race of this species,
being smaller and more richly coloured than the type which
came from the west of Lake Nyasa. (Iilosa, 14. vii. 21.) These
moths are Liparide (Lymantridz), a family not known to include
processionary larve. ‘They may have accidentally pupated in the
web of the true processionaries or were perhaps definitely asso-
ciated with them.
The two other pupe were brown with four longitudinal series
of golden-brown spots. They were 17 mm. in length over all,
except the terminal spine which was a millimetre long. One
moth either failed to emerge or was lost. ‘The pupa of the other
1936 MR. A, LOVERIDGE: NOTES ON
was submitted to Mr. Tams who found that it contained a
parasitic pupa which Dr. Waterston identifies as a species of
Brachymeria (= Chalcis auct.), An anterior prominence on the
moth pupa resembles that found in the Notodontid genus
Thawmetopea, known to have processional larve. The species
may have been 7’. apologetica Strand, from Hast Africa, or
tin allied form,
ORTHOPTERA.
The specimens referred to were received for Nos. 83, 85, 86,
89, 91, 92, 93, 94a, 95, and 98.
81. Orrimoprrra have many enemies. In the stomach of a
Mungoose (A/ungos nuungo colonus) shot to-day were cockroaches,
locusts, grasshoppers, and a number of other insects. (Morogoro,
IY, st, Uo)
A. Kestrel’s (Cerchneis tinnunculus) stomach contents showed
it to have been feeding chiefly on locusts. (Morogoro, 4. xii. 17.)
A Cattle Egret (Bubulcus ibis) had a great number of grass-
hoppers in its stomach. (Morogoro, 14.xi1.17.) |
GRYLLIDA,
82, Bractty¢nyprs MEMBRANACEUS Dru. My first acquaintance
with this huge cricket was at Morogoro, where an oflicer of the
Flying Corps brought me a specimen 23 inches long and ? broad
at the widest part “he rather aptly compared its song to the hum
of a gnome engine. (Morogoro, 3. v.17.)
One of these crickets was shrilling away beneath a shrub two
nights ago: my boy located it, but it dived down its hole; he
made a big excavation, but missed it ; last night presumably the
game cricket was under the next bush twenty feet away. ‘l'o-
night it was at the back of the house a hundred feet from its last
location. I went to see it, and found it was just beginning a
burrow into which it dived, but was easily dug out. It had a
single egg in the oviduct, not round but slightly pointed at each
end. (Kilosa, 12. iv. 22.)
Again, to-night I found a cricket shrilling outside its burrow,
and was able to appreach within a foot of it with an acetylene
lamp, and stand some time before it made off. It also was a
female with an egg ready for oviposition. Within a few inches
of it was a gecko (Hemidactylus squamulatus) perfectly motion-
less as if dazed by, or enjoying, the noise. (MKilosa, 13.iv. 22.)
Each morning one sees little heaps of excavated sand about the
camp, and near them are tracks, not unlike those of a rat at first
glance, which show the extent of the insect’s nocturnal peregrina-
tions. At dawn, and for about an hour afterwards, it is not
uncommon to find stragglers still above ground. If dugout they
ave usually not more than a foot below the surface. Lately I
have been feeding the small Mungoose (Helogale ivori) wpon
BAS! AFRICAN INSECTS. 1037
them, as they show much eagerness for this form of diet, snapping
them from one’s fingers.
83. LIoGRYLLUS BIMACULATUS de Geer. ‘This species is very
widely distributed throughout East Africa, At Mkomasi I dis-
covered two crickets beneath an old battered biscuit tin; one
was already dead and dismembered, the other in the jaws of a
scorpion (Odonturus dentatus Karseh), which was busily engaged
sucking its juices. As 1 was pulling the cricket away from the
scorpion a Sanil-snake’s head was protruded from the tin.
(Mkomasi, 2. vi. 16.)
TEVTIGONIIDH = Locustiva.
84. ENyAuiorsis sp. I caught two of the wingless long-horned
grasshoppers which are very common here. 1 woke up with
one on my face the other night, and several of my comrades have
found them in their blankets. (Dutumi, 20. ix. 16.)
85. Larva of a species of Grytuacrin/. On my table for a
week past has been lying a newspaper cutting—a portrait. This
morning [ found it mutilated in circular fashion as might have
been done by a rose-cutter bee. The cut-out portion had been
turned up flap-wise and carefully cemented down to form a. little
eell without entrance; neither was there any trace of cement or
silk. Opening this I found within it a pink larva. (Lumbo,
25.1x. 18.)
86. Cronia wAuLpencr Stal. A native brought me a specimen
of this peculiar insect, the like of which I had never seen before.
(Morogoro, 11.1.17.) ‘Iwo were subsequently taken at Kilosa,
where they had apparently come to light. (IKilosa, 18. xii. 20
and 18.1. 21.)
87. PskUDORHYNCHUS PUNGENS Schaum. Heavy rainfall last
night and to-day ; the countryside is teeming with these Ortho-
ptera. ‘They were just as thick as locusts, in a rubber plantation
near here. One had to shield the eyes to protect them from
the storm of insects which flew against one’s face and body. The
brown forms were greatly in the minority, being about 5 per
cent, ‘They showed no selection in choosing sites, as they settled
on bright green rubber-leaves just as readily as in the dry brown
grass. (Kilosa, 1. xi. 20.)
The insects are plentiful here also—z.e., some six miles from
Kilosa. (Miombo, 4. xi. 20.)
ACRIDIDE.
88. LamancKiaAnaA sp. I put one of these cryptically-
coloured grasshoppers into a tin as food fora large Mygale spider
(Plerinochilue sp.: Avicularidz), but on opening the tin found
that the er asshopper had killed the spider by biting its abdomen,
(Nr. Moshi, 17. iii. 16.)
1038 MN. A. LOVERIDGE: NOTES ON
89. Scnisvocera GrecaRniaA Forsk. Locusts of this species
travelling in a 5.8.W. direction have literally peppered the sky
to-day, but vagaries in the wind have caused many of them to
settle. (Morogoro, 11. xii. 16.)
The most striking thing about the insect-life to-day were the
locusts of two species (the second being Zonocerus elegans) which
were simply swarming everywhere; the herbage was literally alive
with the stragglers of the swarm of yellow adults that passed
over here three days ago. I met a native woman putting them
into a pail for dinner. On the middens outside the huts are
great piles of wings and hind-legs, the relics of recent feasting.
(Morogoro, 14. xii. 16.)
90. Zonocrrus ELEGANS Thunb, <A rather surprising situation
for immature grasshoppers of gaudy colour were the holes caused
by the larve of many species of longicorn in the thorn-trees,
whose timber was almost as hard to cut as stone. Yet I took many
of these young grasshoppers as high up as seven feet, and might
have found them still higher had I chopped further up the stems.
Immediately their hiding places were laid open they sprang into
the air. (Mbunyi, 17. v.16.)
This species is very interesting on account of the small
proportion that develop wings. The insects frequently mate
while their wings are quite rudimentary. ‘The development of
the wings is, I take it, a matter of no importance to the owner’s
safety, which is guaranteed toa great extent by its gaudy, apose-
matic colouring and offensive smell. (Morogoro, 14, xii. 16.)
These grasshoppers, apparently so warningly-coloured, are eaten
with avidity by hungry lizards (Gerrhosaurus major) and some-
what reluctantly by a Chameleon (C. dilepis dilepis)*. (Morogoro,
22. x. 17.)
This species comprised part of the stomach contents of a
Crested Cuckoo (Coccystes glandarius). (Morogoro, 29.1. 18.)
Also found in the stomach of another species of Cuckoo (Coc-
cystes jacobinus). (Morogoro, 29. xi. 18.)
91, PuHymareus vintprpxs Stal. Captured threelarge Phymateus
insects of the usual vivid green colour with scarlet, blue, and green
under-wings ; there were four of them on the bush, nevertheless
I did not notice them till the first moved. A curious mixture of
protective and aposematic colouring, they are rather slow in their
movements and only take short flights, relying for safety on the
frothy bubbles discharged from glands sitnated just behind the
large jumping-legs. (Ngari Mtoni, near Arusha, 4. iv. 16.)
Took a pair of these Acridians at Morogoro, where they are far
from common. (Morogoro, 10.i.17.)
92. Cysroc@LiIaA ABSIDATA Karsch. Just by my tent someone
caught a very queer orthopteron. Its length over all is 4? inches ;
* Compare Dr. Carpenter's experiments on monkeys with this Acridian (Trans,
Ent. Soc. 1921, pp. 8, 33, 53, 98).
EAST AFRICAN INSECTS. 1039
2% of this is occupied hy the abdomen. A casque of 22 inches
protects the head and well-developed wings. In breadth the
casque is 12 inches, in height 1? inches, and looks out of all pro-
portion to the rest of the insect.
The colour of the creature is a pale green with mid-ribs and
venations of the casque in yellow; the vein-like ribs are punctuated
ati intervals by yellow spots. Two peculiar silver blotches occur
about half-way along the casque, one on each side; these can be
seen from above. The upper wings have each a bright splash of
ultramarine near their junction with the body. (Morogoro,
14. v. 17.)
PHNASMIDA.,
93. PALoruus @rryi Grand., ¢. A native brought me a fine
stick-insect_ measuring 114 inches in length ; antenn 42 inches ;
hind-legs 43 inches ; opened wing 3% inches; elytra 2 inch long,
each having a hollow exerescence resembling a thorn. Large
stick-insects are rare in Kast Africa; I have only come across
two during three years, and these were of different species.
(Morogoro, 1. ii. 18.)
MANTIDS.
94. Prayina Mantips non det. An egg-mass of a mantis,
which I collected a month ago has just hatched out and given
rise to a few score grotesque little black mantids. (Morogoro,
20. iii. 17.)
In the stomach of a Hornbill (Lophoceros deckeni) were two
large and complete mantids. (Morogoro, 1. vi. 17.)
A. Hornbill (Lophoceros nawmannz) which was mobbed by some
small birds had a large mantis in its stomach. (Morogoro,
Usa We)
94a. DacryLorreryx sp. not in Brit. Mus. Took a mantis
which gave me a great deal of trouble tocapture. It harmonized
most perfectly with the bark of the tree on which I found it,
and van round the tree and up and down with the agility of a
gecko. (Morogoro, 2. ii. 17.)
95. PonysriLora #RuGINOsA Goeze. Whilst at my meals all
sort of trifles such as moth or fly wings come floating down. This
is a sign that a praying mantis is also dining. He took up
residence near the ventilating holes at the apex of my bell-
tent, and is between the two covers. Apparently he finds it a
good spot, for he has been there a very long time. (Morogoro,
15. vi..17.)
One of the Black-headed Weavers (Ploceus nigriceps) surprised
me to-day by dodging after a mantis in the fashion of a flycatcher.
Almost immediately after catching it, however, it dropped it again,
presumably owing to the mantis making good use of its toothed
fore-legs and perhaps jaws. (Morogoro, 7. xii. 17.)
1040 Mr. A. LOVERIDGE : NOLES ON
A specimen of this green mantis was brought me at 6 p.m.,
having been found on a tree-trunk. It deposited an egg-mass,
14 inches in length and 1 inch in diameter, on the window frame
before 8a.m. It stayed beside the eggs till dusk the following
evening, when it disappeared. (Nairobi, 7. viii. 19.)
The boy broueht me two more specimens of this mantis—one
‘immature but almost fully-grown and an adult. The latter
measured in head and body 68 mm., abdomen only 45 mm. In
its abdominal cavity, and quite free, I found a Gordian Worm (Gror-
dius sp., Baylis det.) 378 mm. in length and 15 mm. in diameter.
It would be interesting to calculate what length of worm a human
could carry in proportionate body-weights if he were afflicted
with such pests. (Nairobi, 12. viii. 19.)
96. SpHopromantris virtpis Forsk. our or five days ago
I brought a gravid mantis into the house; it spent most of its
time resting on a vase of flowers near the musquito-gauzed
window. This morning a male mantis was on the gauze six inches
from the female. I therefore went outside and brought him
in at 9a.m. The female was clinging to the gauze head upwards,
about 9 inches below the male, who was head downwards. At
12.17 p.m. the female took two short steps in the direction of
the male. At 12.19 p.m, with a rapid jump he sprang upon her,
his head being towards her tail; he turned so quickly that his
fore-legs grasped her just anterior to the middle pair of legs and
base of wings. His second pair of legs rested on her wings, also
his left posterior leg, the right posterior leg being in. space.
The terminal segment of his abdomen was twisted round in a
quite impossible way to form the union. They remained paired
thus from 12.19 p.m. to sundown at 6 p.m, (29.iv. 21). They
paired again for a couple of hours at noon, but after this made
no further attempts (30.iv. 21). Eggs were laid on the window
gauze (24.v.21). Eggs hatched ; young sloughed shortly after
hatehing. They are greyish-fawn in colour, with legs striped
like those of a Stegomyia fasciata, but not in such contrasting
black and white. (Kiloga, 6. vii. 21.)
97. OMomaNtIS zeBRATA Charp. A mantis, being put upon
a mosquito-meshed window where there were three bees, carefully
stalked them, swaying backwards and forwards with every step
just like a chameleon. It is curious to note that two creatures
so entirely different except in their green colour should have
developed precisely the same gait, which to my mind is meant
to simulate the movements of a leaf swaying gently to and fro in
a, breeze, and enables them to approach within, striking distance of
their prey—the one with its tongue, the other with its specially
modified pro-legs. ‘The mantis seized the bee with these; the
abdomen of the bee was held as in a vice in the left, its head in
the right ; without loss of time the mantis bit into and crunched
up the vital thorax, whilst the bee was quite powerless to
retaliate. (Iilosa, 27. i11, 22.)
EAST AFRICAN INSECIS. 1041
98. Manvris reniaiosA L., 9. The wings of this species were
swarming with minute red mites. (Kilosa, 29. iii. 22.)
99. PsEUDOCREOBOTRA WAULBERGI Stal. Arve mantids attracted
to light like moths, or is it because of the moths they can capture
that “they come? I took two examples of this species at a light on
the verandah last night. The eye-spots are seen to best advantage
when the mantis raises its wings and rustles them in its ewan
“warning ” attitude. (Frere flown. 2. vil. 19.)
100. Ipotum biABoLIcuM Sauss. Took an immature specimen
of a chestnut-brown. The leaf-like outgrowth from the head
reminded one of the Leaf-nosed Bats; other extravagant leaf-
like outgrowths occurred from the limb-joints and on the thorax.
(Dutumi, 15,ix. 16.)
I consider the species uncommon in Kast Africa, but as it is so
procryptically coloured my conclusions may be based on my own
poor observation. Ihave only captured two specimens in six
years, the second was taken to-day. (IMilosa, 18. xii. 20.)
ON ''HE GUERNSEY CROCIDURA. 1043
50. On the Guernsey Crocidura.
By Ivor G. S. Mowvacun, I'.Z.8., and Grace Pickrorp.
[Received Octoher 11, 1923: Read November 6, 1923.]
4
Among the material collected by G. E. Pickford in the Channel
Islands in August and September 1923, is a series of twelve
shrews from Guernsey. The examination of this series, and of
four Guernsey shrews already in the British Museum collection,
reveals a number of features which make it necessary to distin-
euish this race from the typical form Crocidura russula russula.
‘The few Guernsey specimens examined by G.8. Miller *, though
recorded as smaller than normal specimens of the large typical
form of the Continent, were insufficient in number to make clear
the characteristic nature of their dimensions. The uniformity
of the small dimension throughout the now extended series leaves,
however, no doubt of the distinct nature of the Guernsey race.
The following tabulation of the diagnostic characters of the
Kuropean races of C. russula indicates their affinities.
C. r. RussuLA Hermann.
Distribution. Central Europe, from Holland and Central Ger-
many to the valley of the Garonne and the coast of South-
Hastern Wrance. Ltaly.
Diagnosis. Size rather large.
(a) Hind foot 11:7-14.
(6) Condylobasal length of skull 19-20-4.
(c) Average of 29 adults: HB. 78°3; T. 38°7: HF. 12:8.
Colour usually dark.
C. x. PuLcHRA Cabrera.
Distribution. Central and Southern Spain. Lowlands of
France south of the Gironde.
Diagnosis. Smaller than C. ». russula.
(a) Hind foot 10:8-13.
(b) Condylobasal length of skull 18-19°4.
(c) Average of 24 adults: HB. 72°8; T. 37-1; HF. 11:9.
Usually paler in colour than Cr. russula.
C. xn. cinrra Miller.
Distribution. Cintra, Portugal.
Diagnosis. Size as in CL r. pulchra.
(a) Hind foot 11°4-12°7.
(6) Condylobasal length of skull 18-19-2.
(c) Average of 10 adults: HB. 67°6; 'T. 37:7; HF. 11:9.
Hairs with a peculiar strong coppery lustre.
* G.S. Miller, Catalogue of the Mammals of Western Europe, B.M., 1912.
1044 ON ‘TE GUERNSEY CROCIDURA,
C. R. PEvA, subsp. n.
?
Distribution. Guernsey, Alderney (?).
Diagnosis. Size as in C. 7. pulchra.
