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COLLECTION 
OF 


WILLIAM SCHAUS 
© 
PRESENTED 
TO THE 
NATIONAL MUSEUM 


MCMV 


y 15 
NE 
Hie 


PROCEEDINGS 


OF THE 
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS 


OF THE 


ZOOLOGICAL SOCIETY 


UE LON DON. 


1917, pp. 1-215, 


WITH 6 PLATES and 82 TExtr-FIGURES. 


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


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


\ >: 
4g perarary geet 
ey RATIONAL ® 


Les 2 


OF THE 


COUNCIL AND OFFICERS 


OF THE 


ZOOLOGICAL 


SOCIETY OF LONDON. 


Loni. 


Patron. 
His Masesry Tue Kina. 


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


‘Tart Hon. Cecrt Barina, M.A. 
R. H. Burne, Esq., M.A. 
ALFRED H. Cocks, Esq., M.A. 


Lr.-Cot. S. Monckton Copsr- 
» MAN, M.D? F.RS. 
CHARLES DrumMmonp, Ksq., 


Treasurer. 

ALFRED Ezra, Esq. 

Capt. Hueu 8. GLADSYONE, 
M.A. 

THE Viscounr Harcourt, P.C. 

SIDNEY Freperic HArMER, KsqQ., 
M.A., Sc.D FRLSs 
President. 

WitiiAM Huntsman, Esq. 

Sir EpmMunp Gites Lover, Br., 
Vice-President. 


Con. Sir A. Henry McManoy, | 


GCM.GAf.8.1. 


Vice- | 


PRINCIPAL 
P. Cuatmers Mircuett, M.A., D.Sc., LL.D., F.RBS., 


Secretary. 


EK. G. B. Mrapg-Watpo, 
Ksq., Vice-President. 

P. CHAuMers MircHe.y, Ksq., 
M.A., D.Se., LL.D., Feiss, 
Secretary. 

THE EARL oF 
Vice-President. 

THEt Marguts oF Stico, F.S.A. 

OLDFIELD ‘THomas,  Ksq., 

AuBYN Trevor-Batrye, Esq., 
M.A. 

ArrsurR Smita Woopwarp, 
Esq@., “LED. F.RoS Vee 
President. ; 

Henry Woopwarbd, Ksq., LL.D., 
F.R.S., Vice-President. 


PoRTSsMOUTH, 


OFFICERS. 


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

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

Lieut. Epwarp G. BouLencErR, Curator of Reptiles. 

Prof, H. Maxweut Lerroy, Curator of Insects. 

Mrs. Peavor, Acting Librarian and Clerk of Publications. 


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


LIST OF. CONTENTS. 


1917, pp. 1-215. 


EXHIBITIONS AND NOTICES. 


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


Wecenn er cal ONO cemaionsas «rele can dae resists Seu e. See aes 


Mr. E. Gerrard. Exhibition on behalf of Mr. Crabb of 
a mounted specimen of a hybrid between a male 
brush and, a temale Blackbird wrx. ....:...c0ccecseetes ss 


The Srcrerary. Report on the Additions to the Society’s 
Menagerie during the month of January, 1917 ...... 


Mr. D. Seru-Smiru, F.Z.8S., Curator of Birds. Exhibition 
of the trachea from a male Anseranus semipalmata... 


Mr. C. J. C. Poot, Assistant Curator of Insects. Notes 
from the Caird Insect House ............. LeAnne eee 


Mr. R. I. Pococx, F.R.S., F.Z.8., Curator of Mammals. 
Lantern-exhibition of the work of the Beavers in the 
mociety s Gardens, ~(Vext-figure 1.)) fo....cc.2.cc... ec 0% 


The Secretary. Report on the Additions to the Society’s 
Menagerie during the month of February, 1917 ...... 


Mr. R. I. Pocock, F.R.S., F.Z.8., Curator of Mammals. 
Exhibition of a young Lion Cub recently deposited 
in the Gardens, and also a Galago from German East 


biel Earl yd A ie oe reer egg orien nD ong nc 


Mr. C. J. Ganan. Exhibition of a living example of the 
“* Death-Watch ” Beetle (Anobsum tesselatwnv) i ee 


Mr. C. Davies SHERBORN, F.Z.S. Exhibition of an auto- 
graph of Captain Bligh of ‘ The Bounty’ ......2.......: 


Page 


96 


160 


209 


209 


209 


210 


iv 


Mr. C. Tarr Reaan, M.A., F.Z.S. Exhibition of a Piraya 
from the Amazon (Serrasalmo pirayd)  .. ..sccceccereeas 

Mr. Atrrep Ezra, F.Z.S. Exhibition of lantern-slides 
of Big-Game Shooting im Ind tai ...6.... 6.0000. -s 0+ ee eee 

The Secretary. Report on the Additions to the Society’s 
Menagerie during the month of March, 1917 ......... 

Mr. C. Davies Suerporn, F.Z.8. Exhibition of the piece 
of hairy Skin of a Gorilla, sent to Richard Owen ...... 

Mr. D. Seru-Smira, F.Z.S., Curator of Birds. Exhibition 
of a female Japanese Pheasant 


any oh 0 Mee ° 
Exhibition of a male specimen of the Carolina Duck... 


evoereer eo eee oer eee eo oeeeese 


Exhibition of some specimens of the trachea of 
various species Of Ducks ....7 2027 .quss say agen eee eee 
Mr. EK. HWeron-Auwen, F.L.S., P.R.M.S., F.Z.S. Remarks 

‘on the Mussel-fishery and Foraminifera of Esnandes, 
illustrated by means of lantern-slides 


weunreseceeerereecee 


The Secretary. Announcement of the death of Mr. Henry 
Peavot 


ereer rere sees eee e ree ee osee tree are eeseosreacseeeeeseseeseseeeosere 


Dr. P CHaumers MircueE.y, F.R.S., F.Z.S. Secretary to, 
the Society. Exhibition of coloured sketch of an 
abnormal Cape Lourie 


eceovoer are eee eee oes ees oe ee ee oee eee eee eee 


Prof. J. P. Hitt, F.R.S., F.Z.8. Exhibition of photograph 
and remarks upon a Gynandromorphic specimen of 
the Karwig 


eoeerecreeose oe ecco es oes ors oee oe eee eeoreeesosesees ore ees 


Dr. A. SmrrH Woopwarp, F.R.S. Exhibition of enlarged 
models of the first molar tooth of a large Chimpanzee, 
Melanesian Man, and Piltdown Man 


oeecuanveeeseeareereaeeens 


Mr. D. M.S. Watson, F.Z.S. Exhibition and diagrams of 
models of the teeth of fossil horses 


e@aneseeneeeeceeee ees eee 


Prof. H. Maxweut Lerroy, M.A., F.Z.8. Remarks on the 
Silkworms and Silk industry of India, illustrated 
with lantern-slides and specimens of the Silk Moths, 
Cocoons, and Silk 


eCeoeocee oe eee tooeee eZee oversee oF OFF HHO HH AEE HOE 


Page 


210 


210 


211 


211 


211 


-211 


211 


212 


212 
212 
213 


213 


213 


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


Dr. P. Cuatmers Mitcuetyi, F-R.S. Secretary to the 
Society. Exhibition of the behaviour of Birds and 
Mammals in the presence of living Serpents............ 


Mr. D. Seru-Smuiru, F.L.S., Curator of Birds. Lantern- 
exhibition of Birds now or recently living in the 
Poclety’s Gardens. .....55.... Be er rSesstile Seem nie theien Wiasati 


Mr. R. I. Pocock, F.R.S., Curator of Mammals. Exhibi- 
tion, on behalf of Messrs. Rowland Ward, Ltd., of 
the skins of two Zebras from British East Africa...... 


Mr. R. E. Howupine. Exhibition and Remarks on a 
WII Tepe Ulnt ss magarenesinaceinnsctaten Males ocorveuns vv ee ques eds oe 


Exhibition of two curiously malformed eggs ............ 


Mr. D. Sersu-Smiru, F.L.S., Curator of Birds. Account of 
the Poultry Exhibition now being held in the 
BS DCT MyM T UE MOMS: 5 ,kiers eh onat 6c hsian do aiplv dd rleren.nerall jeu dies 


PAPERS. ae 


1. On the Pectoral and Pelvic Arches of the British 
Museum specimen of Archeopteryx. By BRANISLAV 
Perronievics, Ph.D., and ArtHuR SmirH Woop- 
wan i) Koes, VP As, (Plate 1.) icc... 


2. On Heude’s Collection of Pigs, Sika, Serows, and Gorals 
in the Sikawei Museum, Shanghai. By ARTHUR DE 
CMRUP ES OWERBN, Br LIISG cet sa o.tucddtaeiuiscehssaicesentouens 


3. Report on the Deaths which occurred in the Zoological 
Gardens during 1916, together with a List of the 
Blood- Parasites found during the Year. By H. G. 
PurmMer, F.R.S., F.Z.S., Professor of Comparative 
Pathology in the Imperial College of Science and 
Technology, London, and Pathologist to the Society . 