(a) Hind foot 11—-12°5.
(6) Condylobasal length of skull 18:1-19:2.
(c) Average of 16 adults: LB 68a. sare nih Tes:
Colour nearly as in C. 2. russula.
CRrocIDURA RUSSULA PEDA, subsp. n.
Crocidura russula russula Miller, Cat. Mamm. W. E., B.M.,
ge yas ON.
Type locality. Guernsey.
Material examined. Four adults (3 3) collected by Bunting in
August 1908 and twelve adults (7.3, 5@) collected by Pickford
in August and September 1923.
Dimensions :—
Type 8.9.2.20, adult 3,23 Aug. 1908. 1B. 63:5. P.39. ILI. 12.
I. 9.
Average of eleven adult males (8.9.2.18, ?8.9.2.19, 8.9.2.20,
8.9.2.21, 1, 3, 4, 5,5, 6, 9, 12) :-—
13118}, G8bi eA weS)s 1aUM, Iikey s 19 ee.
Average of five adult females (2, 11, 23, 24, 25) :—
1BUBY, (i A SM Sito e TEU IIMs Os tehlbe
Condylobasal lengths of skulls:—(8.9.2.18) 19°2; (8.9.2.2)
19:0; (11) 19:0; (8.9.2.20) Type 18°8; (1) 18°8; (8.9.2.19) 18:2;
3) Ise). é
ee he dimensions of this animal show throughout a
uniform smallness which at once distinguishes them from those
of the typical large form C. 7. russula. The normal coloration
is not to be differentiated from that of the dark typical form.
‘he few light specimens, those collected in 1908, are probably
only characteristic of the seasonal conditions in the presence of
which a similar lightness oceurs in GC. 7. russula. This occurrence
of a light summer phase should not be confused with the
lightness of C. +. pulchra which, as Miller (. ¢.) noticed, normally
occurs at all times of the year.
A. Crocidura from Alderney in the British Museum collection
should probably be grouped with the Guernsey form, but is too
young to admit of positive identification.
Noru—The dimensions throughout this paper are in iilli-
metres.
.
REVERSIBLE AND IRREVERSIBLE EVOLUTION, 1045.
D1, Reversible and Irreversible Evolution ; a Study based
on Reptiles. By Dr. Franors, Baron Norvcsa.
[Received October 10, 1923: Read November 20, 1923. ]
(Text-figures 8 & 9.)
Lhe great amount of information that we have about the
evolution of some groups of reptiles, the great amount of varia-
bility that these animals show, the long span of time that the
history of their evolution covers, seem to make it advisable to
base an investigation of the laws of evolution on the history of
these groups. Fejérvdry was one of the first who worked on
these lines (6).
As is well known, in the skeletal structure of Reptilia many
characters oceur that show what has been called the irreversi-
bility of evolution. Such characters are: the development of a
secondary armour in Dermochelys (4), the changes in the pelvis
of the orthopodous Dinosaurs (5), the secondary growth of the
plastron in the Cinosternide (9), and the development of a new
element (prepubis) functioning as pubis in the Crocodilia.
Apart from these changes, some of which have been well studied,
one can detect other less well-known changes tending to prove
that sometimes a reversal of evolution can take place. Changes
of this sort are: the secondary elongation of the anterior limbs
in Dinosaurs, the development of the postorbital bar in Mammals
and theromorphous reptiles, the redevelopment of more or less
plate-like ventral pelvic elements in different reptiles, the occur-
rence of polygonal flat carpal and tarsal bones in highly specialised
reptiles, and the relationship of the frontal to the orbit in
different groups.
The aim of this paper is to give a description of the different
changes of this second type and to draw conclusions.
§(1) The elongation of the anterior limbs in specialised Dino-
SMUrs.
In all primitive diaptosaurian reptiles, such as Rhyncho-
saurians and Parasuchians, and in a Jess marked degree in the
true Rhynchocephalians, the anterior limbs are generally only a
little shorter than the posterior. These animals are exclusively
quadrupedal. In the short-necked Ornithosuchians and in the
long-necked Proterosaurians, which were partly bipedal, a
marked shortening of the anterior limbs can be detected. ‘This
shortening is stronger in the originally bipedal Dinosaurs. 11
is very noticeable in the lightly-built triassie carnivorous Dino-
saurs (Lodokeosaurus, Hallopus, Procompsognathus) but less
marked in the jurassic and cretaceous representatives of this
group. In Procompsognathus (and Podokeosaurus?) the ratio
Proc. Zoou. Soc.—1923, No. LX VIII. 68
1046 BARON FRANCIS NOPCSA ON REVERSIBLE
of the anterior limb to the posterior is 10/27, in the jurassic
Compsognathus 10/18, in the nearly contemporaneous Ornith-
lestes 10/15, and in the cretaceous Struthiomimus 10/16. In
this group, in which not the jaws or the posterior limbs but
the anterior limbs were used for seizing the prey, a decided
lengthening of the anterior limbs occurs. In the heavily-
built carnivorous Dinosaurs, in which, much as in the birds
of prey, a prehensile foot is developed, this elongation does not
occur and the anterior limbs remain small or almost vanish (10).
A relatively short anterior limb is also met with in all bipedal
orthopodous Dinosaurs. The ratio is 10/21 in Hypsilophodon,
10/23 in Thescelesaurus, 10/19 to 10/17 in Camptosaurus, 10/14
in Zywanodon, 10/17 in Kritosawrus, and 10/15 in Corythosawrus,
hough less clearly than the preceding one, this list also shows
that in the more specialised forms, as Jguanodon and Corytho-
saurus, the anterior limbs are a trifle longer than in the more
primitive forms.
In the quadrupedal Sauropoda the anterior limbs are mostly
shorter than the posterior; in one group, however, the Brachio-
sauridze, the length of the limbs is nearly equal. In this case
the secondary elongation is very marked (10).
All these data show that in those specialised Dinosaurs in
which the anterior limb is continuously used a secondary
lengthening of this part occurs. ‘This can be considered as a
reversal to the ancestral pro-dinosaurian type. Diagrammatically
these changes can be expressed in the following manner :—
Anterior limbs long and functioning. Anterior limbs short.
Ancestors of Dinosaurs———-_
Primitive Dinosaurs.
Specialised Dinosaurs <-———— TERT
For one reason the case is not quite conclusive, for it can be
surinised that the apparent secondary elongation is not due to a
renewed growth but simply to the fact that in Dinosaurs the
posterior limbs, on account of their being more used, grew big
first, and that the growth of the anterior ones set in later.
Since also such a hypothesis might explain the temporary dis-
proportion of the limbs, the case must be considered doubtful.
§ (2) The development of the postorbital bar.
The second case to be dealt with is more typical than the
first. While in all more or less primitive reptiles the pestorbital
bar is complete, it is open in some rather specialised forms.
Such Reptiles are: many snake-shaped reptiles, some other
lacertilians, and the Bauriamorpha. Contrary to what is known
in Reptiles, in Mammals the postorbital bar 1s incomplete in the
primitive forms and complete only in the younger Equide, most
of the Artiodactyla, and in the Primates. This being the case,
it may be concluded that all Mammals descended fiom animals
Jacking a postorbital bar. ;
AND IRREVERSIBLE EVOLUTION. 1047
In spite of many mammalian characters, such as the structure
of the teeth, the articulation of the lower jaw, and the shape of
the brain, not the Cynognathidx but the Bauriamorpha must be
consider ea as the ancestors of the Maminalia, for the ribs of the
former show a non-mammalian trend of evolution.’ Curiously
enough the Bauriamorpha have no postorbital bar. ‘Thus the
disappearance of this part in the Bauriamorpha and its reappear-
ance in the higher mammals again points towards a reversal.
Wortmann’s discoveries of a separate postfrontal and even of a
postorbital bar in some Insectivora (19) show that this part of
the mammals is not analogous but homologous with the same
part in reptiles.
‘This change seems again to be nothing else than the reten-
tion of an embryonic character in the adult, for frequently
in embryos of animals characters appear that are later reduced.
Good examples are afforded by the temporary development of a
third cervical rib in the Lacertilia (8) and by the development
of a fourth and fifth digit in embryos of birds (18).
A. process similar to that which accounts for the development
of the postorbital bar in higher mammals is evidently also
changing the development of the claws in Opisthocomus, for
this bird is evidently forgetting how to fly and learning how to
climb (10).
For the history of the development of the postorbital bar in
Reptiles and Mammals the following diagram can be dyvawn :—
Postorbital bar complete. Postorbital bar incomplete.
Primitive Theriodonts----.--— --~.- -—--»Baurinmorpha (or similar
Theriodonts).
Specialised Mammalia<-————————_—_——Pr itive Mammalia.
§ (3) Zhe development of the ventral elements of the pelvis.
As is well known, in primitive Stegocephalians, for example
the Branchiosauride, the ventral elements of the pelvis consist
of four, or sometimes even only of two, small disk-shaped centres
of ossification that were evidently embedded in a large plate of
cartilage. Much the same type of pelvis is found in the recent
Urodeles. In the more specialised Stegocephalians (Hryops,
Cacops) the two ventral elements form a continuous mass of
plate-like bone with a small foramen perforating each pubis. It
is evident that this type of pelvis originated in the complete
ossification of the whole cartilage of the more primitive forms.
This solid type occurs also in the Cotylosaurians (Seymouria,
Diadectes, Labidosaurus (text-fig. 8 (1)), Pareiasaurus); in the
most agile Cotylosaurians (Procolophon), however, and iu the
Pelycosaurians (Ophiacodon, Varanosaurus) a central perforation
and separation of the pubis and ischium appear. From this
latter type were evolved the pelves of the higher reptiles, that
show either one great perforation in the centre and two small
foramina obturatoria passing through the pubes, or one large
68*
1048 BARON FRANCIS NOPCSA ON REVERSIBLE
foramen on each side between each pubis and ischium (text-
fig. 8 (2)). Through this foramen the obturator nerve passes.
As these openings grow larger the central pelvic elements are
more or less reduced to rod-like bones. This change is analogous
to the one that occurs in the skulls of different groups of reptiles,
for also in these the originally plate-like skull bones are reduced,
where they do not cover the brain-case, to rods that correspond
to the different lines of stress and strain.
The tendency to develop more or less rod-like ventral pelvic
elements is fairly well indicated in most tortoises (text-fig. 8 (3)),
with the exception of the marine ones, for the median ossification
is less marked in the modern tortoises than in the Amphichelyde,
The same structure is also observable in the primitive Sauroptery-
gians (Veusticosaurus (text-fig. 8 (5)), Anarosaurus).
Among the Parapsida the rod-like felvis is still missing in
Areoscelis but clearly indicated in Plewrosawrus and well developed
in all Squamata. Among the Rhynchocephalians large ventral
pelvic openings are absent in Howesia and the Rhynchosaurains,
but they are well developed in all other Rhynechocephalians
(text-fig, 8 (7)) with exception of the Proganosauria. In all
‘Thecodontia, all Dinosaurs, and all Crocodiles the pelvic apertures
are always large.
In contrast with this more or less plate-like ventral pelvic
elements are to be found in the specialised Sauropteryeians
(text-fig. 8 (6)), in the Proganosauria (text-fig. 8 (8)), and in the
Pterosaurians. Among the latter this feature is very noteworthy,
for it is’ especially well developed in the Pteranodontide, which
are the most specialised members of the Order (text-fig. 8 (9)).
Dermochelys, which is derived from some unknown chelonian
tortoise, has much smaller foramina obturatoria than all the
Chelonide, and retains in the pelvis a great amount of cartilage
throughout life (text-fig. 8 (4)). In this respect the pelvis of an
adult Dermochelys vecalls somewhat the pelvis of //atteria in an
early stage of development (11, 14). ‘The resemblance which
Baur (1) detected between the pelvis of some Testudinata and
the pelvis of the Rhynchocephalia is, of course, only due to a case
of convergence, for the situation of the foramen obturatorium is
different in the two groups.
Comparing now the relationships of the reptiles mentioned in
the above lines, it becomes clear that in three cases plate-shaped
pelvic elements must have arisen from rod-shaped bones. ‘Lhe
Plesiosaurians must have arisen from Nothosaurian reptiles, the
Proganosauria from jurassic Rhynchocephalians, and the Pteran-
odontide® from triassic Thecodontia. ‘hus these three cases are
quite characteristic cases of reversal. An indication of the same
sort of reversal is afforded by the differences that separate
Dermochelys from the Chelonidie. These differences show more
clearly than the return of the postorbital bar in what manner
such a reversal begins. As suggested in the former case, it
starts by the retention of an embryonic stage throughout life.
AND IRREVERSIBLE EVOLU'ION. 1049
Supposing that in Dermochelys the whole of the cartilage
were to turn to bone, very soon a pelvis would evolve that would
recall the most primitive reptilian pelvis. It would be more
primitive than the pelvis of the Amphichelydx. That such a
reversal can actually take place will be proved in detail in the
following paragraph, here it is enough to emphasise that in
Pteranodon such an ossification actually did occur.
On account of the complexity of the changes in the sania)
elements of the pelvis of reptiles the diagram also becomes
complex.
Ventral part of pelvis to Rod-like bones
great extent cartilaginous, Bony plates with smali with large
Joramen obturatorium small. Soramen obturatorium. apertures.
Peaniierstecocephalra —>f Specialised Stegocephalia; ee
primitive reptiles
wSssaks ¢ Specialised
—@ reptiles.
Primitive marine (and (less, esi aye
volant) reptiles
o ae niarine and
N
volant reptiles
§ (4) Carpus and tarsus of reptiles.
The changes that can be observed in the carpus and tarsus of
reptiles are ‘similar to those in the pelvis.
In primitive Stegocephalia, as in modern Urodeles, the carpus
and tarsus consist of flat polygonal pieces of car tilage with small
disk-shaped ossicles in their centre. In this respect it is sufti-
cient to refer to Uranocenirvodon and to Scincosawrus, the foot-
bones of which have been figured by Broom (3). In other
more reptile-like Stegocephalians carpus and tarsus consist no
longer of cartilage but of more or less flat polygonal bones with
but. little cartilage between thein (text-fig. 8 (10)) (Zrematops).
These tarsal and carpal bones evidently originated by the ossifica-
tion of the whole or of nearly the whole cartilage of the primitive
forms without much change in shape.
With the exception of the Procolophonide, the Cotylosauria
show much the same sort of foot-bones as the Stegocephalia.
In the primitive ones (Diadectes, Disparactus (text-fig. 8 (11),
Limnoscelis), evidently polygonal cartilage plates were present
with disk-like centres of ossification in their middle, while in the
more specialised ones (Pareiasauride) the cartilage is replaced
by polygonal bone. In the Procolophonide the structure of the
foot-bones is different. Instead of cartilaginous or osseous,
neatly immovable elements, ossicles with well-marked concave
and convex surfaces of articulation are present. Probably a fair
amount of cartilage was present, but probably also the surfaces
of the cartilage- bodies were curved.
In the Pelycosaurians the carpus is still sometimes polygonal
with a small amount of | flexibility (Ophiacodon), sometimes
rounded with a fair amount of cartilage (Varanops), and sometimes
1050 BARON FRANCIS NOPCSA ON REVERSIBLE
\ V
be
to
Text-figure 8.
WHA w'
SQV <f x : .
LW
.
® \
ee
NY
LAEY LIPIDS
fs \y
\
\ \\
he
v\ .
wien i \\ Eek VS
\ HR
Plate-like pelvis of carnivorous primitive Cotylosaurian Labidosaurus (from
Case).
. Plate-like pelvis of carnivorous highly organised Theriodont Cynognathus (from
Seeley).
» Rod-shaped pelvis of moderately specialised marine tortoise Chelone (from
Hofmann).
. Cartilaginous plate-like pelvis of highly specialised marine tortoise Dermochelys
(from Volker).
. Rod-shaped pelvis of semi-aquatic Sauropterygian Neusticosaurus (from Fraas).
. Plate-like pelvis of highly organised marine Sauropterygian Peloneustes (from
Linder).
. Rod-like pelvis of terrestrial Rhynchocephalian Sawranodon (from Lortet).
. Plate-like pelvis of aquatic Rhynchocephalian Champsosaurus (from b. Brown).
. Plate-like pelvis of highly specialised Pterosaurian Péeranodon (from Eaton).
. Polygonal tarsus of highly specialised Stegocephalian Zvrematops (from Willis-
ton).
. Disk-shaped cartilaginous tarsus of primitive Cotylosaurian Disparactus (from
Case).
. Disk-shaped carpus of primitive Ichthyosaurian Delphinosaurus (from
Merriam).
. Polygonal carpus of highly specialised Ichthyosaurian Ichthyosaurus (from
Huene, referred to there under the generic name Murypterygius).
> 5 Y. Yi
. Disk-shaped carpus of primitive Sauropterygian Proneusticosawrus (from Volz).
. Polygonal carpus of specialised Sauropterygian Polycoéylus (from Williston).
. Disk-shaped carpus of highly specialised marine tortoise Dermochelys (from
Volker).
i
, ‘
. Spherical carpus of primitive marine tortoise Tlowochelys (from Hay).
. Disk-shaped carpus of primitive Mosasaurian Tylosaurus (from Osborn).
19. Polygonal carpus of specialised Mosasaurian Platecarpus (from Williston).
AND IRREVERSIBLE EVOLUTION. 1051
well ossified with spherical surfaces of articulation (Dimetrodon).
The latter type is to be found also in all terrestrial Squamata
that have well-developed feet. Very little is known until now
about the foot-bones of the Theromorpha; they seem, however,
always to have attained a high degree of perfection.
Turning from the monozygocrotaphous Theromorpha to the
likewise monozygocrotaphous Sauropterygians, which evidently
descended from Pelycosaurians or Theromorpha, one is surprised
to remark that even in those Sauropterygians that are least
adapted to aquatic life (Weusticosaurus, Proneusticosaurus (text-
fig. 8 (14)) the foot-bones are flat and rounded ossicles that evi-
dently formed the centres of cartilaginous, polygonal plates. The
same sort of foot-bones are present in the liassic Plesiosaurians.
Ju the more specialised later Plesiosaurians, instead of the cartila-
ginous plates, polygonal flat bones are present. The flat polygonal
bones observable in Hlasmosaurus, Polycotylus (text-fig. 8 (15))
or Cimoliosaurus recall somewhat the flat polygonal bones of the
specialised Stegocephalians.
Similar changes as those in the Sauropterygians can also be
observed in the Ichthyopterygians. In Jfesosaurus and some
triassic Ichthyosaurians (Shastasaurus, Delphinosaurus (text-fig. 8
(12)) round bony disks occur that were evidently surrounded
by extensive cartilage. In all the more specialised {chthyo-
saurians (text-fig. 8 (13)) polygonal bony plates are present.
These are firmly applied against each other.
Somewhat similar changes as in these groups are to be met
with in the Testudinata. In the terrestrial tortoises polygonal
bones occur with a small amount of mobility between them. In
the Trionychidw these bones show by retaining at their angles
a good amount of cartilage a tendency to round off these angles.
In the Chelonide (text-fig. 8 (17), Toaochelys) this process is still
more marked ; finally, in Dermochelys (text-fig. 8 (16)) instead of
angular bones, polygonal plates of cartilage are present, with
flat bony disks in the centre. These changes show that the
rounding off of the primitive polygonal foot-bones of the Testu-
dinata is due to the retention of an embryonic stage throughout
life (14).
Applying this observation to the fossil marine forms hitherto
discussed, it becomes obvious that also in these first a cartila-
ginous embryonic stage, with small centres of ossification, had
become permanent for some time, and that after this transitory
stage extensive ossification set in, in much the same manner
as in the most primitive reptiles. In this instance it will be
remembered that the hypothesis of a secondary ossification has
already been brought forward in the foregoing paragraph. ye?
Among the marine Squamata the carpal and tarsal bones
retain spherical articulating surfaces in Opetiosaurus, they have
become to a great extent cartilaginous in T'ylosaurus (text-fig. 8
(18)), and are already to some extent replaced hy flat. polygonal
bones in Platecarpus (text-fig: 8 (19)). So also in this group the
same changes occur as in the groups already dealt with.
1052 BARON FRANCIS NOPCSA ON REVERSIBLE
Until the present investigation only the shape of each
isolated foot-bone was dealt with, now it becomes necessary to
consider the whole foot. While each separate foot-bone shows a
decided reversal of evolution, the whole foot as such shows some-
thing else. Although several of the carpal and tarsal bones can
he identified in all groups of reptiles, nevertheless the number
and the relative position of the foot-bones continually change.
This is why the foot of an Ichthyosaurian can readily be distin-
guished from the foot of a specialised Stegocephalian. In conse-
quence of the foot-bones always being differently arranged in the
different groups of reptiles, evolution seems to be irreversible.
Thus the foot-bones of reptiles show in a drastic manner how
in one point of an organ the evolution can be reversible, but
irreversible in another. When such a phenomenon oceurs in
correlated parts of the body, it is admissible to call the case a
mixed one.
The diagram representing the evolution of the foot-bones of
reptiles recalls the one of the evolution of the pelvis :—
Tarsus and carpus
polygonal cartilaginous Polygonal bony Bones with
plates. plates. q spherical swifaces.
Primitive Stegocephalians } eek { Specialised Stegocephalians }
» Cotylosaurians opp Cotylosaurs ( Higher
reptiles.
Primitive marine reptiles<—
a OuG fe sie
> Specialised marine “anes,
~ $(5) he devalaprient of the supraorbital region.
In nearly all the Stegocephalians the postfrontal and the pre-
frontal meet. above the “orbit and exclude the frontal from this
opening. It is only in some highly specialised forms that excep-
tions to this rule can be found. First of all the frontal touches
the orbit in those gigantic and, as Watson (15) proved, specialised
forms, such as Capitosaurus, Mastodonsaurus, and Cyclotosaurus ;
secondly, this occurs in those Labyrinthodonts that show a very
marked: broadening of the skull, such as Plagiosawrus (text-
fie, 9 (4)); thirdly, this occurs in the aberrant Microsaurian
Diplocaulus (text-fig. 9 (2)). In the less aberrant relatives of
Plagiosaurus and Diplocaulus as, for example, Batrachosuchus
(text-fig. 9 (3)), Diceratosaurus, and Batrachiderpetum (text-
fig. 9 (1)), the frontal is yet excluded from the orbit.
Other Stegocephalians, in which the frontal likewise borders
the orbits, are Gephyrosteqgus, which is characterised by the
thinning out of the cranial roof, suggesting the formation of
temporal vacuities, and Z'rematops and Br oiliellus, that both recall
the Cotylosauria. From all this it becomes evident that in the
Stegocephalia the entry of the frontal into the orbits is a sign of
specialisation, For the sake of convenience one can ¢all the 1 type
AND IRREVERSIBLE EVOLUTION, 1053
where it enters into the orbit the neo-orbital type and retain the
expression palco-orbital for the other.
Among the Cotylosauria the Diadectide (text-fig. 9 (5), Dia-
dectes), Paveiasauridee, and Limnoscelide show the same struc-
ture as the primitive Stegocephalians, the Captorhinimorpha and
the Procolophonide the other. Procolophon (text-fig. 9 (6)) is a
very agile Cotylosaurian, showing also many other signs of
specialisation: for example, a small lacrymal bone. In the
Captorhinimorpha, on the other hand, the limbs are specialised
to a rather high degree,
Among the Testudinata that are somewhat allied to the Cotylo-
saurians, the relation of the frontal to the orbit varies. In
some primitive Testudinata, such as Z'riassochelys, Chisternon,
and Kallokibotium, the paleo-orbital type is preserved: in some
other Amphichelydx already the neo-orbital type occurs.
Curiously enough, the palzo-orbital type occurs also in the Pro-
tostegine and the Dermochelyde (text-fig. 9 (8)), while the
Lytolomide and the Chelonide (text-fig. 9 (7), Toxochelys) show
the neo-orbital type. In primitive Chelonide (Zoxochelys) and
in the embryos of Chelone (14) the neo-orbital type is more
marked than in the adult Chelone. In the rest of the Tortoises
generally the neo-orbital type is met with; the palzo-orbital
type occurs, however, in the Platysternine and Chelydride, and
sporadically among the Emydide.
In Platysternum, Dermochelys, and some Chelydride, as
Macroclemmys, the paleo-orbital type is associated with a
secondary enlargement of the bones forming the roof of the
skull-and with the Joss of the power of hiding the head under
the shield. In accordance with this, in Dermochelys, Chelone,
and Chelydra, the posterior excavation of the parietal and the
squamosal is more marked in the embryos than in the adult (14).
Considering that in the most primitive Tortoises the capacity
of withdrawing the head had not yet been acquired, and that it
was but secondarily lest in Dermochelys, Chelone, Platysternum,
and dfacroclemmys, it becomes evident that this feature and the
paleo-orbital type are connected with each other. Evidently
the palso-orbital type of the more specialised Tortoises has been
developed from the neo-orbital type, for the ontogenetical changes
observable in the Chelonide point in this direction.
Among the Theromorpha the neo-orbital type dominates
Microgomphodon (text-fig. 9 (9)). The paleo-orbital type is only
met with in the Cynognathide (Protacmon, text-fig. 9 (10)).
In consequence of this it must be assumed that either the Cyno-
enathide retained a very ancient structure, or that also in this
case a reversal took place. Since in all Pelycosaurians that are
ancestral to the Theromorpha the neo-orbital type likewise occurs,
evidently the latter has to be assumed. As Pelycosaurians, it is
quite enough to mention the genera Varanosaurus, Sphenacodon,
Theropleura, and Dimetrodon.
Among the Placodontid, paleo-orbital genera as Placodus
1054 BARON FRANCIS NOPCSA ON REVERSIBLE
(text-fig. 9 (14)) and neo-orbital genera as Placochelys (text-
fig. 9 (15)) can be distinguished. Unfortunately nothing is known
about the evolution of this group, therefore no conclusions can
be drawn.
Text-figure 9.
\
i 0 a
\X NK
. Palxo-orbital skull of primitive Microsaurian Diplocaulids, Batrachiterpeton
(from Watson).
. Neo-orbital skull of specialised Microsaurian Diplocaulidee, Diplocaulus (from
Douthitt).
. Palxo-orbital skull of primitive Stereospondylous Brachyopidx, Batrachosuchus
(from Watson).
. Neo-orbital skull of specialised Stereospondylous Brachyopid, Plagiosawrus
(from Fraas).
. Palwo-orbital skull of primitive Cotylosaurian Diadectes (from Huene).
. Neo-orbital skull of specialised Cotylosaurian Procolophon (from Woodward).
. Neo-orbital skull of primitive marine tortoise Zovochelys (from Hay).
. Palwo-orbital skull of specialised marine tortoise Dermochelys (from Volker).
. Neo-orbital skull of primitive higher Theriodont Microgomphodon(from Watson).
. Paleo-orbital skull of specialised higher 'Lheriodont Protacmon (trom Watson).
. Palxo-orbital skull of Lepidosaurian Helodermu (from Phisalix).
. Neo-orbital skull of Lepidosaurian Platecarpus (from Williston).
. Tectorbital skull of Lepidosauvian Varanus (from Schmidt).
|. Paleo-orbital skull of Dranitesaurian Placodus (from Broili).
. Neo-orbital skull of Dranitesaurian Placochelys (from Jackel).
. Neo-orbital skull of primitive Archosaurian Huparkeria (from Broom).
. Tectorbital skull of specialised Archosaurian Campéosaurus (from B. Brown).
Among the Sauropterygians, the neo-orbital type is met with
in Anarosaurus, Pistosaurus, and Nothosawrus, the palzo-orbital
type in Cymatosawrus and all Plesiosaurians. In all the Plesio-
AND IRREVERSIBLE EVOLUTION. 1055
saurians the frontal shows a decided tendency to vanish alto-
gether, and, besides this, in the more specialised long-snouted
Plesiosaurians (the Piiosaurians) the tendency exists to develop
large supraorbital bones. This tendency is well observable in
the genera DPeloneustes, Brachyauchenias, and T'rinacromerwn.
In these genera the prefrontals and postfrontals are long and
narrow bones. On account of the reduction of the frontal these
animals revert at first to the paleo-orbital type, but when the
broadening of the head sets in they develop on other lines.
In the Parapsida, that include the Areoscelia, the Acrosauria,
and the Squamata, the frontal nearly always separates the pre-
frontal and the postfrontal. While it borders the orbit in Areo-
scelis, Pleurosaurus, all primitive Chameleons (18) and many
Lacertilians (Platecarpus, text-tig. 9 (12)), it is excluded in some
Lacertilians from the orbit by a supraorbital bone (text-fig. 9
(13), Varanus). Heloderma (text-fig. 9 (11)) and the specialised
Chameleons (17) differ from all the other Squamata in showing
the palo-orbital structure, but this may be due to a reversal.
Thus in this group the structure varies.
In the Diaptosaurians (text-fig. 9 (16), Huparkeria), the Dino-
saurs, and the Crocodiles, the prefrontals and the postfrontals
never meet. In some Crocodiles however, and in the ortho-
odous Dinosaurs supraorbital bones are developed (text-fig. 9
(17), Camptosaurus).
For the neo-orbital type, in which a supraorbital bone is
present, Fejérvary’s term, tectorbital (7), can be adopted. Since
the supraorbital bone is only developed in few groups of reptiles,
the tectorbital type is evidently new.
Proceeding now to group the primarily palso-orbital, the neo-
orbital, the secondarily paleo-orbital, and the tectorbital types
according to chronological order, it is soon seen that the primarily
paleo-orbital types are either permian reptiles or such that are
closely allied to permian reptiles. The neo-orbital type occurs
in different groups from the Permian upwards, it is most marked
in the most advanced reptiles ; the secondarily palzo-orbital forms
are found from the Trias upwards, but mostly among compara-
tively low postcretaceous reptiles; finally tectorbital types occur
exclusively from the Jurassic upwards.
The average conclusion to be drawn is that in primitive
reptiles (Cotylosauria, Tortoises) a reversal could easily occur from
the neo-orbital type to the paleo-orbital type; that, however, in
the more highly developed reptiles (Crocodiles, Dinosaurs) the
broadening of the skull could no longer be attained by a reversal
but only by the development of a new bony element. Compara-
tively primitive reptiles, as Sauropterygia and Squamata, seem
to be intermediate between the two extremes. ‘The primitive
nature of the Squamata is best shown by the circulatory and
respiratory organs.
In two most important papers Weidenreicli (16, 17) pointed
out that in some living animals characters occur that are very
1056 BARON FRANCIS NOPCSA ON REVERSIBLE
characteristic and date back at least to Pliocene time, but that
are all the same not yet perfectly fixed. Such characters have
each time to be acquired by a special stimulus in every individual.
When such a stimulus is lacking a reversal to the ancestral type
takes place. Such characters are, for example, among many
other ones, the blindness of Proteus (16) and the shape of the
calcaneus in man (17).
Other characters, as the development of the foramen of the
operculum, through which, in the Urodela, the extremities are
protruded (16), or the scrotum of man (16), are even then deve-
loped to a certain degree, when inciting stimuli, as the pressure
of the extremities against the operculum or the descent of the
testicles are not acting, but in such cases these characters ave
less marked than when the stimuli ave acting.
A third group of characters is always developed in ontogenesis,
and even apparently without reason. ‘These observations show
that in the fixing of new characters quite different stages occur.
Comparing now these stages with the changes found in the
orbital region, it is evident that the reversal of the neo-orbital
structure to the paleo-orbital type in permian or primitive
reptiles (Tortoises) is entirely analogous to the case when a not
yet fixed character is lost again. The undecided condition pre-
vailing among the Sauropterygians and the Squamata can be
well compared to the changes in Weidenreich’s second group,
and the development of the tectorbital type shows that in the
highly developed reptiles the neo-orbital type had become fixed
to such an extent that a reversal was no more possible.
In this way paleontological observations corroborate zoological
research, and the interest of this case lays in that it is correlated
with geological time.
The changes may be shown diagrammatically as follows :—
Palao-orbital type. . Neo-orbital types Tectorbital type,
Primitive Stegocephalians
49 reptiles Specialised Stegocephualians,
NCU: prejurassic reptiles
Specialised prejurassic ‘
de
reptiles Specialised post- Us lela { Specialised post-
jurassic reptiles § jurassic reptiles.
(The explanation of the abbreviated terms “ prejurassic ” and
“¢ postjurassic ” is given in the text.)
CoNnCLUSION.
After having discussed five cases of reversible evolution, four
of which are beyond question, and after having mentioned at the
beginning of the paper several cases of irreversible evolution,
conclusions may now be drawn.
The first certain case of reversible evolution shows how an
ossification, which was interrupted during the course of evolution,
AND IRREVERSIBLE EVOLUTION. 1057
sets in again. The second case shows the persistence of a
primitive stage of development in later more specialised forms,
and shows the subsequent development of another Stage of
evolution through which the ancestral forms had passed long
ago. ‘The third case, the mixed one, shows a similar change
coupled with “irreversible” evolution,and finally in the fourth case
three phases can be discerned—one phase, in which a character
is not yet fixed, so that a reversal is possible, a second undeter-
mined phase, and a third, in which a reversal is impossible. In
the latter case a particular function can only be attained by the
developmeut of a new organ.
Reviewing the “irreversible” cases, it can easily be detected
that the apparent “irreversibility” is always due to the fact, that
either an adjacent organ or an organ having a similar function
is called upon to replace a degenerating organ or that in the
absence of such parts a new organ is developed.
From the combination of these observations the following
statements can be deduced :—
(1) An apparent irreversibility will occur when a certain
character is already so strongly fixed that it cannot be altered.
Such a fixation will occur all the sooner if the dismissed organ
acquired a new function.
(2) An apparent irreversibility will oecur when some function
is nob perfectly concentrated in a special organ, so that similar! \
functioning organs are ready at hand to replace each other.