214 


214 


val 


4. On the Structure and Function of the Mouth-parts of 
the Palemonid Prawns. By L. A. Borrapaixe, M.A., 
E.Z.S., Lecturer in Zoology in the University of 
Cambridge, Fellow, Dean, and Lecturer of Selwyn 
College. “(Text-figures) 1 [oll s..c2aii5. .24-+ 000m 


5. On the Scolex in the Cestode Genus Duthiersia, and on 
the Species of that Genus. By Frank EH. Bepparp, 
M.A., D.8c., F-R.S., F.Z.8:. (Text-figures 1-5.)...... 


6.—1. The Coleoptera of the Family Cisside found in 
Britain, with Descriptions of two new Species.—2. A 
new Species of the Coleopteran genus Cryptorrhyn- 
chus Mlliger. By C. J.C. Poon, Assistant Curator 
Caird’ Insect” Hious¢ <..22......s20--0:+. canta eee eee 

7. The Prechordal Portion of the Chondrocranium of 
Chimera collie. By Epwarp PuHeEtps ALLIs, Jun., 
E.Z:S. (Plates 1.AIV).).. 2.0502: cee eee 


8, On the Lizards of the Genus Philochortus Matschie. 
By G. A. Boutencer, F.R.S., F.Z.8. (Plates I. & IT.) 


9. An Experimental Investigation of the Migration of 
Woodcock breeding in the West of Ireland. By S. 

R. Doveuas, M.R.C.S., L R.C.P.Lond., Capt. I.M.S. 
(retired), F.Z.8., lst Assistant Bacteriological Depart- 
ment, Medical Research Committee, National In- 
surance Act! icsuptatichis ss ccaaen cope ceetee ee eee ee 

10. A Sketch Classification of the Pre-Jurassic Tetrapod 
Vertebrates. By D. M. 8. Watson, M.Sc., F.Z.S., 

- Lecturer in Vertebrate Paleontology in University 
College, London. (Vext-fleures 1 & 2:) -. ae 

11. Notes on some of the Viscera of an Okapi (Okapia 
johnstom Sclater). By R. H. Burne, M.A., F.ZS. 
(Text-figures 1-232) 25.2. erst ces ee Naat ce eee ee 


Alphabetical List of Contributors 


eco noeseeereeoseeeeese ee ore ose eeeeee 


Index 


eevee eoeoeres ee eres eer eoeoeees ees eee FP HSE HHEHHEHH HOHE H SEH HEHEHE EO HE OF OLEH H ETE HES 


Page 


37 


73 


83 


105 


145 


159 


167 


187 


SERHABE LCA LISt 
OF THK 


CONTRIBUTORS, 


With References to the several Articles contributed by each. 


(1916, pp. 1-215.) 


Page 
AuLEN, E. Heron-. See HERoN-ALLEN, KH. 
Auiis, Epwarp PuHe ps, Jun., F.Z.S. 
The Prechordal Portion of the Chondrocranium of 
Chimeera@ collier. (Plates IolL1.). 2...0cc..cceeescecesceoeees 105 
Bepparp, Frank E., M.A., D.Sc., F.R.S., F.Z.8. 
On the Scolex in the Cestode Genus Duthiersia, and on 
the Species of that Genus. (Text-figures 1—-5.)............ 73 
Borrapaie, L. A., M.A., F.Z.S. 
On the Structure and Function of the Mouth-parts 
of the Palemonid Prawns. (Text-figures 1-51.) ......... 30 


Boutencer, G. A., F.R.S., F.Z.S8. 


On the Lizards of the Genus Philochortus Matschie. 
Peeler ecm re MBE eee ete et as cnr esaesevhec ends Chi costed ees 145 


Vill 
Burne, R. H., M.A., F.Z.S. 


Notes on some of the Viscera of an Okapi (Okapia 
johnstoni Sclater. (Text-figures 1-23.) .............ccceeeee 187 


Doveras, 8.R., M.R.C.S., L.R.C.P. Lond., Captain I.M.S. 
(retired), F.Z.S. 


An experimental Investigation of the Migration of 
Woodcock breeding in the West of Ireland ............... 159 


Ezra, ALFRED, F.Z.S. 
Lantern exhibition illustrating Big-Game Shooting in 


Tndta, 9.255. Bee chess a eedsstes eS ee 2V0 


GAHAN, C. J. 
Exhibition of a living example of the ‘“ Death-Watch ” 
Beetle Wi. caw nen penn lot teen once eee eet oe ee en 209 


GERRARD, EpwARrp. 


Exhibition, on behalf of Mr. Crabb, of a mounted 
specimen of a hybrid between a male Thrush and a 
female? Black omd ai, bicvack atin 's)..teigseb ethene rae 95 


Hrron- Aen, E., F.L.S., P.R.M.S., F.Z.S. 


Remarks on the Mussel-Fishery and Foraminifera of 


Esnandes, illustrated by means of lantern-slides ......... 212 
Mini, Prof. J. P., D.Se., ER.2 CZs. 
Exhibition of photograph and remarks upon a Gynan- 
dromorphic, specimen of the Harwig .:....0.... .....teas- 213 
Ho.pine, R. E. 
Exhibition and remarks on a Cirriped................000- 215 


Exhibition of two curiously malformed eggs ............ 215 


1K 
Lerroy, H. Maxwett, M.A., F.Z.S., Curator of Insects. 


Remarks on the Silkworms and Silk Industry of India, 
illustrated with lantern-slides and specimens of the Silk 


Norns. Cocoonss ad: Silks.) ci5sssee te ocd a iaav enticed evs ceeets 


MircHewi, P. Caatmers, M.A., D.Sc., LL.D., F.B.S., F.Z.S., 
Secretary to the Society. 


Report on the Additions to the Society’s Menagerie 
‘during the months of November and December, 1916 ... 


Report on the Additions to the Society’s Menagerie 
dtnine theamonth-of January, 1917 ....00)...casccsscseecseosh 


Report on the Additions to the Society’s Menagerie 


Gucincsthe month of Mebruary, LOL( ........¢.20.8 avis: ot 


Report on the Additions to the Society’s Menagerie 
Gunine the month of March* 1917 coc... .isecccsccecsesoaced 


Announcement of the death of Mr. Henry Peavot 


Exhibition of coloured sketch of an abnormal Cape 


IONIC Ce ee os ko as une hllee bale vavonewaebaewwwe ts 


Report on the Additions to the Society’s Menagerie 
donne the month of Aprils loll sii.scicerse cst sectensss cece 


Exhibition of the behaviour of Birds and Mammals 


im the presence of living Serpents  .../.......cc.ssseneeeeseds 


PrtTroniEvics, Branisitav, Ph.D., and Woopwarp, ARTHUR 
Sei. (lb Dy HRS. -V.P:Z.S. 

On the Pectoral and Pelvic Arches of the British 

Museum Specimen of Archwopteryx. (Plate I.)............ 


PurMer, H. G., F.R.S., F.Z.8., Pathologist to the Society. 


Report on the deaths which occurred in the Zoological 
Gardens during 1916, together with a List of the Blood- 
arasiues ound, during the Year's .)2.5 0). ...e.cceeeneseseed ess 

Proc. Zoot. Soc.—1917. b 


Page 


213 


95 


96 


209 


Aah 


212 


212 


213 


214 


x 


Page 
Pocock, Reernaup I., F.-R.S., F.LS., F.Z.8., Curator’ of 
Mammals. 
Work of the Beavers in the Society’s Gardens. (Text- 
fioure U.) vis .c.30Seeh eae eenercea cei eee ce oe eee 08 
Exhibition of a young Lion Cub, and of a Galago from 
German’ Hast Africa 22s. iscsi eee ee ee 209 
Exbibition, on behalf of Messrs. Rowland Ward, Ltd., 
of the skins of two Zebras from British East Africa ...... 214 
Poors, C. J. C., Assistant Curator Caird Insect House. 
The Coleoptera of the Family Cissidee found in Britain, 
with Descriptions of two new Species ’...............eseeeeees 83 
A new Species of the Coleopteran Genus Cryptorrhyn- 
Chas MN ers cee uate steams anasto ee eee REA SEE G4 93 
Notes from the Caird Insect House........ ee a ci. 96 
Rea@an, C. Tats, M.A., F.Z.8. 
Exhibition of a Piraya from the Amazon ............... 211 
Sera-Sairu, D., F Z.8., Curator of Birds. 
Exhibition of the trachea of a male Anseranius 
SCI PANIMGLG so 9) 5. sr erage each nc ee eaes ace nen ene 96 
Exhibition of a female Japanese Pheasant ............... 211 


Exhibition of a male specimen of the Carolina Duck... 211 


Exhibition of some specimens of the trachea of various 


species "Ola uckis.. ecg hone SS sins Geese exer oee seamen 214 


Lantern-exhibition of Birds now or recently living 


in ‘the Society s' Gardens 205.25.) saeaeeeh eset ne wea 214 


Account of the Poultry Exhibition now being held 


in the Society’s Gardens .............0+5 Pane ain ae gee A 5 215 


Page 
SHERBORN, C, Davigs, F.Z.S8. 
Exhibition of an Autograph of Captain Bligh of ‘The 
Bounty, ene MANE ets os Perera e en haalts eahan eats 210 
Exhibition of the piece of hairy skin of a Gorilla, sent 
OMMOMATORO WET Wo 24aiscccswas beste saves ye bdentes deeeanscnaees’ 211 
SowErBY, ARTHUR DE CaRLE, F.Z.S. 
On Heude’s Collection of Pigs, Sika, Serows, and 
Gorals in the Sikawei Museum, Shanghai .................. 7 
Watson, D. M.8., M.Sce., F.Z.S. 
A Sketch Classification of the Pre-Jurassic Tetrapod 
Nerteprates:, (Rext-fguros' ] dy 2.) ve... cecssececeesuessormaes 167 


Exhibition of models of the Teeth of Fossil Horses .... 213 


Woopwarp, ArTHUR SmirH, F.R.S. See Perronievics, 
BRANISLAV. 


Exhibition of enlarged models of the first molar tooth 
of a large Chimpanzee, Melanesian Man, and Piltdown 
Iya Oo ee Ee oe ee sie di 


= 


P.Z.S. 1917. PETRONIEVICS & WOODWARD, PL. I. 


ARCHAOPTERYX. 


PROCEEDINGS 


OF TIIE 


GENERAL MEETINGS FOR SCIENTIFIC BUSINESS 


OF TIIB 


ZOOLOGICAL SOCIETY OF LONDON. 


PAPERS. 


1. On the Pectoral and Pelvic Arches of the British 
Museum Specimen of Archwopterys. By BRANISLAV 
PeTrontievics, Ph.D., and ArtHur Smira Woopwarp, 
iD. ERS... V.P.Z.S. 


[Received and Read November 21, 1916.] 


(Plate I. *) 


INDEX. Page 
MCC Gal aleAU Cis 015 ae tlhe icess aueners tease sabrina, 2 
REV ACM ACIP ete nuk Penta te. has cndow ks cies Me ciate Sets ose e8S oD 
Bibliography 6 
Introductory. 


Some further preparation of the British Museum specimen of 
Archeopteryx recently done by Mr. F. O. Barlow, under the 
direction of myself and Dr. Woodward, has resulted in the com- 
plete uncovering of the right coracoid bone and the discovery of 
the pubes. The work was undertaken at my request after I 
had convinced myself, by a prolonged study of this specimen in 
connection with Dames’ two memoirs on the Berlin Archeopteryx, 
that the bones in question must exist buried in the limestone 
matrix. On the results I have prepared an exhaustive paper, 
which I hope to publish later; but the following preliminary 


* For explanation of the Plate see p. 6. 


Proc. Zoou. Soc.—1917, No. I. 1 


2 DRS. B. PETRONIEVICS AND A. S. WOODWARD ON THE 


notice, written by Dr. Woodward, who has made use of our 
joint observations, gives a general account of the important 
additions to our knowledge of the pectoral and _ pelvic arches of 
this primitive Jurassic bird which are now made possible. Some 
differences of opinion between us are indicated in footnotes. 

I have to thank Dr. Woodward and also Dr. C. W. Andrews 
for the valuable help and advice they have given to me.—B. P. 


Pectoral Arch. 


The right scapula and coracoid are completely exposed on their 
outer face (Pl. I. figs. 1, 1@) and apparently almost uninjured. 
They meet in a very wide angle and are fused together, though 
their limits are marked by a line across the glenoid cavity and a 
notch at its outer edge. 

The scapula (sc.), which has already been’ described by Owen 
(1863) and Dames (1884), is typically avian and about twicé as 
long as the coracoid. Its distal end is slightly expanded, as in 
Casuarvus ; while its proximal end bears a well-developed acromial 
process (a), from which a very thin lamina of bone, with 
roughened surface, extends to the coracoid, forming the floor of a 
hollow (f.) in which the furcula evidently articulated. 

_ The coracoid (co.) is a quadrilateral plate of bone longer than 
‘wide, and not much narrowed at the upper or humeral end, 

where it is somewhat thickened. The bone is bent at an obtuse 

angle along a hne extending obliquely from the outer end of the 
‘humeral border to the inner end of the sternal border, so that the 
infero-external half must have been almost in the same plane as 
the expansion of the sternum, while the sapero-internal half faces 
inwards. The wide upper portion of this internal half is pierced 
near’ the middle by the usual coracoid foramen. The thickened 
upper or humeral border bears two eminences, a larger adjacent 
to the margin of the glenoid cavity and a smaller at the outer 
angle of the bone, where it extends a little down the outer border 
(fig. la). Below this eminence the outer border is thin and 
slightly excavated, meeting the equally thin but straight lower 
(or sternal) harden in an obtuse angle. The inner border, which 
meets the lower border in a right angle, appears to be still 
thinner and forms a somewhat jagged edge in which two indents 
-are probably natural*. The upper and larger of these, which 
is semicircular, occurs at the upper end; the smaller indent 
- further down is bounded below by a slight prominence yt. The 


* T consider that the upper indent is certainly natural.—B. P. 

+ I think I can distinguish a faint line extending from the hollow between the 
upper eminences to the middle of the lower indent of the inner border. This may 
' perhaps mark the division between the precoracoid and the true coracoid—in which 
case the glenoid eminence would be on the former, the second eminence on the latter. 
The eminence on the coracoid would thus be homologous with the spina coracoidea 
of the Ratites and the acrocoracoid of the Carinates, while the intervening hollow 
would correspond with the fossa supracoracoidea of the Ratites (cf. M. Fiirbringer, 

‘Untersuchungen zur Morphologie und Systematik der Vogel,’ vol. i. (1888) pp. 36, 
39, 40).—B. P. 


PECTORAL AND PELVIC ARCHES OF ARCH HOPTERYX. 3 


irregularities | in the rest of the Morden are probably due to 
imperfections i in the fossil. 

As remarked by Huxley (1868), the furcula (PI. 1. He. 2) is 
exposed from its ventral face. The new preparation ‘of the 
specimen has only defined more clearly the shape of the upper 
articular end on the left side * 


TABLE OF MEASUREMENTS. 


Scapura: mm. 
Greatest length to.apex of acromion ..................020000008 46 
Greatest breadth of distal end) s.0......acl.etsc veces eseaess, 6 

= MnOmtMm allen aes .a57; ssuasnes. otees setae ook 
Simallest Mreadth:. s..8c2.ccu eee re 
Greatest diameter of scapular fart a ileaoid ara Be Rinse 5 

CORACOID: 

Greatest length from prominence above glenoid fossa to 
ITLOHO- Mater NA ANC... ce syis. yet dese tosses eseeegeoGe Rageoy 29 
Breadth at upper (haenetall) Bouin Ea AE aes ait ye 
Breadth at middle.. BOER, Berio ae EO ae cracls AL 
' Breadth at lower aberualh homers pron beter poe ele 
Greatest diameter of coracoidal part of alenoid Pisce picthe’ 3 

FURCULA: 

PEM SHON OL Nena ieatecnstoaiesav cs lem ev.se@eaungevecreclere med ind! 2 
MGeMMC SU WIC malin tein caht sens testestamcassuatena sceessacconasss OUND 
2 KGnreaucetenTedativOr DONE! oi). hs ecacesmecudscecens ch tcesetcavesss ° @ 


Inner angle of furcula about 90°. 


It would be interesting to compare the pectoral arch of 
Archeopteryx, as now made known, with the corresponding parts 
in the second specimen in Berlin; but, unfortunately, according 
to the description by Dames (1897), the several bones in the 
latter fossil are so imperfect, crushed, and obscured that no 
satisfactory comparison is possible. It must suffice to note that, 
although Archeopteryx was obviously a bird of flight, its scapulo- 
coracoid is more closely similar to that of certain Mesozoic 
reptiles than that of any other known bird, and the nearest 
approach in shape to its coracoid is found in the existing Ratites 
and the Cretaceous Hesperornis. In the fusion of the scapula 
with the eoracoid and the very wide angle at which these bones 
meet, it agrees only with the ratite birds and the reptiles. 


Pelvic Arch. 
In the pelvie arch (Pl. I. fig. 3) the right ilinm (é.) and 


ischium (és.) are exposed on their outer face, while the two 
ae (pu.) are displaced and seen from above. The bones were 


* A small prominence on the stone below the middle of the furcula may perl; ips 
mark the anterior end of the sternum (BIOA. fiet 2.5.) B, P. 
+ [ venture, however, to make the comparison, ‘notwithstanding naperfeetions In 
the Berlin specimen.—B. P. 
1 


4 DRS. B. PETRONIEVICS AND A. S. WOODWARD ON THE 


evidently all separate, and the illum must have been only loosely 
articulated with the sacrum, which is proved to be absent by an 
exploratory excavation made in the stone beneath the pelvis. 