(3) An apparent irreversibility will occur when an adjacent
organ is ready to replace the more or less degenerated one. As an
example of this sort the pelvis of the Crocodiles may be men-
tioned, for in this case the posterior ventral ribs assumed the
function of the degenerated pubis and became the prepubes (new
hypothesis of the author),
(4) Evolution will appear irreversible when in some organ the
possibility of development still exists—z. ¢., if the organ is yet in
a primitive unspecialised state. In such a case this part is ready
to develop new features that can replace another degenerating
organ. An example of this case is afforded by the development
of the secondary dermal armour in Dermochelys. As Schmidt's
investigations (12) showed, the skin of the Tortoises ig not yet
strongly modified in the young, and so it has evidently not yet
lost the general faculty of developing dermal ossifications.
When one of the four enumerated ways of solving a bio-
logical problem has become impossible, an animal can only be
saved from extinction by a reversal to an embryonic stage.
This will only be possible—-
(1) if the state to be given up is not yet fixed by heredity.
That means if this state is not very far back in the history. A.
good example is afforded by the development of the supraorbital
region ;
(2) if the embryonic state to be called upon has not in the
meantime acquired a new and vital function. For this case
1058 BARON FRANCIS NOPCSA ON REVERSIBLE
Salamandra atra is a good example, for if the. gills of, this
Urodele were more adapted to the interuterine breathing than
they actually are, the rearing of its larvee in water would become
impossible (16). Even in Tortoises evidently the embryonic gills
ave already modified to such an extent that the Mud-Tortoises
were incapable of falling back on the use of their gills, and had in
the course of their aquatic adaptation to develop new pharyngeal
organs for breathing under water.
The unexplainable but important fact, that the life-history of
each individual is always a distorted recapitulation of the history
of its whole phylum, gives the clue by which we can understand
why a limited reversal of evolution can occur.
LIvERATURE REFERRED TO,
The purely osteological or paleontological literature must be
looked up for in Nopesa, ‘ Die Familien der Reptilien ? (Berlin,
1923).
1. Baur, G.—‘*The Pelvis of the Testudinata.” Journ. of
Morphology, 1892.
2, Broom, R.--“On the Structure of the Skull in Cynodont
Reptiles.” Proce. Zool. Soc. London, 1915.
3. Broom, R.— ‘On the Structure of the Reptilian Tarsus.”
Proc. Zool. Soc. London, 1921.
4, Dotto, L.—* Sur Vorigine de la tortue luth.” Bull. Soc.
roy. médicales et naturelles, Bruxelles, 1901.
5. Dotto, L.—‘‘Les Dinosaurien adaptées, a la vie quadrupéde
secondaire.” Bull. Soc. belg. Géolog., Paléont. et Hydro-
log., Bruxelles, 1905.
6. Fesirvdny, G.—-'Quelques observations sur la loi de Dollo
et I’épistrephogenése.” Bull. Soc. Vaudoise des Se.
nat. Lausanne, 1921.
7, Fesirvdny, G.—‘ Phylogenetical and Morphological Notes
on primary and secondary dermal bones.” Archiv f.
Naturgesch. Berlin, 1922.
8. Ktane, K.—*‘ Variationen der Wirbelsiiule und des Extremi-
tatenplexus bei Lacerta.” Morpholog. Jahrbuch, 1913.
9, Norcsa.—‘‘ A case of Secondary Adaptation in a Tortoise.”
Ann. & Mag. Nat. Hist. London, 1922
10. Norcsa.—‘‘ On the Origin of Flight in Birds.” Proc. Zool.
Soe. London, 1923.
11. Scuaurnsuann.—‘ Weitere Beitriige zur Entwicklungs-
geschichte von Jatteria.” Archiv f. microscopische
Anatomie, 1900.
12. Scumipr, W. J.—‘‘ Ueber Schuppenrudimente und Haut-
sinneorgane bei Hmyda granosa.” Zoolog. Anzeiger,
1920.
13. Srrmer, H.—‘ Die ontogenetische und phylogenetische Ent-
wicklung dev Vogelfliigelskelettes.” Acta Zoologica,
Stockholm, 1922.
AND IRREVERSIBLE EVOLUTION. 1059
14, VoriKer, H.—‘ Uber Stamm-, Gliedmassen- und Hautskelett
von Dermochelys.” Zoologische Jahrbiicher, Jena, 1913.
15. Warson, D. M.8.—‘ The Structure, Evolution, and Origin
of the Amphibia.” Phil. Trans. Roy. Soc. London,
1915.
16. Wemeynreicu, '.—“ Das Evolutionsproblem.” Roux, Vor-
trige und Aufsaitze tiber Hntwicklungsmechanik,
Berlin, 1921.
17. Werpernrercn, F’.—-“‘ Die Typen und Artenlehre der Verer-
bungswissenschaft.” Paleontolog. Zeitschrift, Berlin,
1923.
18. Werner, F.—“‘ Zur Kenntnis des Skelettes von Lthampho-
leon.” Arbeiten Zoolog. Institut Universitit, Wien,
1902.
19. Worrmann, J. L.—‘‘ On some hitherto unrecognised reptilian
characters in Insectivora.” Proc. U.S. Nat. Mus.
Washington, 1920.
NEW AND RARE REPTILES FROM SOUTH AMERICA. 1061
52. On New and Rare Reptiles from South America. By
Joan B. Procter, F.Z.8., F.L.8., Curator of Reptiles
to the Society, formerly in charge of the Reptile
Collections in the British Museum (Natural History).
[Received November 12, 1923: Read November 20, 1923,]
(Text-figures 1 & 2.)
The British Museum has recently acquired two interesting
collections of Reptiles and Batrachians from South America,
The first was made by Mr. A. C. McDougall in Trinidad, on
the River Mamore, Bolivia, and presented in 1922. It
includes a very interesting new snake of the genus Oxybelis
which I have named boulengeri after Mr. G. A. Boulenger, who
has given me so much help and encouragement in my work
during the past seven years. The second is a large collection
from Marajo Island, at the mouth of the Amazon, This was
made by Herr W. Ehrhardt, and it includes many rarities and
a new species of Amphisbena which I have much pleasure in
naming mitchell: after Dr. P. Chalmers Mitchell.
Complete lists of the species collected are appended for their
geographical interest, but for the sake of brevity, notes on the
more common species are omitted.
I.—The McDougall Collection from Bolivia.
ECAUDATA.
BuFoNniIpDas.
]. BuFo GRANULOSUS Spix.
LACERTILIA.
TEILIDA.
2. PANTODACTYLUS SCHREIBERSII Wiegm.
IGuAaNIDa.
3. PoLYCHRUS MARMORATUS L.
OPHIDIA.
GLAUCONIIDA.
4, GLAUCONIA ALBIFRONS Wag].
Proc. Zoou. Soc.—1923, No. LXIX. 69
1062 MISS JOAN B. PROCTER ON NEW AND
ILYSIID 4.
5, Inysta scyraLe L.
CoLUuBRIN:.
6. Henicors potyLeris Gthr.
One half-grown specimen is beautifully marked; the white
spots on the dark brown ventrals form a paired series, and the
throat hag a transverse white band as well as the usnal spots.
The throat of this specimen is puffed out in a similar manner
to that of an angry Dispholidus or Spilotes.
7. Herperopryas CARINATUS Ith
8. Liopmis paxcrLoayrus Wied.
9, RADIN@A COBELLA L.
10. XENODON MERREMI Wag].
DIPSADOMORPHINS.
11. Lerroprra ANNuLATA L.
12. Oxyruorus Peronarius L.
13. O. Brrorguarus Gthr,
14, O. coronarus Schneid,
15. OXYBELIS BOULENGERI, sp. n.
Habit very slender; eye large.
Snout twice as long as eye, tapering, truncated at tip. Tostral
broader than deep, scarcely visible from above ; internasals
slightly shorter than preefrontals ; frontal long, elegantly shaped,
three times as long as broad, as long as the purictals, as broad as
the supraoculars; loreal small, elongate, three to four times as long
as deep ; preocular large, widely separated from the frontal. Hye
large, diameter three times its distance from lip, going five and
a half times into total head-length ; two or three postoculars
(differing on the two sides); temporals 14+2; six upper
Jabials, the fourth very large and surrounding the lower third
of the eye-rim; four lower labials in contact with the anterior
chin-shields, which are much shorter than the posterior pair.
Seales smooth, in 17 rows; ventrals deeply rounded, 102 in
number ; anal divided ; subcaudals in 190 pairs.
Body olive above, with two darker narrow lateral streaks on
the fifth scale-row of each side, fading into bright green on the
third and fourth and cream-colour on the first and outer
edge of ventrals. A broad bright dark green band occupying
the true ventral surface with a median pale green streak.
Tail olive-green above and below, with a cream lateral stripe
which fades into pale green and disappears on the posterior half.
Tlead brown-olive above, bright green on the sides; a dark
NARE REPLILES FROM SOUTH AMERICA. 1063
brown-olive streak from snout-tip, through eye, continuing down
neck as the lateral streak ; lips and chin pale yellow-green.
Type-specimen a half-grown male collected in Trinidad, River
Mamore, Bolivia.
Text-figure 1.
Oxybelis boulengeri, sp. n.
IIead: a. dorsal view ; 6. ventral view ; c. lateral view.
This charming snake is allied to O. argenteus Daud., which it
resembles in the proportions of the snout and in the fourth upper
labial alone entering the eye-rim. It differs in many characters,
however, which are best shown by tabulation.
O. argenters. | O. boulengeri, sp. n.
Bye: moderate, 7 times in total | largo, 5} times in total head-length.
head-length.
Frontal: 3 times as long as broad. _| 4 times as long as broad.
Anal: entire. divided.
Throat: with regular punctations. immaculate.
Coloration : upper surface light | upper surface olive; ventrals dark
brown ; ventrals cream, with 2 wide | bright green with pale median
green lateral and a slender median | streak,
streak.
)
ELAPIN#.
16. Exars spixit Wag.
One female measuring about 3 feet.
So far as I am aware this rare Hlaps has not previously been
recorded from Bolivia. When the British Museum Catalogue
was published the collection only contained three small specimens,
in which the eye was larger in proportion, measuring two-thirds
its distance from the mouth. In fully adult specimens the
diameter of the eye is only half its distance from the mouth.
AMBLYCEPHALIDA,
17. LrproGnarius CAtESBYI Sentz.
69*
1064 MISS JOAN B, PROCTER ON NEW AND
IL.—The Ehrhardt Collection from Marajo Island, at the
mouth of the Amazon.
BATRACHIA ECAUDATA.
CYSTIGNATHIDA.
1, PALUDICOLA SIGNIFERA Gir.
9, LerroDACrYLus PENTADACTYLUs L.
3. L. mysracrnus Burmeist.
4, la. vypHontus Daud.
BUFONIDS.
5. Buro aranurosus Gthr.
5 specimens ¢ Q.
One male has lost the right hand, and the nuptial excrescences
are enormously developed on the inner side of the wrist stump,
so much so that they form a protuberant mass which is obviously
as useful for holding the female as a normal hand.
Tynip «&.
G. Wyna vaunina Fitz.
7. H. venvunosa Laur.
8. H. rupra Daud.
9), PilyLLOMEDUSA HYPOCIIONDRIALIS Daud.
LACERTILIA.
GuCKONID™.
10. GonaropEs nUMERALIS Guich.
11. THEcADACTYLUS RAPICAUDUS Hontt.
12. HemrpactyLus MABOUIA Moreau.
I@uaNiID«.
13. ANOLIS orronit Cope.
14, Potycurus MARMoraAtUS L.
15. Opimryorssa suprrciirosa L.
16. Unoornrron Azureum L,
[ive specimens of this rare lizard, all brilliantly marked.
17. Ieuayxa tuBERCULATA Laur.
18. AMEIVA sURINAMENSIS Laur.
A magnificent series of specimens.
RARE REPTILES FROM SOUTH AMERICA. 1065
AMPHISBANID &.
19. CopmrAs FLAvEscENS Bonn.
One adult female of this extremely rare species. It agrees in
every particular with the specimens already in the British
Museum, and with the description in the Catalogue of Lizards,
vol. iti. p. 418.
20. AMPHISBENA ALBA L.
21. AMPHISBENA MITCHELLI, sp. 0.
Habit as in A. vermicularis.
Snout obtusely pointed, rather prominent. Rostral small, as
deep as broad, scarcely visible from above ; nasals large, forming
a suture behind the rostral, which is slightly longer than the one
between the prefrontals; frontal suture longer than the pre-
frontal suture; two pairs of small parietals. Eye distinct
through the ocular; no preocular ; postocular large, pentagonal ;
a single triangular anterior temporal ; three large and one small
upper labial on each side, third in contact with ocular and post-
ocular. Symphysial totvagonal followed by a large heptagonal
median chin-shield ; two large posterior chin-shields in contact
Text-figure 2.,
Amphisbena mitchelli, sp. 0.
Head: a. dorsal view; 6. ventral view ; c. lateral view.
d, Anal region: ventral view.
with second and third sublabials, separated from each other by
4 small shields. Body slender, composed of 213 annull; 14
segments above and 14 below the lateral grooves, which are well-
defined. Each segment above about twice as long as broad ;
median ventrals twice as broad as long; a vertebral groove.
Tail composed of 30 annuli. Anal 4 segments wide, the two
median enlarged ; 2 preanal pores.
Upper surface a soft brown, each segment light-edged. Snout
and lower surfaces écru.
Described from a single adult from Marajo Island, mouth of
the Amazon.
1066 ‘MISS JOAN B. PROCTER ON NEW AND
This very distinct species may at once be recognized by its
relatively long nasal suture, large anterior temporal, and single
pair of preanal pores. It does not’ seem to be very closely
related to any other species. ,
SCINGCID 2.
22. Mapua AuRAtA Schneid.
OPHIDIA.
TyYPHLOPIDSA.
93, TypuLops rericuLatus L.
CoLUBRIN &.
24, Henicops LEorarpinus Schleg.
The British Museum has only received one specimen of this
rare snake since 1869.
25. DryMopius BODDAER'T f, t. Sentz.
26. CoLuBER corals Boie.
27. Herveropryas oartnatros L., var. E (FLAVOLINEATUS Jan).
28. Leproruis Liocercus Wied.
29. Liormis pa@ciLoayrus Wied.
30. L. reaine L.
Five hgr. and yg.
‘These specimens are brown on the upper surfaces, salmon-pink
blotched with black beneath, and strikingly similar to Lthadinwa
cobella in general coloration.
DIPSADOMORPHIN &.
31. Hsmanropes cencuoa L.
32. OxyRHOPUS TRIGEMINUS D. & B.
Two males.
The markings in this species are widely variable; the middle
black band of each set of three may be much broader than the
outer two, and rounded like a hugh lozenge-shaped spot. The
paired white rings may be very narrow indeed, or equal to the
black ones in width.
33. PHILoDRYAS viRIDISsIMUS L.
34, OxyYBELIS FULGIDUS Daud.
35, O. acumtnatus Wied.
36. HoMALOCRANIUM MELANOCEPHALUM L.
RARE REPLILES FROM SOUTH AMERICA. 1067
MW UAPIN A,
37. Wnars spixit Wagl.
One 6d, similar to the specimen already noted in the McDougall
collection. Both these specimens have very much _ swollen
cheeks, and minute eyes half or less than half their distance
from the mouth in diameter.
38. Hnars MArcaraAvit Wied.
The white band across the prefrontals and 2nd and 8rd labials
is well marked; eleven sets of annuli on the body, the white
rings being 4 the width of the black ones.
NEW AND RAR REPLTLES AND BATRACHIANS, 1069
53. On New and Rare Reptiles and Batrachians from the
Australian Region. By Joan B. Procrsr, F.Z.S.,
F'.L.8., Curator of Reptiles to the Socicty.
[Received November 12, 1923: Read November 20, 1923.1
(Text-figures 3 & 4.)
Several interesting collections from the Australian Region
have recently been received by the British Museum (Nat. Hist.).
I shall include four in this one paper, and limit it for the sake of
brevity to notes on rarities and descriptions of new species.
I—NEW GUINEA.
Mr. W. Potter recently collected for the Museum in N.E. New
Guinea, and besides large series of the commoner things, obtained
several rare lizards and a new skink. ‘The latter I have named
after Mr. C. J. Battersby, who has been a most helpful assistant
to me throughout the time that I was in charge of the Museum’s
collections of Reptilia and Batrachia.
LACERTILIA.
GECKONIDS.
GYMNODACTYLIS LORIE Blgr.
A male (length: head 45 mm., body 110 mm., tail missing),
at least twice as large as the male type-specimen, the only
individual already in the Collection; in other particulars the
two specimens are identical.
GYMNODACTYLUS LOUISIADENSIS dle Vis.
A very Jarge well-preserved female (length: head 35 mm.,
body 93 mm., tail 47 mm.).. The three specimens of this rare
Gecko already in the Museu are from the D’Entrecasteaux Group _
and the Solomon Islands.
PyGoropDID”.
Lranis sicant Bier.
One adult of this extremely rare Pygopod was collected at
Lababia. Each dorsal scale, with the exception of the two
median series, has a small black apical dot, and is also finely
speckled. The species was only known from the three type-
specimens.
1070 MISS JOAN B. PROCTER ON NEW AND RARE REPLLLES
SciNciIDs.
Lygosoma (LIoLEPISMA) virENS Peters.
Four of these sharp-nosed skinks were collected on the banks
of the River Adler, Huon Gulf.
Lyaosoma (LiotErisMA) Lonaicers Bler.
One very well-marked adult: from Hae.
The digits are depressed with smooth lamelle below as far as
the end joint; the distal phalanx is slender and compressed,
and the digits therefore resemble those of a /emidactylus.
Lycgosoma (EMoA) BATYERSBYI, sp. n.
Snout moderate in length, obtusely pointed. Rostral small,
twice as broad as deep, forming a suture with the frontonasal,
which is broader than long and roundly pointed anteriorly ;
frontal three-fifths broad as long, as long as fronto- and inter-
parietals together; frontoparietals fused ; interparietal small,
distinct ; parietals forming a short median suture, and followed
by a pair of enlarged nuchals. Nostril pierced between a minute
nasal, small postnosal, and supranasul ; frontonasals separated
from each other; lower eyelid with a transparent disk ; sixth
upper labial in place of subocular. Kar-opening sinall, not as
Text-figure 3.
Head of Lygosoma (Hmoa) battersbyi, sp. n.
a. dorsal view; 6. ventral view; c. lateral view.
large as the transparent palpebral disk; one auricular lobule.
Digits long, distally compressed ; thirty smooth lamellw under
fourth finger, forty under fourth finger, forty under fourth toe.
Scales in thirty rows; sharply tri- or quinquecarinate. Mar-
ginal anals slightly enlarged.
Habit lacertiform ; the distance between the tip of the snout
and the fore-limb equals the distance between axilla and groin ;
the hind-limb reaches to in front of the shoulder when adpressed.
Tail nearly twice length of head and body.
Olive-green above, lighter on the head; opalescent beneath.
An indistinctly marked light lateral band, black-edged.
AND BATRACHIANS FROM THLE AUSYRALIAN REGION. 1071
A single adult speciinen from Huon Gulf, N.E. New Guinea.
Allied to ZL. tropidolepis Blgr.*, from which it differs in having
the interparietal shield distinct and the marginal przanals
enlarged.
II.— QUEENSLAND.
‘wo coltections ave considered in this chapter. One made by
Mr. 'U. V. Sherrin in Ravenshoe, N. Queensland, includes several
rare species until now unrepresented in the Museum’s collection.
The other, made by Capt. Wilkins in 8... and C. Queensland,
although small, also includes rare things.
ECAUDATA.
CysTIGNATHIDS.
LIMNODYNASTES FLELTCHERI Blgr.
Wilkins Collection. St. George’s District, C. Queensland.
One male specimen of this rare frog was obtained. The skin
is glandular and the markings clearly defined, spots in paired
lateral series beginning on the loreal region and continuing from
the temporal; a large shield-shaped occipital blotch and large
median dorsal and sacral markings. Basal web between the toes
well marked.
Originally described from New South Wales tf, but recently
recorded from 8. Queensland.
ENGYSTOMATIDS.
SPMENOPHRYNE VARIABILIS Blgr.
Sherrin Collection. Ravenshoe, N. Queensland.
One male specimen of this rare frog was collected. It differs
from typical specimens in having a slightly longer snout, but I
hesitate to give it a distinct varietal name on this account. In
all other respects it agrees with the type-series, and the charac-
teristic markings are well defined. This species was known only
from Celebes.
TLyLip&.
Hyxa Lesveurn D. & B.
Wilkins Collection. C. Queensland.
A single male of this extraordinary /Zyla was collected ; the
species is unlike all the rest in the genus, resembling Rana agilis
in general appearance.
(Epura tryont de Vis (ocELLATA Blgr.).
Sherrin Collection. Two specimens from Ravenshoe, N.
Queensland,
* Boulenger, Trans. Zool. Soc. xx. 1914, p. 260, pl. xxix. figs. 4, 4a.
+ Boulenger, Ann. Mae. N. H. (6) ii. 1888, p. 142.
1072 MISS JOAN B. PROCTER ON NEW AND RARE REPTILES
. VARANUS VARIUS Shaw.
‘Wilkins Collection. One specimen (skull and dried skin ; total
length 49”: head and body 19” and tail 30").
In this specimen the characteristic yellow spots are absent on
the anterior half of the body, which is merely speckled ; they are
well marked, however, on the posterior half.
VARANUS VARIUS var. BELLI Gray.
_ Wilkins Collection. Two specimens (skulls and dried skins) of
this beautiful banded variety. The Museum had only one
specimen, ‘The new pair are 50” (tail 30”) and 41” (body 20",
tail truncated) respectively. The four dorsal and three caudal
bands are arranged as follows :—
The first band is lunate, extending from the head backwards
to the shoulders ; the second is shaped like a bolero jacket ; the
third is very broad, with convex borders pointed medially ; the
fourth covers the sacral region and the upper part of the thighs.
There follow three caudal bands, the first 4” wide, the second 8",
the third 5” ; the tip of the tailis also dark. All the intermediate
areas are bright sand-coloured, black speckled.
ScrNCcIDs.
Lycosoma (Hinuxia) Quoyi D. & B.
Sherrin Collection. ‘hree well-marked specimens from Ravens-
hoe, N. Queensland.
Although this species has a wide range of distribution, it is
rare and local.
Lycosoma (TfinutA) reENuge Gray.
Wilkins Collection. One adult collected on Tambourine Mt.,
S.1M. Queensland. ‘This individual is Z. murrayi Blev. of the
British Museum Catalogue (iii. p. 232). The Museum has only
the specimen originally collected by H.M.S. ‘Challenger. On
comparing these two specimens of murrayi, which have 34 scale-
rows, with the series of tenwe the supposed differences in the size
of the ear-opening and in coloration cannot be detected, and
without counting the scale-rows it is impossible to tell them
apart. LZ, tenue is already known to have scale-rows in 28, 30,
or 32 vows, so that it seems impossible to maintain individuals
with 34 rows as a distinct species on this character alone.
L. murrayt Bigr., therefore, should be placed in the synonymy of
L. tenue Gray.
Lyaosoma (HinuL1a) monorropis Bler.
Wilkins Collection. One adult from C. Queensland.
Sandy brown, with ten dark transverse bands on neck and
body and twenty-three on the tail. Some of these bands bifurcate
on the sides.
AND BATRACHIJANS FROM THE AUSTRALIAN REGION. 1073
LyGosoma (LioLurisma) CHALLENGERI Bier.
‘his species is only known from the two type-specimens which
were received from the ‘Challenger’ Expedition in 1882.
Mr. Sherrin has collected two more in Ravenshoe, both agreeing
precisely with ours and with the description in the British
Museum Catalogue (vol. iii. p. 268).
Lycosoma (Itnopona) PuNCTATOVITTATUM Gthr.
Wilkins Collection. One specimen from Thomby Station,
St. George, Central 8. Queensland. Until last year we possessed
only the type-specimen of this species from Queensland. Prof.
Wood-Jones then sent us one collected on Flinders Island in the
Great Australian Bight. Zeitz records it from Queensland,
Victoria, and Tasmania, so that, although so rare, its range
probably extends all over the eastern half of the continent. The
present specimen agrees. in every particular with the type and
with the description in the British Museum Catalogue (iii. p. 335).
Lyaosoma (Homotnpma) ma6pprer Lonnb. & And.
Sherrin Collection. This species is new to the British Museum.
One adult was collected, minus half its tail but otherwise well
preserved. It agrees well with Lonnberg and Anderson’s
description *.
'TROPIDOPILORUS QUEENSLANDLE de Vis.
Sherrin Collection. his species is also a great rarity new to the
Museum’s collection,. An adult and one very young specimen were
obtained which agree well with the minutely detailed description
given by de Vist in everything except markings. They have
very pale alternating cross-bands, which give a. reticulated effect
dorsally and continue on the tail. Beneath, the pale tint pre-
dominates, with reticulations on the chin, down the sides and
beneath the tail. A median dark wavy band from the chest to the
vent. The two specimens are marked exactly alike.
OPHIDIA.
CoLUBRID®.
HornocermaALus BIrorquatus Jan.
Wilkins Collection. One young specimen from Thomby
Station, near St. George, Central S. Queensland. It has a well-
defined cream-coloured nuchal collar and the characteristic black
markings on the head. The Museum has not received a specimen
of this rare snake since 1876.
HLAPINA.
PSEUDECHIS AUSTRALIS Gray.
Wilkins Collection. One chestnut-brown adult from Thomby
Station, Central S. Queensland. Aberrant in having 19 scale-
rows on the body; 21 on the neck.
* Lénn. & And. Kungl. Sv. Vet.-Akad. Handl. lii. 1915, p. 6.
t Proc. Linn. Soc. N.S. Wales, 1889, vol. iv. p. 1034.
1074 MISS JOAN B, PROC'TER ON NEW AND RARE REPTILES
III._SOUTH AUSTRALIA.
Tn 1921 Prof. Wood-Jones made a very valuable collection of
lizards in Nuyt’s Archipelago and the Investigator Group.
These lizards, which he presented to the Museum, were mostly
rare skinks, amongst them a new species—Lygosomu (/lomolepida)
wood-jonesi. The description of the latter, with notes on that
collection, appear elsewhere *. ‘This year Prof. Wood-Jones has
sent us a supplementary collection mostly from the coast of the
mainland, including a new Amphibolurus and several rave geckos
and skinks. Only species of special interest in this second
collection will be noted here.
LACERTILIA.
GECKONIDA.
Herrnonora piort Gray.
Whilst working at Prof. Wood-Jones’s specimens I compared
our series of H. binoet and derbiana Gray, and came to the con-
clusion that they cannot be maintained as distinct species. The
arrangement of the dorsal tubercles is subject to great individual
variation, the derbiana form completely intergrading with that
proper to binoet.
DIPLODACTYLUS TESSELLATUS Gray.
There are but two specimens already in the Museum’s col-
lection. Prof, Wood-Jones obtained another on a journey from
Hergath Springs, N.E., to Kilalpaninna, Cooper’s Creek. Unfor-
tunately all three are males.
PEROPUS VARIEGATUS Dum. et Bibr.
Prof. Wood-Jones’s specimens would formerly have been called
Gehyra australis, but Fry has pointed out 7 that the genus should
be Peropws Wiegm., and series of specimens now show that
australis intergrades with variegatus.
AMPHIBOLURUS IMBRiICcA'tUS Peters.
Prof. Wood-Jones collected one female at Berri, River Murray.
Its scales are extremely lanceolate and the characteristic markings
are indistinct. ‘The Museum until now had only two specimens
of this very rare species—both females—so that in this case the
reverse of the state of things noted under Diplodactylus tessellatus
has occurred, and we have now three females of A. imbricatus
and no males, and three males of D. tessellatus and no females.
* Procter, “Flora and Fauna of Nuyt’s Archipelago and the Investigator Gro
No. 6—Lizards.” Trans. Roy. Soc. 8S. Australia, 1923. ban an eeniee
+ Fry, Rec. W. Aust. Mus. i. 1914, p. 178.
AND BATRACIIANS FROM 'THE AUSTRALIAN REGION. 1075
AMPILIBOLURUS FIONNI, Sp. n.
Habit slender, limbs very long.
Head moderate; snout as long as the orbit, with strongly-
marked canthus rostralis and concave loreal region; nostril
nearer the eye than tip of snout, situated below the canthus edge;
tympanum as large as eye-opening; upper head-scales tubercular,
keeled on the snout, smallest on the supraocular region. Sides
of neck stvongly plicate; a fold set with small spines from the
angle of the mouth to the shoulder, others below the eye and
above the tympanum, and one in the form of a low nuchal
Text-figure 4.
Sass
Head of Amphibolurus fionni, sp. n.
a. lateral view; b. dorsal view; c. ventral view.
erest, flanked by a pair reaching from the occiput to the shoulders,
Gular scales minute. Body covered with small keeled scales,
which are minute on the sides and enlarged on the vertebral
region. No enlarged ones scattered on the sides. Ventral
scales small, smooth, the ones under the fore-arm and under the
tibia keeled. Caudal scales small, strongly keeled.
Limbs very long, the fourth toe reaching to the nostril when
the hind-limb is adpressed, scales on the limbs slightly enlarged,
strongly keeled. Digits long, distinctly fringed. About 40
femoral pores, slightly interrupted in the middle. ‘Tail slender;
depressed at the base; twice length of head and body.
1076 MISS JOAN B. PROCTER ON NEW AND RARE REPTILES
Upper surfaces all olive-grey with darker variation. A lateral
series of large dark blotches. Greyish white beneath, the throat
reticulated with grey.
Described from a single female. Head and body 60 mm. in
length; tail 118 mm.
This species is allied to A. ornatus Gray, from W. Australia,
from which it may easily be distinguished as follows:—
A, ornatus Gray. ; A, fionni, sp. n.
Lateral scales heterogeneous. Lateral scales homogeneous.
No nuchal crest or spinose A low nuchal crest and
folds on head and neck several lateral spinose
(described from an adult folds on neck (although
male). described from a_ small
female).
Toes feebly fringed. Toes strongly fringed.
Since the type is a small femalo the spinose folds make a
distinguishing character, as we know that adult males in ornatus
do not have them, and it is to be expected that the male of
the new species would haye them still more developed. The
coloration and markings are also quite distinctive, but this
might be sexual.
Tt is also allied to A. rufescens Stirling, but the characters in
which it differs from this ave more numerous,
AMPILIBOLURUS ADELAIDENSIS Gray.
Prof. Wood-Jones caught three specimens of this beautifully-
marked species, which has not been received here for the past
fifty years.
‘YMPANOCRYPTIS CEPHALUS Gthr.
Zeitz in his Catalogue (p. 198) places this species in the
synonymy of 7’. lineatus Peters. The two species are, however,
quite distinct ; lineatus has sharply-keeled scales and spinose
tubercles ; cephalus has smooth, more cycloid scales and _ less
spinose tubercles. ‘I'he present collection included a female and
young,
Lycosoma (Lionepisma) EntREcAsrEauxit D, & B.
The adult collected by Prof. Wood-Jones is the first to be
added to the Museum’s collection since the publication of the
Catalogue in 1887. As far as I am aware, it has so far only
been recorded from Tasmania. The present specimen was caught
on Pearson’s Island, Investigator Group.
The new specimen has tricarinate dorsal scales, and is excep-
tionally well marked,
AND BATRACHIANS FROM THE AUSTRALIAN REGION. 1077
Lyeosoma (RiopoNna) BouGAINVILLI Gray.
One adult was collected at Port Lincoln.
LyGosoma (Ruopona) Brves Fischer.
This species is very rarely met with in South Australia.
ABLEPHARUS LINEO-OCELLATUS var. ADELAIDENSIS Peters.
One adult of this rare variety from Port Lincoln.
OPHIDIA.
KLAPIN«a,
DIEMENIA TEXTILIS D. & B.
Kreft states that young specimens from Adelaide lack the
typical dorsal bands met with in young from elsewhere. A
specimen in the present collection from Mt. Lofty bears this out,
but it is well worth noting that there is a specimen from
N. Queensland in the Museum’s collection which is precisely
similar in this respect.
Proc, Zoou. Soc,— 1923, No. LXX, i)
vem
Soha f
eh gee
-
eye
ADDITIONS TO THE SNAKE FAUNA OF EGYPT. 1079.
54. On Additions to the Snake Fauna of Heypt.
By Major 8. 8. Frowrr, O.B.E., I.L.8., F.Z.8.
[Received August 31, 1923 :. Read November 6, 1923.]
1. Summary.—Snakes of the genera Typhlops and Lycophi-
dium ave vecorded for the first time from Hgypt, remarks are
made on the status of Zycophidium abyssinicus, and the Protero-
glyph Snake Walterinnesia egyptia is for the first time definitely
proved to be an inhabitant of Hgypt.
2. Acknowledgements.—1 am indebted to Dr. Walter Francis
Innes Bey, formerly Curator of the Zoological Museum, School
of Medicine, Cairo, to Prof. Edward Hindle, F.L.S., F.Z.S.,
Biological Department, School of Medicine, Cairo, and to Mr.
Michael J. Nicoll, F.2Z.8., Assistant Director, Egyptian Govern-
ment Zoological Service, for kindly giving me many opportunities
of examining Heyptian snakes.
I wish also to express my thanks to Miss J. B. Procter, F.1L.S.,
I.Z.8., for her kindness in Jooking up, and allowing me to
examine, certain snakes in the collection under her charge in the
British Museum.
3. Typhlops.
When Mr. Boulenger published his “ List of the Snakes of
North Africa,” P.Z.S. 1919, p. 299 et seg., no representatives
of the family Typhlopidee were known from Egypt.
Among snakes lent to me for examination in April 1923 were
two individuals of the genus 7’yphlops.
lst. A spirit specimen definitely proves that Z’yphlops occurs
in Egypt. The discovery is due to Dr. Innes, who caught the
snake himself on 6th March, 1906, among the roots of a tamarisk
tree at Marg, on the eastern outskirts of Cairo.
The dimensions of this specimen in spirit, 17th April, 1923,
were :—
mm. | per cent. Remarks.
Length, snout to vent .........) 267 100 Scales in 22 or 24 rows.