The relatively large preacetabular portion of the ilium (/.) is 
elliptically rounded in front, and the hinder half of its lower 
border is gently excavated where it forms the sharp outer edge 
of the concave anterior face of the large deep pedicle for the 
articulation of the pubis. The middle of the outer face of the 
bone is, deeply concave, while its lower margin is convex. The 
acetabulum is deeper than wide and, though perforate, is partly 
closed by a thin marginal lamina of bone, which is preserved at 
its anterior border. Above the acetabulum the ilium 1s slightly 
convex, but bears no trace of an anti-trochanter. The post- 
acetabular portion of the bone is a relatively small narrow exten- 
sion, slightly arched and tapering to its hinder end, which is 
imperfect in the fossil. Its outer face is gently concave. The 
posterior pedicle is also small, and obliquely truncated for the 
articulation of the ischium. 

The ischium (2s.) appears to be in its natural position, but is 
distinctly separated by suture from the ilium. It is expanded 
proximally, though slender behind, and bends inwards at its 
distal end, where it bifurcates into two sharply separated pro- 
cesses. The upper of these processes is the longer and curves 
upwards; the lower and shorter process is nearly straight. The 
anterior border of the proximal expansion is slightly excavated to 
form about half of the posterior rim of the acetabulum. Above 
and below this there is a short articular surface for the ilium and 
the pubis respectively. Shortly behind the former articulation 
the upper border rises into a quadrangular process ; and there isa 
second but less elevated process of nearly the same shape further 
back. The anterior portion of the lower border is strongly 
convex. The outer face of the proximal expansion of the bone is 
marked by an ovoid depression, at the bottom of which there is a 
foramen. 

The pubes (pu.) are long, slender, and nearly straight bones 
projecting backwards far beyond the ischia, where they meet with- 
out fusion in an extended symphysis. The proximal articular 
end, best preserved on the right side, is convex and ovoid, its 
long diameter being transverse. The slender shaft of the bone 
curves slightly upwards to this articular end, and in the bend 
there is a large foramen, well seen on the left side. The lower 
face being obscured, it 1s impossible to determine whether or no 
a pectineal process is present. As the bone approaches the hinder 
symphysis, it gradually widens in a hor izontal plane at its inner 
edge until its maximum width is reached at the beginning of the 
symphysis. The apposed pair of nearly horizontal lamine taper 
gradually backwards until they end in a slightly upturned point 
and are lost in an irregular mass of calcite («.) which probably 
represents original cartilage. 


PECTORAL AND PELVIC ARCHES OF ARCH AOPTERYX, 5 


TABLE OF MEASUREMENTS. 


ILium: mm. 
Greatest length (as preserved) ............... Parc OO) 
Lezgth from middle of aekerion aeetabalae bards to 

RCIBIOLE (NG AAA ee eee ei seen an enn orca Meier 23 
Same measure to posterior end (as sa a, Berea ean LO 
Width of acetabulum ........... Pee ee eet O 
Greatest depth (at articulation for ane Mo mostGeeae LO 

IscHIUM: 

Greatest length .. tes Bee ener ener “76 
Greatest breadth (at proce re Be ktoenmes te asain ce (iL () 
Weastubreadth (near middle): ) ..tii. csc esctsgntaeeeccccene 8 
Pusis: 
Greatest length ............... pe he aaa meg, 
Length of median symphysis Sot fades expansion ......... 19 
wleast-breadth (near proximal end) 2... .c0...0..ccseec-ceeees = 2 
Greatest breadth of hinder expansion ..............0.c00008 A 


The pelvis of Archwopterya, as now described, can be more 
satisfactorily compared with that of the second specimen in 
Berlin than the pectoral arch to which reference has already 
been made. In the pelvic region, however, the Berlin specimen 
is Somewhat differently crushed, all three elements being exposed 
in direct side-view, with the femur in its natural position over- 
lying them and obscuring the proximal end of the ischium. The 
figure published by Dames, 1897 (copied in Pl. I. fig. 4), is, 
indeed, rather diagrammatic, and suggests imperfections in the 
state of preservation of parts of the bones, which must be taken 
into account. Small differences in shape between the ilium and 
ischium of the two specimens have already been noted both by 
Seeley (1881) and by Dames (1897), who include them among the 
marks of at least specific distinction which they recognise in 
the two skeletons. Still more striking differences now appear, at 
first sight, between the pubes; but 1t must be remembered that 
in the British Museum specimen these bones are seen from 
above, while in the Berlin specimen the right pubis alone is 
exposed in outer side-view. Making allowance for the imper- 
fection of the proximal ends in the latter case, the relative length 
of the ischium and pubis is approximately the same in the two 
fossils, and the terminal expansion in the Berlin specimen may 
well be the apparently cartilaginous mass («#.) of the British 
Museum specimen crushed from side to side. Essential differ- 
ences, therefore, are not yet demonstrated *. 

Among existing birds, the pubes meet in a symphysis only in 

* T am of opinion that future discoveries will result in demonstrating such 
differences. I do not think that a side-view of the pubes in the British Museum 
specimen would correspond with that shown in the Berlin specimen: I consider that 
the cartilaginous end of the British Museum specimen is not homologous with the 
bony expansion of the Berlin specimen ; and I conclude that there cannot have been 
any pubic symphysis in the latter. I therefore refer the so-called Archeopterys 


siemensi to a distinct genus Archeornis, which will be defined by differences in the 
pectoral arch as well as in the pelvis.—B. P. 


6 ON THE PECTORAL AND PELVIC ARCHES OF ARCH HOPTERYX. 


one genus, the Ratite Struthio; and here there is no resemblance 
in shape’to the corresponding symphysis in Archwopteryx. The 
nearest approach to the form of the pubis in the fossil appears 
to occur in Colymbus, where the incurved distal end of the long 
slender bone expands inwardly into a delicate triangular lamina, 
but does not meet its fellow of the opposite side. 

While distinctly avian, therefore, the pelvis of Archwopteryx, 
like the pectoral arch, is more closely similar to that of certain 
reptiles than that of any other known bird; and the only exist- 
ing bird repeating its most characteristic feature, the symphysial 
union of the pubes, is a ratite. 


BIBLIOGRAPHY. 


1863. R. Owmn, “On the Archeopteryx of von Meyer, with a 
Description of the Fossil Remains of a Long-tailed 
Species, from the Lithographic Stone of Soienhofen.” 
Phil. Trans. 1863, pp. 33-47, pls. i.-iv. 

1868. T. H. Huxury, “ Remarks upon <Archeopteryx lithogra- 
phicw.” Proc. Roy. Soc. vol. xvi. pp. 243-248. 

1881. H. G. Sesuny, ‘On some Differences between the London 
and Berlin Specimens referred to Archeopteryx.” Geol. 
Mag. |[2} vol. vill. pp. 454-455, pl]. xii. 

1884. W. Dams, ‘“ Ueber Archeopteryx.’ Paleont. Abhandl. 
vol. 11. pp. 119-196, pl. xv. 

1897. W. Damss, ‘‘ Ueber Br ustbein, Schulter- und Beckahaneel 

der Archaopter Uae Sitzungsb. k.-preuss. Akad. Wiss., 
Jahrg. 1897, pp. 818-834. 


EXPLANATION OF THE PLATE. 


Fig. 1. Archeopteryx macrura Gwen; fore limb as displayed on the slab of stone, 
about four-thirds nat. size —Lithoeraphic Stone (Lower Kimmeridgian) ; 
Solenhofen, Bavaria. {British Museum no. 37001. | 
Coracoid of same in front view. 

oe Furcula of same specimen, ventral face, about four-thirds nat. size. 
3. Pelvis of same specimen as displayed on the slab of stone, about four-thirds 
nat. size. : 
4. Archeopteryx siemensi Dames; pelvis, right side view, about nat. size.— 
Ibid. [Berlin Museum of Natural History.]| After Dames, 1897. 


a., acromial process of scapula; c., carpus (with distal extension probably of 
calcified tendon); ¢o..-coracoid; f., facette for furcula; fe., femur; /., humerus ; 
il, ilium ; és.,ischium; pw., pubis; 7., radius; s., supposed anterior end of sternum ; 


se., scapula; 2, uina ; Le caleite probably representing inpertectly calcified car 
tilage. 


= 


ON HEUDLK’S CHINESE MAMMALS. 


2, On Heude’s Collection of Pigs, Sika, Serows, and 
Gorals in the Sikawei Museum, Shanghai. By Arruur 
DE CARLE SOWERBY, F’.Z.S. 


{Received October 9, 1916: Read February 20, 1917. | 


INDEX. Page 
PETS PECIES* OLS US a! g.caeuis cay sesh ss teen eee ee LL 
3 ne WEPEUS Herth na ee Se eee HELO 
5 3 COMENCOT MIST hn tel fe RN ee OU) 
a 5 INGINOTMCAUS A oo see vuahinceteabnninateoas, Om 


For many years the numerous species of mammals described 
or named by Pere Heude in his ‘ Mémoires concernant |’ Histoire 
Naturelle de Empire Chinois’ have been a stumbling-block in 
the path of naturalists who have tried to arrive at a proper under- 
standing of the mammalogy of China and adjacent countries. 