Bee tr Ne ces anisad che odaponooG 4 149 Tail ends in a sharp point.
Por totale Men er ez 101°49
Diameter of body ...............| aah 187 | Goes 5t times in total length.
PY tail, ab vent ...... 5 | 187
(lengthy of shielded }
Head portion, m median 4. 1-49
THT® con noo cou ene oupotce
» totallength ............ 7(P) 2°62(?)
my. WACGIED, coo cco nso codisas casos 45 1:68
gy depth oo. c.. sc. cee een e= =o: 4 1:49
ee 70*
1080. . MAJOR S. 8S. FLOWER ON ADDITIONS
This individual appears to me referable to the Typhlops vernii-
cularis of Mr. Boulenger’s British Museum Catalogue of Snakes,
and so extends the range of that well-known species.
9nd. On the 15th of April, 1923, Prof. Hindle sent me a live
snake which he had purchased in Cairo from an Arab vendor of
reptiles. The Arab said the snake came from Damietta. ‘This
may or may not have been so. As Damietta is a seaport its
trade with other parts of the Mediterranean and the local habit
of careening sailing craft on the shore make it possible for a
snake found there to have been brought, by accident, with cargo
or ballast from almost anywhere. Seaports are not ‘ good
localities ” for recording reptiles from.
As far as I could examine this live individual it was also a
specimen of Z'yphlops vermicularis, but it was of most unusual
size for that species, being about 383 mm. in total length. The
diameter of body was 7 mm. (i. e., goes about 53 times in total
length).
A, Lycophidiwm.
In “A Hist of the Snakes of North-East Africa” by Mr.
Boulenger, P. Z.S. 1915, p. 646, two species of Lycophidiwm are
mentioned :—
Lycophidiwm capense from “Tropical and South Africa.”
Lycophidium abyssinicus from “ Abyssinia.”
In “A List of the Snakes of North Africa” by the same
author, P. Z.S. 1919, the genus Lycophidiwm is not mentioned.
5. Lycophidium abyssinicus Boulenger, Brit. Mus. Cat. Snakes,
i. p. 342, appears to be known only from the type-specimen in
the British Museum, and to differ from Z. capense only in having
the rostral shield narrowed, almost pointed, behind. On ex-
amining the type in London, July 17, 1923, it appears to be
inseparable from L. capense. Two females of L. capense from
the Blue Nile have their rostral shields pointed.
6. Lycophidium capense.
When Mr. Boulenger wrote the Catalogue of Snakes there
were no specimens of LZ. capense from North-Hast Africa in the
British Museum, and so the single individual Lycophidium from
Abyssinia was separated by a very wide tract of country from
the known range of L. capense.
In July 1923 in the British Museum there were four specimens
from North-East Africa entered in the catalogue in manuscript
by Mr. Boulenger, under :—
Lycophidium capense, var. B.
Female. V.188. C. 31. Roseires, Blue Nile. S.S. Flower. 1909.
Female. V.191. C. 40. Deesa, 5 “ i
Male. V. 166. C. 35. Onaramalka, Abyssinia. G. A. F. Abercromby. 1916.
Female. V. 180. C, 33. i Fs i
These four specimens have ad/ subcaudal shields paired,
TO THE SNAKE FAUNA OF EGYPT. 1081
7. An Egyptian Lycophidium.
Dy. Innes told me that he had in spirit a supposed young
individual of Walterinnesia egyptia, collected in agricultural land
in the Fayim, Upper Egypt, in 1904, by the Rev. Father
Teillard, and that there was no doubt whatsoever as to the
locality. In answer to my enquiries Dr. Innes told me that
Father Teillard had never collected in the Sudan, but had, from
Cairo, gone to stay at a farm in the Faytim, and while there
came across this snake and brought it back with him for
Dr. Innes’s collection.
On 19th April, 1923, Dr. Innes lent me this specimen for iden-
tification. For fear of damaging it, I did not examine the teeth,
but externally there appears no reason not to consider it a
Lycophidiam capense.
Scales 17 rows. Ventrals circa 196. Subcaudals 39: these
subcaudals consist of three pairs at base, then three undivided
scales, then thirty-two pairs.
It was on the strength of these undivided subcaudals that the
snake had been referred to Walterinnesia.
It is a most remarkable fact that this South and Tropical
African snake should occur in the Fayim, but a parallel case
appears to exist in the Rough-keeled, or Egg-eating, Snake
Dasypeltis scabra, of which the late Dr. John Anderson recorded
a single specimen from the Fayum.
8. Walterinnesta egyptia Lataste, 1887.
Dr. Innes discovered this species: he saw a snake in the hands
of a native ‘“snake-charmer” in Cairo, and recognizing that it
was something different from the ordinary Cobras of the genus
Naia, purchased the specimen and sent it to France to Monsieur
Fernand Lataste.
On 7th February, 1887, this snake was described as a new genus
and species by M. Lataste (‘ Le Naturaliste,’ Paris, 1887, p. 411
et seq.), the generic name being in honour of the discoverer.
9. “Habitat” of Walterinnesia.
As the type-specimen was purchased by Dr. Innes from a pro-
fessional ‘‘ snake-charmer ” in Cairo, it was in itself no evidence
of being of Egyptian origin. A trade in live reptiles between
the resident and wandering performers in Asia and North Africa
exists, and probably has existed for many centuries.
Subsequently Dr. Innes purchased two more individuals from
the same man. So three specimens of Walterinnesiw appear to
have been known when Dr. John Anderson wrote on the
herpetology of Egypt, but Dr. Anderson does not mention the
number. ;
Dr. Anderson wrote (‘ Zoology of Egypt,’ Reptilia, 1898,
p- 325): “The only specimens on record of this species were
purchased by Dr. Walter Innes from a snake-charmer in Cairo,
1082 MAJOR §. S. FLOWER ON ADDITIONS
and there is nothing to fall back upon, beyond the statement of
the juggler, to establish it (Walterinnesia) as an Meyptian species.
I have made the most careful enquiries about its presence in the
neighbourhood of Cairo, without having been able to throw any
light upon the subject, and Dr. Tnnes’s endeavours to procure
more specimens have been fruitless.”
_ Since Dr. Andergon’s time several other visitors to Hgypt,
interested in Zoology, have made special endeavours to obtain
specimens of JValtéerinnesia, but without success.
On 12th November, 1908, the man who had sold the first speci-
mens to Dr. Innes brought to Giza a live Walterinnesia, which
IT purchased for the Giza Zoological Gardens. This man said
that this species of snake was “ies found in Keypt, and could
only be obtained near the river Atbara, in the Berber Province
of the Sudan. ‘This statement was probably. made to enhance
the value of the specimen he offered to sell, but it influenced me
at the time. I informed Mr. Boulenger, anu in G. Z. G. “ List
of Animals (2nd edition),” 1910, p. 328, wrote of this species
“ Habitat probably Upper Nubia.” S80 Mr. Boulenger, 2.4.5.
1915, p. Gd6, Bye the distribution of Walterinnesia cequpbid as
“Nubia 2 Heypt 2? 2” and, P.Z.S. 1919, p. 306 :—‘ Heypt ?
Nubia?”
Meanwhile, in 1904, Dr. Innes had received from the Fayim
a black snake supposed to be an immature MWaléecrinnesia. In
April 1923 I had an opportunity of seeing this specimen, and, as
uientioned above, it belongs to a very different genus.
10. Rediscovery of Walterinnesia.
On Sth April, 1923, Mr. M. J. Nicoll shot a black snake about
21 miles east of Cairo, on the Cairo-Suez road, which he handed
over to me the same day for examination. It is the fifth in-
dividual of Walterinnesia egyptia of which we have record, and of
great importance as being the first specimen obtained with
reliable data as to locality, ete. It proves that M. Lataste was
not in error when he gave the specific name egyptia. Mr.
Nicoll’s find also brings to notice the interesting fact that
Walterinnesia is found in the desert, far from water, in the same
kind of country where the Vipers of the genera Ce rastes and
Echis occur.
The only Proteroglyph Snakes, besides Walterinnesia, which
ue known from Keypt ave two species of Cobra, Naia haie and
Nuia nigricollis, neither of which, as far as my pr esont experience
goes, are ever found more than about half a mile’s distance from
permanent water.
11. Known specimens of Walterinnesia.
(i.) The type. Female. Described by Lataste. Now in
British Museum.
(ii.) Male. Described by Boulenger (Cat. Snakes, iii. p. 392)
and by Anderson (1898, pp. 324. 825). Now in British
Museum.
TO THLE SNAKE FAUNA OF EGYPT. 1083
(iii.) Obtained by Dr. Innes (as were Nos. i. and ii.). Was in
the Cairo School of Medicine Museum, now in the Giza
Zoological Museum.
(iv.) Obtained alive (v. sup.) November 12, 1908, lived in Giza
Zoological Gardens for eight months nineteen days.
Male. Now in the British Museum.
(v.) Male. Collected by Mr. M. J. Nicoll, April 5, 1923, on
desert, about 21 miles east of Cairo. Now in Giza
Zoological Museuin.
12. Notes on 5th specimen of Walterinnesia egyptia,
by 8.8. F., April 5, 1923.
Male. Collected to-day by Mr. Nicoll.
General impression :—A heavily built, short-tailed snake, with
a large flattish head, a small black eye, and a conspicuously
large nostril.
Colour :—Upper surfaces shiny black; lower surfaces grey.
Ventrals 190. Anals 2. Subcaudals 51 (actually 51 right,
50 left. First, at base, divided. Second to sixth entire. Seventh
divided. Highth entire. Remaining forty-two divided, and one
extra half on right side).
| nn. per cent. | Remarks.
Length, snout to vent ......... | 943 100 |
ayo ACHE colemoddonuguacseacellt sale 16°9 Tail ends in a sharp point.
. OGRE a eee coucns | 986 1169
Diameter (greatest), body ...! ¢. 23 e.2°72
length of shielded Head-shields agree with de-
Hest) portion, in metinn 24 2°84. } scription by Mr. Boulenger,
Tine rece aaneseteests Cat. Snakes, iii. p. 392.
» totallength ............ 32 3°79
py WAG EMG corer eee ees aesaeon ce 23 2°72
Be GED brwastccwms case Wascel 13 154
Number of scales round body :—
At about 5th ventral circa 29
a 2 Obi Sunes 21
mee OAN A Tos ps 21
ai. SOth eel mn heen Res
WOE GLY leek, 23
Pn OO tents. Dis
i Wuaaloothy (nes 21
avrmenilin(O thivea: 4% i
By eul OME Ya at a 17
ny
ue
CERVICAL VERTEBR® OF A GIGANTIC BLUE WHALE. 1085
55. Cervical Vertebree of a Gigantic Blue Whale from
Panama. By Sir Sipney F. Harmer, K.B.E., Se.D.,
A ree ey
[Received November 19, 1923 ; Read November 20, 1923. j
~ (Text-figure 1.)
The bones under consideration (which were exhibited) were
the second and third cervical vertebrae of a Blue Whale (Bale-
noptera musculus V.), and had been presented to the British
Museum (Natural History) by Mr. F. A. Mitchell-Hedges, who
had given the following account of their history :——The whale
entered the harbour at Cristobal, the northern entrance to the
Panama Canal, in January 1922. It passed up the canal towards
the first locks at Gatun, and having become a menace to shipping,
it was killed with machine-guns. It was towed by tugs to the
Cristobal docks, where unsuccessful efforts to raise it from the
water were made with powerful 75-ton cranes. It is said to
have been carefully measured, at this stage, as having a length
of 98 feet, and its weight was estimated at 100 tons. It was
Jater towed out to sea, but it drifted ashore again, and after
having been towed out once more, it was bombed from the air
by United States army planes. Parts of the carcass subse-
quently came ashore at Santa Isabel, between Nombre de Dios
and Cape San Blas, where the vertebrae were found by Mr.
Mitchell-Hedges.
The specimen is of special interest from several points of
view. The Blue Whale is commonly considered an ice-loving
species, and it has been found in large numbers on the fringe of
the Arctic and Antarctic ice. Although it is known to travel
considerable distances from the poles, as shown by its frequent
capture by whaling companies off the Finmark, Newfoundland,
and South African coasts, records of its occurrence in or near the
Tropics are rare. In his memoir, “The Whalebone Whales
of New England” *, Mr. Glover M. Allen states vhat the Blue
Whale is essentially a ‘‘ cold-water” species, and that New
Jersey “ perhaps represents nearly the normal] southward limit ”
on the Atlantic coast, though it may eventually be found to.
follow the inshore waters as far south as the Carolinacoast. The_
specimen under consideration occurred in lat. 9° 30'N., and the
record is of importance as bearing on the possibility of a migration
across the equator of Blue Whales from the Northern Hemi-_
sphere to the Southern, or vice versd.
“# Allen, G. M., Mem. Boston Soc. Nat. Hist. vol. viii, No. 2, p. 265 (1916).
1086 SIR SIDNLY WARMER ON CERVICAL VERTEBRA
In structure the vertebre show an unusual peculiarity.
Although Balenoptera is ordinarily distinguished from Balena
by the fact that all the cervical vertelze are completely free,
these two vertebre are firmly ankylosed to one another by a
union between their centra, principally on the right side, at the
periphery, where there is some indication of a diseased condition
of the bone.
The specimen is further of interest, from the information it
gives as to the size which may be reached by the Blue Whale in
Northern waters. Recent unpublished records of the Southern
Whaling Companies establish the fact that the length of 100
feet may be exceeded in that part of the world, and the opinion
has been expressed that the Southern race differs from the
Northern by attaining a greater size. ‘Thus Allen (op. cit. p. 249),
relying on measurements of Northern Blue Whales made by
Text-figure 1.
te}
Blue Whale (Balenoptera musculus). Anterior view of the ankylosed axis and
third cervical vertebra; the neural arch of the latter just visikle through the
neural canal of the axis. Panama, I. A. Mitchell-Hedges Coll.
A seale nearly of 5 feet in length is seen resting on the somewhat injured
neural spine of the axis.
True and at the Norwegian whaling. stations, comes to the con-
clusion that 77 feet 2 inches ‘is probably nearly a true maximum,”
although he mentions a Norwegian record of 87 feet 6-5 inches.
The question had previously been discussed at length by True *,
who concluded (p. 156) that the maximum length of Newfound-
land Blue Whales ig less than that of Norwegian specimens,
pointing to a difference in size of Blue Whales on opposite sides
of the Atlantic. This opinion, which has been quoted in support
of the view that the Eastern and Western schools are distinct in
that ocean, is obviously affected by the present record.
* True, I. W., “ Whalebone Whales of the Western North Atlantic,” Smithson.
Contr. Knowledge, vol. xxxiii. (1904).
OF A GIGANTIC BLUE WIHTALE MROM PANAMA. 1087
The material available for comparison in the British Museum
is unfortunately scanty, and the best is a skeleton of a Blue
Whale which was stranded at Rosslare, Wexford Bay, on Mar. 25,
1891. According to the information given by W. Crouch* and
G. KH. WH. Barvett-Hamilton t, this specimen was probably a
female 82 feet long. The examination of its vertebral column
shows that tho opiphyses of the centra ave free, or nearly free,
Mong the whole length of the column. The process may
perhaps have commenced in the caudal region. It should be noted,
however, that True quotes an observation by Guldberg, who
measured a male Blue Whale as 78 feet 9 inches long, and subse-
quently ascertained that all the epiphyses were ankylosed to the
bodies of the vertebre. Flower § has pointed out that ankylosis
of the vertebral epiphyses in Cetacea commences in the cervical
and caudal regions, extending from both ends towards the middle
of the length of the column, where the process is finally completed.
He distinguishes animals in which this process has commenced
but is incomplete as being in the “adolescent ” stage, which has
thus haraly been reached by the Wexford whale. The Panama
vertebra, on the contrary, have their epiphyses completely
united; and it may be inferred from their condition that the
animal was fully adult, although it is obvious that they give no
. complete answer to the, question whether all the vertebral
epiphyses were thus united.
The axis of the Panama specimen measures 4 feet 74 inches
from tip to tip of the transverse processes, and it thus greatly
exceeds in size those of (1) the Wexford whale, with a corre-
sponding measurement of 3 feet 81 inches, and (2) the female
specimen recorded by Sir Willian Turner ||, of an estimated
leagth of “70 to 80 feet or upwards,” stranded at Longniddry,
Firth of Forth, November 1869, whose axis measured 3 feet
8 inches across. The Panama axis is strikingly more massive
than that of the Wexford whale, as is shown by a comparison of
the weights. The ankylosed axis and third cervical vertebra of the
Panama whale weigh 112 lbs., while the corresponding bones of
the Wexford whale are only 53 lbs. This gives a fair comparison
of the actual volume of the bones, even taking into account the
possibility of a slight error due to differences in the amount of
animal matter left after cleaning. It is further in accordance
with expectation, in an aniinal increasing from 80 to 100 feet in
length, a proportion of 4 to 5. The increase in volume should
be in proportion to the cubes of these numbers, and the cube of
5 1s almost exactly twice the cube of 4.
- It is of interest to attempt to verify the recorded length of the
Panama whale. True has recorded the measurements of a
* <The Zoologist,’ 1891, p. 216.