Pére Heude in the eighties and nineties of the last century, 
with the help of numerous Catholic missionaries in the field, 
gathered together a fine collection of mammals, birds, reptiles, 
and other forms of animal life in the Sikawei1 Museum at 
Shanghai. With his peculiar ideas on what constitutes specific 
characters in animals, he set about classifying and naming 
such mammals as came into his hands, with the result that he 
enormously multiplied the number of species in China, especially 
in the genera Sus, Cervus, Capricornis, and Nemorhedus, thereby 
reducing the subject to a state bordering on chaos; subsequent 
workers finding themselves confronted with such bewildering 
facts as eight species of pigs and eleven species of sika (six from’ 
one locality and seven from another) scattered over China, not 
to mention some seventeen species of goral and seven or eight 
bears. That such could not really be the case was obvious, but 
without good series of specimens from Heude’s type-localities, or 
at least his own specimens for examination, the matter could not 
be cleared up. 

During the past few years collectors and sportsmen have secured 
a few specimens of the larger mammals such as pigs, serows, gorals, 
and bears, but the material has been altogether insufiicient to be 
of much help. 

I believe attempts have been made to get hold of Heude’s 
collection, either by purchase or exchange, for some of the more 
important museums of Europe and America, but without success. 

There remained therefore only two things to be done—either 
the securing of series of specimens from all “of Heude’s collectin g- 
grounds, or the revision, on the part of someone fitted for the 
task, of his collection in the Sikawei Museum. 

In 1914 Mr. Gerrit S. Miller, jun., of the Division of Mammals, 
Smithsonian Institution, suggested to me that I should attempt 


8 MR, A. DE CARLE SOWERBY ON 


both of these tasks. JT agreed to try, but so far have found the 
collecting of the mammals in question from the saine or near-by 
places as those whence Heude’s specimens were secured an almost 
hopeless task. 

In the first place, many of the species have become very rare in 
the past few years. Then, too, the conditions under which one 
has to work can never yield the same results as were obtained by 
Heude’s collectors. A protracted residence in each type-locality 
would be necessary, while one would have to enlist the services of 
local native hunters with a knowledge of the country and quarry 
to secure the required specimens. For instance, the difficulty 
of securing specimens of the serow may be gathered from the 
account of Fenwick Owen’s recent trip through Central China. 

Four successive trips into Manchuria on my part have yielded 
only one specimen of pig, four bears, three wapiti (or red deer), 
one roe, and two gorals. No sika were secured, though specimens 
in captivity were seen. ‘These trips have taken a considerable 
amount of time and money, and the poor results, so far as the 
larger mammals are concerned, have been disappointing in the 
extreme; but, with the great increase of settlers, the cutting 
away of the big forests, and the unceasing hunting on the part 
of natives, to which the animals are subject, one could expect 
nothing else. 

On the Yang-tze, where, in Heude’s time, the sika seems to 
have been so plentiful, that beautiful deer is now very rare, while 
wild pigs are noticeably fewer than they were, large specimens 
being particularly difficult to secure. 

A trip made by me last spring (1915) into the high moun- 
tainous region of South Shensi after big game yielded only a 
couple of takins. Serows and gorals, although said to be plentiful 
by the natives, were not even seen, Several hunting and col- 
lecting expeditions into Central and West China in the past few 
years have yielded little better results. Fenwick Owen, already 
mentioned, got one serow, but neither goral nor wild pig. 
J. W. Brooke got a goral and a couple of serows, which the late 
Mr. Lydekker mentioned in a paper read before the Zoological 
Society of London in 1908. 

Mr. Malcolm P, Anderson’s last expedition yielded no speci- 
mens of the serow, though he went right through the country 
inhabited by those animals. He secured a goral and a wild pig 
in the Tai-pei-shan region of South Shensi, 

On the Clark expedition in 1909 I secured a goral in the 
mountains south of Si-an-fu and a fineadult boar in North-Central 
Shensi, near Yen-an-fu, I have also secured gorals and wild pigs 
in North and West Shansi, but the mammals of this province, 
with the exception of the sika, seem to have escaped Heude’s 
attention, so that my specimens can have little or no bearing 
upon the subject. 

Since, then, the collecting of series of specimens from Heude’s 
type-localities was going to prove a long if not a hopeless 
task, there remained only the alternative of going over his 


HEUDE’S CHINESE MAMMALS. 9 


material in the Sikawei Museum. Through the courtesy of the 
Jesuit Missionaries in charge I was able to do this, and the 
following is an attempt to arrive at some definite conclusion as 
to the validity or otherwise of Heude’s numerous species, and to 
determine therefrom the number of distinct species contained in 
the genera Sus, Cervus, Capricornis, and Nemorhedus in China, 
ores, Me Aare, and ihe adjacent portions of Thibet. 

Without intending to cast any slur upon the present curator 
of the Museum, I must state that I found the specimens in some 
confusion, due to the fact that Heude died in the midst of his 
labours, leaving no one to fill his place or carry on his work. 
The specimens were not numbered ; some of them even lacked 
labels, so that it was with great difficulty that I found many 
of the specimens of the species described and figured in the 
publication already referred to in the beginning of this paper. 

Owing to the inevitable ravages of time and insects in such a 
climate as that of Shanghai, I found the condition of the skins 
very poor, and so was forced to confine my observations almost 
entirely to the skulls of the specimens, but where possible I went 
over the skins as well and drew conclusions from them. In 
regard to the pelts I found that Heude had evidently regarded 
seasonal differences in colouring as specific, just as with the skulls 
it was obvious that he had made specific differences out of those 
due to age and sex. How so able a naturalist came to make 
these errors it is difficult to understand ; and I want to make 
it clear that in spite of his peculiar views on the classification 
of species, Heude must be considered as one of the fathers of 
Chinese mammalogy just as Pere David was the father of Chinese 
ornithology. 

As regards the names of the species upon the labels attached 
to the specimens, it was no uncommon thing to find that the 
original name had been erased and substituted by another, which, 
in some cases, was again erased and replaced by a third and 
fourth. Sometimes one name would appear on the label, while 
another would be written on the skull itself, and only in a very 
few cases was the word type to be found. 

However, after going over the specimens carefully, I am 
satisfied that I found most of the skulls figured by Heude. 

Another point to be noticed is that many of the specimens are 
labelled from certain localities, while, in the descriptions of the 
species, other, and in some cases somewhat misleading, localities 
are assigned. Thus, of the sikas from the Poyang Lake District, 
seven species are labelled by the original collector ‘ Aventé.” 
In Heude’s descriptions of these species, he gives the localities 
variously as “hills of Kienté,” ‘‘ territory included between the 
north-east of the Poyang Lake and the Blue River,” “ Central 
China,” “ Poyang Lake,” etc. As a particular instance, take 
the specimen marked Sika lachrymosus. In the collection it is 
labelled Aienté by the collector, yet Heude’s description of it 
gives its locality as Central China, which, to say the least of it, is 
misleading. 


10 


1 


2 


10. 


ie 


16. 


17. 


. Kuangsi 


MR. A. DE CARLE SOWERBY ON 


Before going into details about the specimens themselves, it 
would be well to get an accurate idea of just where Heude’s 
collections were made, and, in making out the following list, L 
have relied more upon the localities on the labels of the actual 
speimens than upon those given in the descriptions. I have been 
at some pains to get the list as accurate as possible. 


. Ning-kuo-fu ...... 


EO INTEMtO ie eee 


ay Lomelicouuee. sec. 


. Tchen-kiang 


Tchang-cheou 


. Chao-hing-fu ...... 


aKa lisienieeeeree 


. Kouang-tung ...... 


Miechanoy ee aa 


Kiun-tcheou ...... 


. Tchen-kou-ting ... 


. The Upper Han 


Valley. 


. Mountains south of 


Si-an-fu, Shensi. 


. Paoki 


West Ssuchuan ... 


MiG api) seer ae eas 


S. by S.E. of Nanking, Anhui Province. 
Pig and sika were collected here. 


(Chienté) N.E. of Poyang Lake, S. of Anking-fu, Anhui 
Province. 
Pig and sika were collected here. 


(Tang-liu) a little to the N. of Kienté on the bank of the 
Yang-tze, Anhui Province. 
Sika collected here. 


(Chinkiang) on the Yang-tze, below Nanking, Kiangsu 
Province. (The specimen of Sus paludesus described as 
from the Kiangyin River at Nanking was secured by 
Carnie and is marked on the label as from T’chen-kiang.) 

Pig coliected here. 


(Chang-chou) on the Yang-tze between Chinkiang and 
Shanghai, Kiangsu Province. 
Pig collected here. 


(Hsao-shing-fu) S.E. of Hangchow and W. of Ningpo, 
Chekiang Province. 
Serow secured here. 


(Chi-hsien) probably near Hsao-shing-fu, Chekiang Pro- 
vince. 
Goral secured here. 


Province of Kuangsi, South China: 
The specimen Sus meles is labelled Kuangsi, with no further 
particulars as to exact locality. 


(Kuang-tung or Canton) Nan-iong or Nanyang, where the 
specimen of Capricornis collasinus was secured, lies in 
the extreme north-eastern corner of the Province of 
Kuang-tung. 

(I-chang) a well-known Treaty Port on the Yang-tze, above 
Hankow, in the Province of Hupei. 

Goral secured here. 


(Chuin-chou) probably in the vicinity of Yun-yang-fu, 
N.W. Hupei. (Lhe specimen of Nemorhedus fantoza- 
tianus was labelled N.W. Hupei.) 

(Chén-k’ou-ting) in the extreme N.E. of Ssuchuan, close to 
the Shensi-Hupei border. 

A large number of serow and goral came from this spot 
collected by Paul Farges. 

Near Han-chung-fu, 8.W. Shensi. 

Pig and serow collected here. 


Goral collected here. 


(Pao-chi) W. of Si-an-fu, S.W. Shensi (not north, as stated 
in Heude's description) near the Tai-pei-shan region. 

Bear collected here. 

(No more specific locality on label.) 

Serow and goral collected here. 


North-west Ssuchuan and adjacent portion of Thibet. 
Goral, serow, and bear collected here. 


18. Kechen, Keso, and Serow collected here. 


Along Bay, Tongkin. 


HEUDE’S CHINESE MAMMALS. ll 


19. Tsing-lo Hsien .... W. Shansi, near Ning-wu-fu. 
. Sika horns from here. 
20. Mukden ............. Manchuria. 
Pig and sika collected here. 


21. Sungari ............ Valley of the Sungari River, Manchuria. 
Pig and sika collected here. 
22. Vladivostok ...... Primorski, on the Maritime Province near S.E. Manchurian 
border. 


Pig and sika collected here. 
23e-Oussour ......:5..:- (Ussuri) Ussuri Valley, between Kirin Province, Manchuria, 
and Primorski, Siberia. 
Bear, pig, and goral collected here. 
24. Fusan and Ousi .... Corea (Eastern Coast). 
Pig collected here. 


25. Central Japan...... Pig from here. 
26; Halodate:.......-... (Yeso) North Japan Island. 


Bear from here. 


27. Marche................ From which place there is a skull of a pig with the name 
Sus angusticeps inscribed on the label. I cannot discover 
it on the map, but from the specimen I should judge it to 
be in the Yang-tze region. 


Hende also has described two sika from Shantung, but I 
tailed to find the specimens in the Museum. 


Genus Sus. 


Altogether I examined the skulls of thirty (80) pigs from 
thirteen (13) localities, amongst which there were no less than 
eighteen (18) species according to Heude’s classification. After 
careful comparison, however, I could not find grounds for distin- 
guishing more than five (5) distinct species. The following list, 
which gives details of the skulls examined, incidentally comprises 
practically the entire collection of wild pigs’ skulls from China, 
Manchuria, and Japan in the Museum :— 


I 3 skulls. o-¢ of Sus collinus from Kienté. 
2. 4skulls oo of Sus chirodontus from Kienté. 
3. 2skulls oo of Sus stricticeps from Kienté. 
4. 1skull ¢ of Sus acrocranius from Kienté. 
5. 3skulls2¢ 9,14 of Sus flavescens trom Ning-kuo-fu and Li-yang. 
6. 2skulls oo of Sus paludosus from Tchen-kiang (Chinkiang). 
7. 2skulls1¢,12 of Sus nigricans from Tchang-cheou. 
8. lskull ¢ of Sus laticeps from Shensi. 
DA skull ig ot Sus eurtidens trom Shensi. 
10, _1-skull ,3' of Sus oryodontus from Shensi. 
lite skull +2 of Sus moupinensis Milne-Edwards, from Shensi. 
ieee alkene, of Sus meles from Kuangsi. 
13. 2skulls oo of Sus gigas from Vladivostok. 
14. 1skull @ (juv.) of Sus songaricus from Sungari Valley. 
Hos Leskull- of of Sus ussuricus from Ussuri Valley. 
163 92 skulls og of Sus coreanus from Fusan and Ousi, Corea. 
17. lskull ¢ of Sus angusticeps from Marche. 
18. Mskulk ¢ of Sus nipponicus from Japan Central. 


In this list it will be noticed that I have given the localities 
marked on the labels, which, as already mentioned, do not always 


12 MR. A. DE CARLE SOWERBY ON 


seem to agree with the locality given in the description of 
the species. 


1. Sus pALubosus Heude. 


In comparing the skulls of the first seven species in the above 
list (i. «. the species inhabiting the Yang-tze Valley) one was 
struck by the fact that they were practically indistinguishable 
one from the other, except that some were lar ‘ger than others, 
slightly heavier, and with more worn teeth. 

I have no hesitation in saying that the differences occurring 
amongst the skulls of this series were due either to individual 
variation or age. Thus, the skull labelled S. paludosus from 
Chinkiang, being the oldest specimen, as evidenced by the wearing 
of the teeth, was very heavy throughout and proportionately 
broader than any of the others. The type-specimen of S. chiro- 
dontus was longer in proportion to its width than any of the 
others, but not sufficiently so to warrant its separation from them 
as a distinct species, for in this respect it also differed from 
the other specimens labelled chirodontus. One of the specimens 
labelled S. chiodontus from Kienté has S. stricticeps written in 
ink on the palate, and was (judging from erasures on the label 
and marks on the skull) also originally the type of S. collinus. 
S. acrocranius was originally labelled S. chirodontus. One of the 
specimens of S. flavescens from Li-yang was very high in the 
forehead, giving a strongly concave outline to the cranium. It 

was or iginailly labelled S. leucorhinus, but the describer evidently 
Changed his mind about it. This all goes to show that Heude 
himself had considerable doubt about the specific differences of 
his species. 

Taking the whole of the series from the Yang-tze Valley, I 
should not hesitate to class them as one species, with the note 
that the specimens from Kienteé district (Poyang Lake District), 
where the feeding is probably good and_ plentiful, are somewhat 
larger than those from further east (7. e. Chinkiang and Ning- 
koufu districts). 

My own experience in the field while hunting wild pigs goes to 
show that there is a great deal of individual variation amongst 
these animals, some being larger and heavier, though younger, 
than others. Other sportsmen and hunters, natives as well as 
Kuropean, have informed me that their experience is the same 
as mine. 

As the specimen from Chinkiang was the first to be described, 
the name paludosus* must stand for the species occurring in the 
Yang-tze Valley. Chirodontus and other names were mentioned 
earlier in Heude’s publications, but without descriptions. That 
the Yang-tze pig is not Sus leucomystax T. et 8., to which species 
it was originally referred by Swinhoet, is certain from a com- 
parison of Heude’s specimens with a skull of about the same size, 


* Mém. conc. I’ Hist. Nat. de l’Emp. Chin. tome iii. p. 193. . 
+ P.Z.S. Lond. 1870, pp. 639, 640. 


HEUDE’S CHINESE MAMMALS, 13 


wrongly labelled S. ntpponicus, from Central Japan. The skulls 
of the Yang-tze pigs are not only considerably larger than this 
specimen from Japan, but are proportionately much longer. It 
is interesting to note, however, that the Yang-tze pigs, in common 
with most of the pigs in China, have the ‘peculiar whisker-lke 
white patch on the cheek, which gives its name to the Japanese 
pig S. lewcomystax. 


2. Sus MOUPINENSIS M.-Edw. 


In comparing the three adult male skulls from Shensi, labelled 
S. oxyodontus, SN. laticeps, and S. curtidens respectively, with each 
other I can find little grounds for separating them. All three 
were collected in 1883 by the same collector, M. Vidi, and, as 
there are no data to the contrary, one must presume from ‘the 
same locality in the region of the U1 pper Han River, S. Shensi. 

Compared with the type of S. paludosus the Shensi skulls are 
all smaller and proportionately shorter and broader—in fact, the 
broadest of the three (S. daticeps) is actually a few millimetres 
wider than the Yang-tze skull, though it is fully 4°5 centimetres 
shorter. As, however, no description accompanies any of the 
three names Payodontiis. laticeps, and curtidens, while the speci- 
mens appear to agree with Milne-Kdwards’ Sas moupinensis px 
further west, I should suggest considering the Shensi pigs as 
belonging to that species, Aillforthen andl more relable data as 
to their ‘representing a distinct form are forthcoming. Indeed, 
Heude himself applied this name to a skull of a sow from 
the same district from which the other Shensi skulls came. 
Mr. Gerrit 8. Miller has referred a specimen of a wild boar shot by 
me near Yen-an-fu in North-Central Shensi to Milne-Edwards’s 
species. 

It is interesting to note that the wild pigs from Shansi also 
have wide skulls, ane specimens sent by me from the Ning-wu 
district in the west of that Province to the Smithsonian Tnsti- 
tution have been determined by Mr. Miller as S. moupinensis. 
An old boar examined by mein 1910 gave the following skull- 
measurements :— 


Condylo-basal length .......0.......68. 395 mm. 
Zygomatic SGC He ¢ es geRenae iy ok CRCen REE EE 5 175 mm. 
Width across postorbital processes of 

HE OUMU Sbar he reieey eete a see, ake, (27m, 


These measurements show a greater proportional width than 
an even larger skull from Chinkiang secured December 1915 
by “Mr. A. HL: Rasmussen, whose property it remains, which 
measured :— 


Condylo-basalilensth ......2.001...4... 400 mm. 
My SOMAbIC WIG x... eu. cees see cses sees 166 mm. 
Width across postorbital processes 

OM OMUAIS fs ear os cease. <a ae TiO mm, 


14 MR. A. DE CARLE SOWERBY ON 


It will be noticed, however, that in this comparative breadth of 
skull the Shansi pig is intermediate between the Shensi and 
Yang-tze species: for, whereas the skull of the Shansi pig is 
broader than that of the Yang-tze pig, by nearly a centimetre, it 
is only half a centimetre shorter, while the skull of the Shensi pig 
that was broader than that of the Yang-tze pig was some four and 
a half centimetres shorter. iu 

For this reason, and also because of its geographical position, 
it being entirely cut off from the Shensi and Moupin species by 
the Yellow River, the Shansi pig would appear to be a distinct 
form. It certainly does not belong to the Manchurian species, 
which is a very much larger animal with a proportionately much 
longer and narrower skull. 