+ Tbid. p. 306. i
t Op. cit. p..161. ; :
§ Proc. Zool. Sov. 1864, p. 385.
|| ‘Marine Mammals Anatom. Mus. Uniy. Edinburgh,’ London, 1912, pp. 40-60.
“| Op. cit. pp. 180, 184.
1088 SIR SIDNEY HARMER ON CERVICAL VERTEBR&
Blue Whale stranded at Ocean City, New Jersey, in October
1891. Although not completely satisfactory, in view of the
small size of the animal, this may be taken as a basis for com-
parison. The measurements it is necessary to notice are as
follows :—-
Whale, Skull, Mandible, Mandible, Axis
length. length. along curve. straight. width.
New Jersey... 66’ 2" 14’ 74" Ne? 312 15’ 2" 3 gr
(Catimated )
Wexford ...... 82’ 0” 4 200 ieee TS) Shean O and
Longniddry ... @ = Oe OU Da ite Sr. Se
Panama 44:.. — = == —= AT}
The Wexford skull is partially disarticulated, and it is not
possible to state its exact length. According to True’s measure-
ments of the New Jersey specimen, the total length of the
animal is approximately given by multiplying the length of the
skull by 4:5, of the mandible (along curve) by 3°9, of the mandible
(straight) by 4-4, and of theaxis-breadth by 22. The proportions
in other whales are known to differ as a result of age or individual
variation, but it is not without interest to ascertain how far the
factors indicated above will apply in the present connection.
Taking these proportions, the estimates of the length of the
Longniddry whale, derived from the axis, the mandible (straight)
and the mandible (along curve) are respectively 80' 8”, 85' 5",
and 82' 6", with an average of 82’ 10’, which agrees well enough
with ‘Turner’ s vague estimate of ‘*70 to 80 fect or upwards. 55
The same three measurements of the Wexford whale are respec-
tively 81’ 7", 82' 1", and 78’ 4”, with an average of 80’ 8”, as
compared with the recorded length of 82 feet. The only available
measurement of the Panama whale is that of the axis; but,
applying the same proportion, the estimated total length of the
animal is 101’ 33”. I think this may be regarded as a substantial
confirmation of the recorded length of 98 feet. he skull of
this animal may have been about 23 feet long.
Scoresby *, a particularly reliable authority, mentions the
following records of large Blue Whales, which he describes as
Balenoptera Gibbar :--A specimen found dead in Davis Strait,
105 feet; an individual stranded on the banks of the Humber
in September 1750, 101 feet. Another old record of a large
Blue Whale is that of the well-known ‘‘ Ostende Whale,” which
was found floating in the North Sea and towed into the harbour
of Ostende on Nov. 4, 1827, its length having been variously
estimated as 80 to 102 feet. It has been customary to discredit
such measurements, but the evidence of the Panama specimen
tends to confirm theiraccuracy. The point is one of great impor-
tance, in view of the inclination of naturalists to distinguish a
* © An Account of the Arctic Regions,’ vol. i. pp. 481, 482 (1820).
OF A GIGANTIC BLUE WHALE FROM PANAMA. 1089
Southern race of Blue Whales by their superior size. The
evidence now submitted shows that Blue Whales of a size much
larger than that generally accepted may occur to the north of the
equator, and thus throws doubt on the assumption that the
Northern Blue Whale is smaller than those that frequent
the Southern Ocean. It may be suggested that owing to the
great intensity of whaling which has occurred in Northern waters,
few Blue Whales of the largest size have been permitted to
survive, but that in the South, where whaling has been practised
only since 1904/5, a greater number of these individuals have
remained. It may be anticipated that the enormous destruction
which is at present taking place in the South will result in a
diminution of the number of these very large specimens,
NOTES ON THE AFRICAN CRESTED RAT. 1091
56, Notes on the African Crested Rat (Lophiomys imhaust).
By G. H. Gonprincu.*
[Received July 9, 1923: Read November 6, 1923.]
Native (Wandorobo) name “ Monget” or “ Krmonget.”
The first one of these animals I got was when I was stationed
at Nakuru; it came from the Aberdare side. It was taken out of a
hole in a tree by a Wandorobo whom I had sent to collect a hyrax.
I found that the Wandorobo had the superstition about this
animal that if anybody got bitten byit he died. I think this may
be taken to mean that somebody got bitten and got blood poisoning
and died, as their bite is harmless. However, after this old man
found he was unhurt, I had no difficulty in getting all J wanted,
and at one time I had something like a dozen of them. I forget
whether I had one born in captivity, but J remember one
delightful toto. With regard to food, I have come to the
conclusion that besides eating leaves etc. they probably also
want, or rather eat, insects, small mammals, young birds, ete. I
used to feed mine on sweet potato leaves mainly, but found
they would also eat lucerne, wild clover, ete., and would eat the
potatoes themselves, but did not seem to care much about
them. On this diet I have had them live six months, as far as
Leanremember. Why I think they are also carnivorous is this :
I kept them in a sort of aviary with a row of boxes for
them to go into, and I had got hold of a winged Guinea pigeon
and thought it would be 2 good place for him, but in the morning
all that remained were a few feathers. They are, I think, purely
nocturnal animals, but run about much more on the ground
than tree-hyrax. ‘They are most cunning in escaping from
cages. I have met mine, when they got out, trotting along
the road to Nakuru in the middle of the night. I gave two
or three to the late C. W. Woodhouse to try and get home to the
Zoo, but they died on the voyage. I daresay what troubles them
is the heat, as they come from the cold forests. I think they are
pretty hardy really, and there seems to be no bother about
getting them to feed when they are freshly caught. Mine
used to come out of their boxes just as it began to get dark.
The best method of handling them is by the tail. There would,
I think, be no difficulty in getting them if they were wanted.
This, by the way, is not the animal that gnaws old tusks of ivory,
as Sir F, Jackson suggested. What does that is the porcupine.
* Communicated by the SECRETARY.
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THE SECRETARY ON ADDITIONS 'lO THE MENAGERIE. 1093
EXHIBITIONS AND NOTICES.
October 23rd, 1923.
Dr. A. Smrrn Woopwarp, F.R.S., Vice-President,
in the Chair.
The Srcrerarny read the following Report on the Additions
made to the Society's Menagerie during the months of June,
July, August, and September, 1923 :—
JUNE.
The registered additions to the Society's Menagerie during
the month of June were 210 in number. Of these 91 were
acquired by presentation, 13 were deposited, 69 were purchased,
and 37 were born in the Menagerie.
The following may be specially mentioned :—
1 Chimpanzee (Anthropopithecus troglodytes), 2, from West
Africa, purchased on June 12th.
1 Black Mangabey (Cercocebus aterrimus), 3 , from the Congo,
purchased on June 29th.
2 Pumas (Felis concolor), ¢ Q, from South America, pur-
chased on June 22nd.
1 Goliath Heron (Ardea goliath), from Kerna Marshes, Meso-
potamia, presented by Major Wilson.
2 Kurdistan Chukar Partridges (Alectoris grwca hurdistanicus),
from Kurdistan, new to the Collection, presented by Dr. Sinder-
son on June 8th.
1 Syrian Ostrich (Struthio camelus syriacus), from Jauf, Central
Arabia, new to the Collection, presented by Sir Percy Cox,
G.C.M.G., and Capt. R. EB. Cheesman, I.A., on June 8th.
2 Sundevall’s Lizards (Lygosoma swndevalli), new to the Collec-
tion, from Dodoma, K. Africa, presented by Arthur Loveridge,
C.M.Z.8.
JULY.
The registered additions to the Society’s Menagerie during the
inonth of July were 316 in number. Of these 148 were acquired
by presentation, 146 were deposited, 9 were purchased, 8 were
received in exchange, and 5 were born in the Menagerie.
The following may be specially mentioned :—
A collection of animals from Australia, containing Opossums,
Bandicoots, Regent-Birds, Bower-Birds, Brush-Turkeys, and
others, presented by the Marquess of ‘Tavistock, F.Z.8., on
July 3rd. :
Proc. Zuon. Soc, —19238, No. LX XI, fal
109£ THE SECRETARY. ON ADDITIONS ''O, THE MENAGERIE.
A collection of animals from Gambia, containing an Ostrich,
a Porcupine, an African Tantalus, Owls, and others, presented
by H.E. Capt. C. i, Armitage, C.M.G., D.8.0., P.ZS., on
July 24th.
1 Llama (Zama glam), bred in the Menagerie on July 7th.
A. collection of Indian Snakes, including an Indian Sand-
Snake (Zamenis arenarius) and a THodgson’s Snake (Coluber
hodgsonz), both new to the Collection, presented by A.A. L. Flynn,
Hsq., C.M.Z.8., on July 26th.
_ Avoust.
‘Nhe registered additions to the Society’s Menagerie during the
month of August were 205 in number. Of these 58 were
eon by pr esentation, 78 were deposited, 5 were purchased,
2 were received in oxchange, and 62 were born in the Menagerie.
The following may be peel, mentioned :—
1 South African Giraffe (Giraffa camelopardalis capensis), pur-
chased on August 2|st.
2 Ocelots (Felis pardalis), 1 Hensel’s Cat (Felis pardinoides),
2 Orab-eating Dogs (Canis thous), and 2 Amazonian Skunks
(Mephitis amazonicus), the last new to the Collection, from
Brazil, presented by George Chalmers, C.M.Z.S., on August 9th.
2 Gelada Baboons (Zheropithecus gelada), from Abyssinia,
received in exchange on August 30th.
1 Black Howler (ih ycetes caraya), from the Amazons, pr esented
by Walter Goodfellow, F.Z.8., on August 21st.
1 Puma (felis concolor), from South America, presented by
‘Arthur R. T. Woods on August 15th.
1 Fossa (Cryptoprocta ferox), from Madagascar, presented by
Percivale Helyar on August 11th.
6 Lion Cubs (felis leo), born in the Menagerie, 2 on August Ist
and 4 on August 25th,
SEPTEMBER.
The registered additions to the Society’s Menagerie during the
month of September were 221 in number. Of these 135 were
acquired by presentation, 60. were deposited, 18 were purchased,
5 were reeeived in exchange, and 3 were born in the Menagerie.
The following may be specially mentioned :—
1 African Elephant (Loxodon africanus), from ‘Tanganyika
Territory, presented by H.E. Sir Horace Byatt, K.C.M.G.,
Honorary Member of the Society, on September 14th.
2 Great Anteaters (J/yrmecophaga jubata), from South America,
purchased on September 15th.
1 Yellow-throated Marten (Martes flaviqula), from India, pre-
sented by the Bombay Natural History Society on September 12th.
LION-CUBS BORN IN THE GARDENS. 1095
Dr. C. W. Anprews, F.R.S., F.Z.S., exhibited, and made
some remarks upon, an imperfect phalangeal bone belonging to
one of the curious clawed Perissodactyls, the ‘Ancylopoda, (Chalz-
cotheroidea). Although only a scrap of bone, it is quite charac-
teristic, and there is no doubt about the determination. The
interest of the specimen lies in the fact that this is the first
recorded occurrence of the group in Africa: members of it are
known from Europe, Asia, and America in beds of ages ranging
from the Kocene up to the late Pliocene or early Pleistocene,
the Pleistocene forms being confined to Asia. The beds from
which the present specimen comes are of late Pliocene or, more
probably, Pleistocene age, and are situated on the Bunyoro side
of Lake Albert: remains of Hippopotamus phacochwrus, Crocodile,
and fish were also found. It is interesting to note that a species
of Chalicotherium is found in the Pliocene beds of Samos asso-
ciated with Samotherium, a close relative of: the existing Okapi,
and it may be suggested that possibly a still surviving Chali-
cothere may be the basis of the persistent rumours of the
existence in Central Africa of a large Bear or _Hyzna-like
animal. The specimens were collected and sent to the British
Museum by Dr. Wayland, Director of the Geological Survey of
Uganda.
Mr. D. Seru-Smiru, Curator of Mammals and Birds, exhibited
n series of lantern-slides of Lion-cubs born in the’ Society’s ©
Gardens. He said that since 1887 no lion-cubs born in the
jardens had lived more than a very short time. At the present
time there were two families of cubs, one of two and the other of
four, the former being over three months and the latter about
nine weeks old. Both families had been born and were being
reared in the outside cages of the Lion House, and at present
appeared to be thriving.
Mr. Seru-Smirm also exhibited a living specimen of the
Mangabey described in 1899 by Dr. Sclater as Cercocebus eongicus
(P.Z.8. 1889, p. 827), and pointed out that it was without
doubt a semi-albino variety of Cercocebus aterrimus.
Dr. Cant Axsoton exhibited a large series of drawings and
photographs of Cave Animals from the Balkans.
Mr. E. A. Spaut, B.Sce., F.Z.S., gave an account of his experi-
ments on acceleration of metamorphoses of Frog-tadpoles by
injection of anterior-lobe pituitary-gland extract and iodine,
1096 LIVING SPECIMEN OF A PIGMY HIPPOPOTAMURS,
November 6th, 1923.
Sir S.F. Hanmer, K.B.E., F.R.S., Vice-President,
in the Chair. a sast,
The Secretary exhibited, and made remarks upon, a photo-
graph of a Ratel from Lake Magadi, Kenya,
————
_ My. D. Seru-Surrn, F.Z.S,, Curator of Mammals and Birds,
exhibited a living specimen, believed to be about six months old,
of the Pigmy Hippopotamus (Cheropsis liberiensis) which had
heen purehased by the Society from Mr. Harry Smith, of
Monrovia, and arrived on November 2nd. Its weight on arrival
was 40 Ibs., its length 24 inches, and height 14 inches. It was
fed entirely upon milk, and appeared to be in excellent condition.
Since its arrival in the Society’s Gardens it had spent most of
Text-figure 1.
Baby Pigmy Hippopotamus ( Cheeropsis liberiensis),,
the daytime asleep in its box, taking to the warmed water of the
tank towards evening. Although so young it proved to be an
excellent swimmer,
Great credit was due to Mr. Harry Smith for having so
successfully reared this little animal since it was but a few weeks
old, and to Mr, R. M. Mitchell, chief officer of the s.s. ‘ Melville,’
who carefully tended it on the voyage home.
Mr, F. A, Mivcnenu-Hepers, 8.2.8, F.LS., F.BGS., gave
an account of his recent expedition through the jungle-region of
Panama, and illustrated his remarks with a series of lantern-
slides of scenery, natives, and large fish,
THE SECRETARY ON ADDITIONS TO THE MENAGERIE, 1097
November 20th, 1923.
Dr. A. SurtH-Woopwarp, F.R.S., Vice-President,
in the Chair.
‘The Srcrerary read the following Report on the Additions
made to the Society’s Menagerie during the month of October,
1923 :—
The registered additions to the Society’s Menagerie during
the month of October were 219 in number. Of these 50 were
acquired by presentation, 26 were deposited, 29 were purchased,
and 114 were born in the Menagerie.
The following may be specially mentioned :—
1 Puma (felis concolor), from Bolivia, presented by Mr. Robert
Macdonald on October 20th.
1 Jaguar (Felis onca), from Parnahyba, Brazil, presented by
Mr. Louis F. Hamley on October 16th.
1 Serval (Felis serval), from Sierra Leone, presented by Capt.
F. R. Ellis on October 22nd.
The Srcrerary exhibited, and made remarks upon, a collection
of Autographs recently presented to the Society by Mr. Hugh 8,
Gladstone, F.Z.S.
Mr. R. 'T. Gunrnen, M.A., F.Z.S8., exhibited, and made remarks
upon, (1) Vertebree of Mesozoic Crocodile showing the course of
intercostal arteries, and (2) a Jaw-bone of Ursus anglicus Gunther,
from the Cherwell Gravels.
No. 244.
ABSTRACT OF THE PROCEEDINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON.*
October 23rd, 1923.
Dr. A. Surru Woopwaprp, F.R.S., Vice-President,
in the Chair.
The Secrerany read a report on the Additions made to the
Society’s Menagerie during the months of June, July, August,
and September, 1923.
Dr. C. W. Anprewers, F.R.S., F.Z.S., exhibited, and made
remarks upon, an imperfect phalangeal bone belonging to one
of the curious clawed Perissodactyls, the Ancylopoda (Chali-
cotheroidea).
Mr. D. Seru-Smiru, F.Z.8., exhibited, and made remarks upon,
a series of photographs of Lion cubs recently born in the Society’s
Menagerie. Mr. Seth-Smith also exhibited some rare varieties
of the Black Mangabey Monkey, Cercocebus aterrimus.
Dr. Cart Axnsoton exhibited a large series of drawings and
photographs of Cave Animals from the Balkans.
Mr. E. A. Spaut, B.Sc., F.Z.S., gave a résumé of his experiments
on acceleration of metamorphoses of Frog-Tadpoles by injection
of Anterior-lobe Pituitary-gland Extract and Iodine.
A paper on “ Observations on the Development of the Sympa-
thetic Nervous System and Suprarenal Bodies in the Sparrow,”
by A. Suppa Rav, B.A., M.Sc., and P. H. Jounson, B.Sc., F.Z.S.,
was communicated by Mr. A. Suspa Rav.