3. Sus giaas Heude. 


One was confronted with some difficulties in comparing the 
pigs from Vladivostok, the Ussuri Valley, and the: Sungari 
Valley. ‘The two skulls from Vladivostok were of adult males i in 
good condition, that from the Ussuri Valley being of a younger 
male with the cranium missing and that from the Sungari Valley 
being of a young female. 

However, considering that these skulls were all Heude had to 
gO upon in creating his three species, there seems to be no reason 
why the specimens should not be used legitimately as a_ basis 
for a reconsideration of his deductions. Considering the great 
individual variation that occurs amongst pigs of the same species, 
Heude was hardly justified in creating these three species, when 
it is also considered that the regions from which the specimens 
in question were secured are topographically identical. The 
Primorsk or Maritime Province in. the vicinity of Vladivostok, 
the Ussuri Valley, and the source and upper, as well as the lower, 
reaches of the Sungari, form one more or less continuous fovest 
spreading over low hills and marshy valleys. 

Compared with the Yang-tze skulls, the Vladivostok cals 
were distinguishable at onee by their enormous size and propor- 
tionate narrowness. What there is of the Ussuri skull also shows 
this very plainly, while the narrowness of the Sungari skull 
(imm. 2) when compared with a small female skull of the 
Yang-tze species is noticeable. 

T could not discover specimens of iS. canescens or S. mandchuricus 
in the collection, but, as Heude says that specimens were bought 
in the Peking market, having come from Mukden, they must 
originally have been secured in the Manchurian forests, either in 
the vicinity of Harbin or well to the east of Mukden itself. In 
any case they would be inhabitants of the same topographical and 
faunal area as the other three species. I have no doubt that 
there is in this area but the one species of pig which must bear 
the name Sus gigas*, as it was the earliest given. This is a good 


#* Mém. cone. l’ Hist. Nat. PEmp. Chin. tome i. p. 189. 


HEUDE’S CHINESE MAMMALS. 15 


name, not only because of its priority, but because it suggests the 
ereat size of the species, and also because the type-skull and the 
topotypical one are complete and fully adult. 

The two skulls from Vladivostok and that from the Ussuri 
exceed in size any skulls hitherto obtained in China proper, and 
also show a wonderful tusk-development. ‘The lower tusk of the 
smaller of the two Vladivostok skulls measured 10", notwith- 
standing the fact that it was broken off at the tip, while the tusk 
of the larger skull must have exceeded this considerably, but I 
could not extract it for measurement. However, it measured 12” 
in diameter. The largest tusks from China that I have measured 
-were 10” in length (not broken at all) and 13" in diameter. 
None of the fine skulls*in Heude’s collection from China bore 
tusks over 84" in length or 14" in diameter. 

In regard to the Manchurian pigs Heude says (page 111, 
vol]. ii.) that the eastern forms partake somewhat of the dental 
characteristics of S. scrofa, but differ more widely as one goes 
westwards. This may be so, but, as I have pointed out, his 
specimens canescens and mandchuricus in all probability come 
from the same forest area as his specimens wssuricus, songaricus, 
and gigas—themselves representing but the one species. 


4. Sus coREANUS Heude *. 


TI should consider this a valid species. Its skull is altogether 
smaller than that of the Manchurian species, and is also propor- 
tionately shorter than that of S. paludosus, the Yang-tze’ form, 


but it is larger and proportionately longer than the Japanese 
S.. leucomystax. 


5. Sus metEs Heude ft. 


Represented by a single skull from Kuangsi, South China, this 
species I should have no hesitation in confirming, were it not for 
the fact that no description accompanies the name. The skull 
has shorter nasals and a much broader forehead than the type of 
S. paludosus, besides being much smaller. A remarkable differ- 
ence occurs in the upper tooth-row, where there is one premolar 
Jess than in any of the foregoing species. The skull is not so 
short and heavy as that of the specimen from Japan. 


Besides the fine series of adult male skulls mentioned above, 
there are several adult female and young male skulls in the 
collection. A female skull labelled S. moupinensis from South 
“Shensi (7. e¢. somewhere in the region where the other Shensi 
skulls came from) differs but little from a female skull from 
Kienté, though the latter is a little higher in the crest, and is 
also slightly longer and narrower. The height of the crest, how- 
ever, is confined to this single specimen, all the other female 


* Mém. conc. l’Hist. Nat. de Emp. Chin. tome iii. p. 191. 


+. Not described in Mém. cone. ]’Hist. Nat. de_l’Emp. Chin., though the name is 
five). 


16 MR. A. DE CARLE SOWERBY ON 


skulls in the collection being the same in this respect as the 
South Shensi skull. 

Colour.—As regards the colour of the various species I could 
learn nothing from Heude’s collection, as I could not find any 
skins. However, I had the opportunity of looking over several 
skins in the possession of Mr. A. H. Rasmussen of Chinkiang. 
All those he showed me had been secured by himself in the 
vicinity. IL was struck by the fact that though the underwool 
in these peits was brown and the long hairs black, yet the latter 
had such a large portion of their length white or cream that it 
gave the whole skin a very ight appearance. This characteristic 
occurs only in extremely old specimens of the pigs from Shansi. 
There the young males are brown and the old males nearly black. 
Very young specimens of the Yang-tze species have the coat of a 
rich chestnut-colour, which is not the case with the Shansi species. 
I have been told by Mr. Lewis Broome, who has shot wild boars 
in Shansi as well as in Manchuria, that those from Manchuria are 
even blacker than the Shansi ones. The white whisker-like pateh 
on the cheek is more pronounced in S. paludosus than in the pigs 
from Shansi. 

From the foregoing comparisons and notes it would seem that 
there are the following distinct species of Pig represented in 
Pére Heude’s collection, all others given in his papers and 
descriptions being invalid under the present generally accepted 
system of classification :— 


1. Sus paludosus Heude. Yang-tze Valley. Type from Chin- 
kiang, Kiang-su, China. No number. 

2. Sus moupinensis M.-Edw. The Upper Han, South Shensi, also 
Ssuchuan and Kansu, China. 

3. Sus gigas Heude. Forests of Manchuria and the Primorsk, 
Type from Vladivostok. No number. 

4. Sus coreanus Heude. Fusan, Corea. Type from Fusan. No 
number. 

5. Sus meles Heude. Kuangsi, South China. Type same. No 
number or description, but a distinct species. 

6. Sus lewcomystax 'T. et S. Central Japan (specimen in Hende’s 
collection wrongly named S. nipponicus). 


Genus CERVUS. 


The Cervus-sika group of the genus Cervus is represented in — 
the Sikawei Museum by a fine series of skulls and skins, but as 
the latter are now in a poor condition from the ravages of insects 
and the effects of damp, the following notes and observations are 
based chiefly upon an examination of the skulls. 

The range of the Sika in China was formerly very extensive, 
but, owing ‘to the i increasing demand for “‘ hartshorn,” made from 
the horns, when in velvet, of this deer, the animal 1s practically 
extinct in most places and is nowhere plentiful. 


HEUDE’S CHINESE MAMMALS. ; 17 


The Sika is known to occur in a few isolated regions in North- 
west Shansi, in the Imperial Hunting-grounds north-east of 
Peking in Chihli, in the more remote parts of the Manchurian 
forests, in North Corea, in one or two isolated areas along the 
Lower Yang-tze, and probably also in Tonkin and parts of South 
China. 

Heude’s collection contains specimens from the Sungari Valley 
and Mukden in Manchuria, Vladivostok in the Maritime Province 
or Primorsk, Kienté, Tonglieou and Ning-kuo-fu on the Yang-tze, 
Phu-lang-thuang in Tonkin and Tsing-lo-hsien in Shansi. 

Following is a list of Heude’s species, whose skulls were 
examined, giving Heude’s names together with the localities as 
found on the labels :— 


1. Cervus imperialis. Songari Valley, Manchuria. 
»  microspilus, Mukden, Manchuria. 

5 dybowskii, Vladivostok, Primorsk, 

»  manchuricus. Mukden, Manchuria, 
cycloserus. 


»  brachyrhinus., 
»  lachrymosus. 