* 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.
2
In the absence of the Authors, the following papers were taken
as read :—H, C. Abrauam, F.Z.S., “ A new Spider of the Genus
Liphistius from the Malay Peninsula, and some Observations on
its Habits”; Matcoum A. Surru, M.R.C.S., L.R.CP., F.ZS8.,
“A Review of the Lizards of the Genus 7'ropidophorus on the
Asiatic Mainland”; J. G. H. Frew, M.Sc., I'.Z.S., “On the
Larval Anatomy of the Gout-fly (Chlorops teniopus Meig.) and
two related Acalyptrate Muscids with Notes on their Winter
Host-plants” ; Arrnur Lovreriper, F.E.S.,C.M.Z.8., “ (1) Notes
on Mammals collected in Tanganyika Territory, 1920-1923,”
and ‘¢(2) A List of the Lizards of British Hast Africa (Uganda,
Kenya Colony, Tanganyika Territory, and Zanzibar) with Keys
for the Diagnosis of the Species.”
‘The next Meeting of the Society for Scientific Business will be
held on Tuesday, November 6th, 1923, at 5.30 v.m., when the
following Communications will be made :—
F. A. Mirenurt-Henors, ¥.Z.8., F.LS., F.B.GS.
The Primeval Jungle of Panama.
G. H. GoLprincu.
Notes on the African Crested Rat (Lophiomys imhausi).
Haron G. Jackson, F.Z.S.
A Revision of the Isopod Genus Ligidiwn Brandt [Crus-
tacea ].
Ivor G. S. Monvraau, F.Z.S.
On some Mammals from Jugo-Slavia.
Ivor G. 8. Monvaau, F.Z.S., and Grace Pickrorp,
—— eee eee Eee
On the Guernsey Crocidura.
Major S. 8. Frower, O.B.E., F.L.S., F.Z.S.
On Additions to the Snake Fauna of Egypt.
Srantey Hrirsr, F.Z.S.
On some new or little-known Species of Acari.
Cuas. F. Sonnrac, M.D., F.Z.S.
Sie A
On the Pelvic Muscles and Generative Organs of the Male
Chimpanzee. ;
The following Papers have been received :—
Cynrit Crossianp, M.A., D.Sc., O.N., F.Z.S.
Polycheta of Tropical Kast Africa, the Red Sea, and Cape
Verde Islands ; and of the Maldive Archipelago,
Matcoutm A. Smirn, M.R.C.S., L.R.C.P., F.Z.S.
New Tree-Frogs from Indo-China and the Malay Peninsula.
Artuur Loveniper, J.E.S., C.M.Z.S.
SS a
(1) Notes on East African Birds (chiefly Nesting-habits and
Endo-parasites), collected 1920-1923.
(2) Notes on East African Snakes, collected 1918-1923.
(3) Notes on Hast African Tortoises, collected 1921-1923,
with Description of a new Species of soft Land-tortoise.
(4) Notes on East African Lizards collected 1920-1928,
with Descriptions of Two new Races of Agama lionotus Blgr.
(5) Notes on Hast African Insects collected 1915-1922.
R. R. Mout, C.M.Z.8.
The Trinidad Snakes.
Mary L. Hert, F.Z.8.
On the Family Linguatulide.
Dr. Francis, Baron Norsca.
Reversible and irreversible Evolution; a Study based on
Reptiles.
J. R. Garroop, M.D.
Two Skeletons of the Cetacean Pseudorca crassidens from
Thorney Fen, Cambridgeshire.
I. W. Swann, B.Sc., F.Z.S.
Investigations on the Comparative Myology of Fishes.
4
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,
ReaEnt’s Park, Lonpon, N.W. 8.
October 30th, 1923.
No. 245.
ABSTRACT OF THE PROCEEDINGS
ZOOLOGICAL SOCIETY OF LONDON.*
November. 6th, 1923.
Sir S. F. Harmer, K.B.E., F.R.8., Vice-President,
in the Chair.
The Secrerary exhibited, and made remarks upon, a photo-
graph of a Ratel from Lake Magadi, Kenya.
Mr. D. Serit-Smitru, F.Z.S., exhibited, and made remarks upon,
(1) a yong Pigmy Hippopotamus purchased by the Society ;
and (2) a specimen of a 'Tree-Shrew found in a tin of Pineapple
Chunks in syrup.
Mr. F. A. Mrrcureni-Hencrs, F.Z.S., F.L.S., F.R.G.S., gave
an account of his recent expedition through the jungle-region of
Panama, and illustrated his remarks with a series of lantern-
slides of scenery, natives, and large fish.
Mr. Arruur Loverives, F.E.S., C.M.Z.S., gave a réswmé of his
papers :—
(1) Notes on East African Birds (chiefly Nesting-habits and
Endo-parasites), collected 1920-1923,
(2) Notes on East African Snakes, collected 1918-1923.
(3) Notes on East African Tortoises, collected 1921-1923,
with Description of a new Species of soft Land-Tortoise.
(4) Notes on East African Lizards collected 1920-1923, with
Descriptions of Two new Races of Agama lionotus Blgr-
(5) Notes on East African Insects collected 1915-1922.
* This Abstract is published by the Society at its offices, Zoological Gardens,
Regent's Park, N.W., on the Tuesday following. the date of Mecting 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 obtnined on the
day of publication at the price of Sixpence, or, if desired, sent post free for
the sum of Six Shillings per anni, payable in ydvance.
6
The Hon. Ivor G. §. Monraeu, F.Z.S., communicated his
paper ‘‘ On some Mammals from Jugo-Slavia,” and gave a résumé
ot his investigations in collaboration with Miss Grace Pickrorp,
on “The Guernsey Crocidura.”
In the absence of the Authors, the following papers were taken
as read :—G. H. Gotprincn, ‘‘ Notes on the African Crested Rat
(Lophiomys imhaust) ” ; Haroun G. Jackson, I'.Z.S., ‘ A Revision
of the Isopod Genus Ligidiwm Brandt [Crustacea]” ; Major 8.8.
Fiowrr, O.B.E., F.L.S., F.Z.S8., “On Additions to the Snake
Fauna of Egypt”; Sranuey Hirst, F.Z.8., “On some new or
little-known Species of Acari” ; Cuas. F. Sonnac, M.D., F.Z.S.,
“On the Pelvic Muscles and Generative Organs of the Male
Chimpanzee.”
‘The next Meeting of the Society for Scientific Business will
be held on ‘Tuesday, November 20th, 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 October, 1923.
The Secrerary.
Exhibition of a Collection of Autographs recently presented
to the Society by Mr. Hugh S. Gladstone, I’.Z.S.
W. E. Le Gros Crang, F.R.C.8.Eng., F.Z.8.
Notes on the living Tarsier.
Dr. Francis, Baron Nopsca.
Reversible and irreversible Evolution ; a Study based on
Reptiles.
J. R. Garroop, M.D.
Two Skeletons of the Cetacean Pseudorca crassidens from
Thorney Fen, Cambridgeshire.
Cyr Crossianp, M.A., D.8c., O.N., F.Z.S.
Polycheta of Tropical Hast Africa, the Red Sea, and Cape
Verde Islands ; and of the Maldive Archipelago,
7
Miss Joan B. Procrer, F.Z.S., F.L.S.
co e
(1) On new and rare Reptiles from South America.
(2) On new and rare Reptiles and Batrachians from the
Australian Region.
The following Papers have been received :—-
Mazscoum A. Suirn, M.R.C.S., L.R.C.P., F.Z.S.
New Tree- Frogs from Indo-China and the Malay Peninsula.
R. R. Moir, C.M.Z.8.
The Trinidad Snakes.
Mary L. Hert, F.Z.S.
On the Family Linguatulide.
i. W. Suann, B.Sc., F.Z.8.
Investigations on the Comparative Morphology of Fishes.
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.
ZooLoaicaL Society or Lonpon,
Ruceni’s Pars, Lonpon, N.W. 8.
November 13th, 1923.
1h
2
No. 246.
ABSTRACT OF THE PROCEEDINGS
ZOOLOGIGAL SOCIETY OF LONDON.
November 20th, 1923.
Dr. A. Suita Woopwarp, F.R.S., Vice-President,
in the Chair.
The Secretary read a Report on the Additions to the Society’s
Menagerie during the month of October, 1923.
The Secretary exhibited an autograph letter of Sir Stamford
Rafiles, one of the principal founders of the Society, which had
been presented by Mr. Cosmo Bevan, and a collection of auto-
graphs of 161 Naturalists, including the first President, Treasurer,
und Secretary of the Society, presented by Mr. Hugh 8. Glad-
stone, I°.Z.8.
Sir Stamford Raffles’ letter was dated August 13, 1821, and
was written to Sir Everard Home. It announced the dispatch
of the head of an elephant, preserved in alcohol, stated that the
Babirussa, then believed to have a wider distribution, was
restricted to Celebes and Moluccas, and discussed other matters
of zoological interest.
In Mr. Gladstone’s gift there were included autographs of
H. W. Bates, the Naturalist of the ‘ Amazon,’ de Blainville,
Bleeker, Francis Day, Sir Joseph Hooker and his father (Sir
William Hooker), D. W. Mitchell, Secretary of the Society 1847-
1859, and Temminck.
* 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 Sia Shillings per annum, payable in advance.
~
10
My. BR. 'T. Gunrieir, M.A., 1.Z.8., exhibited, and made remarks
upon, (1) Vertebrie of a Mesozoic Crocodile showing the course
of intercostal Arteries, and (2) the jaw-bone of Ursus anglicus
‘Gunther, from the Cherwell Gravels,
Mr. W. HK. Le Gros Crank, F.R.CS.Kng., 1Z.8., com-
municated his paper “ Notes on the Living ‘larsier (Zarsius
spectrwir).”
Sir Srpney F. Warwer, K.B.E., F.R.S., Vice-President of the
Society, communicated his paper on ‘Cervical Vertebre of a
gigantic Blue Whale from Panama.”
In the absence of the Author, Dr. J. Ro Ganroon, a réswmé of
his paper on ‘“ ‘wo Skeletons of the Cetacean Pseudorca crassidens
from ‘Thorney Fen, Cambridge,” was communicated by Sir SripNEY
F. Harmer.
Dr. Francis, Baron Noprsca, communicated his paper on
“Reversible and irreversible Mvolution ; a Study based on
Reptiles.”
Tn the absence of the Authors, the following papers were taken
as read :—Cynit CrossuanD, M.A., D.Sc., O.N., F.Z.8., ‘ Poly-
cheta of Tropical Hast Africa, the Red Sea, and Cape Verde
Islands; and of the Maldive Archipelago.” Miss Joan B.
Procrrer, F.Z.5., (1) On new and rare Reptiles from South
America,” “(2) On new and rare Reptiles and Batrachians from
the Australian Region.”
The next Meeting of the Society for Scientific Business will
ve held on ‘Tuesday, February 5th, 1924, at 5.30 p.m.
A notice stating the Agenda for the Mceting will be circulated
early in January.
The following Papers have been received :—
Matcoum A. Suiru, M.R.C.S., L.R.C.P., ¥.Z.8.
New Tree-Frogs from Indo-China and the Malay Peninsula.
il
R. R. Moun, C.M.Z.8.
The Trinidad Snakes.
Many L. Herr, B.Sc., F.Z.S.
(1) On the Family Iinguatulide.
(2) Zoological Results of the Third Tanganyika Expedition
conducted by Dr. W. A. Cunnington, F.Z.S., 1904-1905:
Report on the Linguatulide.
KE. W. Suann, B.S8e., F.Z.8.
Investigations on the Comparative Morphology of Fishes.
H. C. Cuapwick, A.LS.
On some Abnormal and Imperfectly Developed Specimens of
the Sea-Urchin HLehinus esculentus.
Vhe 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.
ZooLocicaL Socirry or Lonvon,
Recent’s Park, Lonpon, N.W.8.
November 27th, 1923.
PAPERS (cont.).
d : : Page
39. On the Larval Anatomy of the Gout-Fly of Barley (Chlorops teniopus Meig.) and two
Related Acalyptrate Muscids, with Notes on their Winter Host-Plants. By J. H.
Frew, M.Sc., 1.Z.8., etc. (Lext-figures LES esclereavess eon ea ane eT eae 783
40. A Revision of the Isopod Genus Ligidium Brandt—Crustacea. By Warotp Gornon
Jackson, °.Z.8. (Text-figures 1-10.)........ SOOCOD0ddooo nn COogH0OdaoadC0G fee
AL. A List of the Lizards of British Territovies in Kast Africa (Uganda, Kenya Colony,
Tanganyika Territory, and Zanzibar), with Keys for the Diagnosis of the Species.
JEN 7 EIEN LON Aa TIN, I OLIDUSR CHIN EYAISL, © Su Seinen oc aude aa oa nude Mee esnel MM CPLA Thos at
42, On some Mammals from Yugoslavia. By lvor G. 8. Montaau, F.Z.S. sooudn0 chit)
- (3. Notes on Kast African Snakes, collected 1918- 1923. By Arruur Lovermen, F.E.S,,
C.M.Z.S. SoC COO OO CG IMAN hi hes OodicB ing cMidbNslGs BiarhoOIN SOMOINiCo Ea aon a 56 899
41. Notes on East African Birds (chiefly nesting habits and endo-parasites) collected
UGROSGee, ling Aum Ioohyeroenoy TOLIDUSL, OLNVASE “Gogeaces sunscuccsudo nc -. 899
45. Notes on Mast African Tortoises collected 1921-1923, with the Description of a
new species of Soft Land Tortoise. By Artnur Lovertnas, F.1.S., C.M.Z.S.
(Pinfigs Ils ts IU) “Ga ooad oolseen podod0000 00 d00ad0 0000 9010600 00090 0000008000000 923
46, Notes on East African Lizards collected 1920-1928, with the Description of two new
Races of Agama lionotus Blgr. By Artnur Lovuripas, F.E.8.,C.M.Z.8. ........ 935
47. On some New or little-known Species of Acari. By Srantey Hirst, F.Z.8. (‘Lext-
figures 1-24.) ..........66 S000 06 O0S0SG000000 0000 0nd nD 0050000090 0mn00D00000 HY
48, On the Pelvic Muscles and Generative Organs in the Male Chimpanzee... By Cuantxs
EF. Sonntaa, M.D., ¥.Z.8. (Anatomist to the Society). (‘Text-figures 58-64.) ...... 100]
49. Notes on a African Insects collected 1915-1922. By Arrnur Lovenrtnen, E.ES.,
50. On the Guernsey Crocidura. By Ivor G. 8. Montagu, F.Z.S., and Gracz Pickronrp... 1043
51. Reversible and Irreversibie Evolution; a me based on eae By Dr. Franets,
Baron Norcsa. (Text-figures 8 & 9.) .. a rop avon rol ela iateuioNe sts) tena reli A oaecucter el saeeee vee. 1045
52. On New and Rare Reptiles from South America. By Joan B. Procter, F.Z.S., F.L.S.
(Curator of Reptiles). (Text-figures 1 & 2.) 20... 1. ccc cece eee ee ee
23. On New and Rare Reptiles and Batrachians from the Australian Region. By Joay
B. Procrer, F.Z.S., .L.S. (Curator of Reptiles). (Text-figures 3 & 4.) .....-.... 1069
54. On Additions to the Snake Fauna of Egypt. By Major S. 8. Frower, O.B.E., F.Z.8. 1079
9d. Cervical Vertebrae of a Gigantic Blue Whale from Panama, By Sir Stoney I. IfArner,
K.B.E., Se.))., F.R.S., F.Z.8. (Pext-figuro 1.) .,.....2....- Strouse oan cite 1085
56. Notes on the African Crested Rat (Lephiomy's imhausti). By G. IL. Gouprisen...... 1091
Alphabetical List of Contributors eye esis teslaves dpa Riot chon sity =) Meth Soe i adiad | l
Index of Illustrations ......ee..005 Rese Nee aie
Bex eye en ara adore rege yk
Le ig er
.
LIST OF PLATES.
1923, Parr IV. (pp. 661-1097).
Page
769
920
H.C, Asranau: PII. Nest of Liphistius malayanus, sp.n. ..
A. Lovenrpen: Pls. 1-1I. Testudo proctere, sp. n.
NOTICE.
i ‘he ‘ Proceedings’ for the year are issued in four parts, paged consecutively,
so that the complete reference is now P. Z. 8. 1928, p.... The Distribution
is usually as follows :— Ny aes
Part. I, issued in Mareh.
ean ecellh, $5 June.
Sud El aa September.
WV ee kas December,
- ————————
High
‘Proceedings, 1923, Part ITL. (pp. 483-659), was published on September 3rd,
1923. ,
.
The Abstracts of the ‘ Proceedings,’ Nos. 244-246, are
contained in this Part.
- [he dates of Publication of ‘Proceedings 1830-1858 will be found in the
‘ Proceedings’ for 1893, page 436.
Tho dates of Publication of ‘Transactions 1833-1869 will be found in the
* Proceedings’ for 1913, page 814. BS
) aed!
This preservation photocopy
was made and hand bound at BookLab, Inc.
in compliance with copyright law. The paper,
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