So Sas Ek Ne etd 


: Kienté, Yang-tze Valley, Anhui; also Tong- 
»  pouvrelianus. r 2 
: E lieou, Yang-tze, Anhui, 
» «© poretianus. 
HO; »  JSrinianus. 
11. » grilloanus. y 
12. 5»  vrivierianus, Tong-lieou, Yang-tze, Anhui. 
13. 5 (name illegible), Kienté, Yang-tze, Anhui. 
14. »  andreanus, Ning-kuo-fu, Anhui, 
poy »  dugennianus, Phu-lang-thuang, Tonkin, 
16. 3 «6 gvassianus, (Pair of horns only) Tehing-lo-hsien, Shansi. 


An examination of the skulls assigned by Heude to the first four 
species in the list, 7.e. those from the forested areas of Manchuria 
and adjacent Primorsk, convinced me that these forested areas 
are inhabited at the very most by but two species, namely, Cervus 
mantchuricus Swin.* and Cervus dybowskti Tacz. + 

The skull of the adult male labelled ¢mperialis from the Sungari 
Valley was practically identical with the skull of an adult male 
labelled dybowskii from Vladivostok, the only noticeable difference 
being that the former (imperialis) was shorter and broader than 
the latter (dybowskii). Two adult female skulls of imperialis and 
dybowskit showed just the reverse, the imperialis one being longer 
and proportionately narrower than the dybowskii one. However, 
this lengthening out of the skull seems to be a characteristic of 
age, for in both cases it was the older skull that was the longer. 
From this it would appear that the Sika from the Sungari Valley 
is identical with that from the Primorsk, but the latter has been 
described as having a pure white muzzle, which is not the case 
with the former, so that the two are probably distinct. 

The skull of a young male labelled microspilus from Mukden 


* P.7.S. 1864, pp. 168, 169. 
+ P.Z.S. 1876, p. 123. 


Proc. Zoou. Soc. —1917, No. II. 2 


18 MR. A. DE CARLE SOWERBY ON 


was very similar to that of a young male labelled mandchuricus 
also from Mukden, though the former, the younger, was shorter 
proportionately than the latter. In connection with these two 
specimens it should be noted that in all probability they did not 
come from the immediate vicinity of Mukden, but were brought 
there for sale by native hunters or were kept in captivity there. 
The nearest forest country suitable for these deer to inhabit, even 
as long ago as 1899, could only have been a hundred miles or so 
to the east on the borders of the Fengtien (formerly Shenking) 
Province. 

The name Cervus mantchuricus Swin. undoubtedly applies to 
the species inhabiting North and Central Manchuria, and Cervus 
dybowskii to the species inhabiting the Primorsk. To which of 
these two species, if either, the North Corean Sika belongs cannot 
be determined here. 

The specimen labelled Cervus grassianus from Tching-lo-hsien 
(Tsing-lo Hsien), Shansi, consisting of a pair of horns only, 
afforded but little scope for comparison with the other specimens 
in the Museum, but it was noticeable that the horns were large 
and heavy, resembling those from Manchuria rather than those 
from the Yang-tze. 

Whether the Shansi Sika is referable to Milne-Edwards’s Cervus 
mandarinus from the Imperial Hunting-park, Chihli, I cannot 
say. No really good specimens of this deer (from Shansi) have 
been secured. JI have seen summer and winter skins, the former 
being a fine rufous colour, very much like that of the North China 
roe (Cu apreolus bedfordt) in summer, studded with white spots, and 
having a dark median-dorsal line, while the latter is grey, much 
lighter than Milne-Edwards’s figure, with dark median-dorsal 
line. The horns also of this deer attain a large size, and I 
have seen considerably larger specimens than those in Heude’s 
collection. 

‘In dealing with skulls from the Yang-tze I laid out the whole 
series and compared them with each other and with those from 
Manchuria and Tonkin. I could detect no difference that might 
be called specific between the ten species described by Heude and 
represented in the series before me (Nos. 5 to 14 inclusive in the 
list given above). Such differences as did exist could be attri- 
buted to the varying ages of the specimens or to perfectly natural 
individual variation. ‘The variations in length and breadth, such 
as were noticed with the Manchurian skulls, were again notice- 
able here, and as before coincided with the varying ages of the 
animals. 

Of the three male skulls with well-developed horns, labelled 
C. pouvrelianus, C. brachyrhinus, and C. cycloserus, the first had 
the widest spread and the best development, though it fell 
considerably short of the Manchurian and Shansi horns. 

A single specimen of a fully adult male labelled C. andreanus 
from Ning-kuo-fu had well-developed horns, which, however, 
were smaller and lighter than those of any of the three specimens 


ee 


HEUDE'S CHINESE MAMMALS. 19 


just mentioned from Kienté, though they agreed in size with 
those of some males from I iene of Baur the same age. 

C. rivieriauus was represented by a single female skull from 
Tong-leou, which was indistinguishable, except that it was 
younger and therefore slightly shorter, from the skulls of other 
females from Kienté. The skull of a young male with small 
horns labelled C. grilloanus bears a label similar to that upon the 
C. rivierianus skull, and was collected by the same person at 
the same place (‘Tong- lieou), apparently at the same time, while 
the skull of an adult male with horns cut away, labelled 
C’. grilloanus, was collected at Kienté Central. 

Many of the labels on the skulls bear evidence that the specific 
names have been erased and rewritten several times, giving clear 
proof of the doubt and confusion existing in the mind of Heude 
himself as to which of his specimens belonged to which of his 
species. 

Under these circumstances I fail to find any reason for admitting 
the existence of more than one species of Sika in the Yang-tze 
Valley, and therefore I would suggest that, until further and 
more substantial evidence than the. shape of deers’ horns (capal ble, 
as is well known, of almost unlimfted minor variation in a sin aie 
species), which seems to have been the chief characteristic used 
by Heude in his classitication, is forthcoming, the whole of his 
names for the Yang-tze Sikas be suppressed in favour of Swinhoe’s 
much earlier name kopschi*. 

As an example of the extraordinary state of confusion that 
exists amongst Heude’s specimens was a skull of a fine fully adult 
male Sika with good horn-development. On the label of the skull 
the name oaxycephalus has been scratched out and replaced by 
another which is illegible. ‘Yhe word type was also written on 
the label. This skull was not specifically different from any of 
the other Kienté skulls. 


CERVUS DUGENNIANUS Heude ?. 


The two skulls from Phu-lang-thuang, Tonkin, I found to 
closely resemble those from the Yang-tze, but on account of the 
geographical position of the species, and without going into any 
further details, I should consider it valid. 

I could not find any specimens of the following species named 
by Heude :-— 

Cervus cyclorhinus. Shantung. 
»  A&yemalys. North Shantung. 
»  tgnotus. Central China. 
» yuanus. Found in the gravel in caves near 
Tong-lieou, facing An-king-fu. 

Regarding the first two I very much doubt if Sika have 
pecuived in a wild state in Shantung for hundreds of years. 
That province is closely settled and contains no forested regions 

* P. ZS. 1873, p. 574. 


+ Mém. conc, ? Hist. Nat. de ?Emp. Chin. tome ii, p. 156. 
5 6) 


20 MR. A. DE CARLE SOWERBY ON 


suitable to the requirements of these deer. The two specimens 
referred to by Heude must have been kept in captivity in some 
temple, or else had been brought for sale as game from some other 
province. 

Cervus ignotus might be the illegible name on the specimen 
(No. 13 in the list) referred to above, while Cervus ywanus, sup- 
posed by Heude to be an extinct species, may or may not belong 
to the same species as the rest of the Yang-tze Sikas. 

Thus, I propose dividing the Sikas inhabiting China, Man- 
ehuria, and the Primorsk or Maritime Province into the following 
valid species :— 

1. Cervus kopschi Swin. Yang-tze Valley. 
2. » mantchuricus Swin. Ussuri Basin to the 
plains of Western Manchuria. 
3. » dybowskii Tacz. The Primorsk. 
4. » mandarinus M.-Edw. Chibli and North 
China. 
5. 5 dugennanus Heude. Tonkin, 8. China. 

The Sika that inhabits Shansi might temporarily be referred to 
C. mandarinus, while that occurring in Northern Corea might be 
placed with C. mantchuricus till specimens have been secured for 
comparison and description. 


Genus CAPRICORNIS. 


In going over the collection of Serows in the Museum I[ 
examined 26 skulls, with a number of the corresponding skins. 
Altogether I found 24 species as follows :— 


1. Capricornis brachyrhinus. 1 skull. ) 


2. :, chrysochetes. ie eas | 
3. 3 erythropygius. 1 ,, | All collected by Farges at T'chen- 
A, 3 Sargesianus. 2 skulls. r kou-ting in N.E. Ssuchuan. 
5 longicornis. 1 skull. | 
6. S platyrhinus. Ths Ra , 
ie < cornutus. 2 skulls. West Ssuchuan and Moupin. 
8, 3 nasutus. 1 skull. 
9. a ungulosus. par {afoupin, collected by M. Briand. 
10. e microdontus. ie 
11, vidianus. Ines Han-chang-fu, Shensi. 
12, im collasinus. Le 3 Kuang-tung (Canton Province). 
13. ‘3 argyrochetes. | eee Chao-hing-fu (Hsao-shing-fu), 
14, * maxillaris. Lie. Chekiang. 
15. " maritimus. 1 ye oh ator’) 
16. 5 benetianus. 1 cain? 
Wife > bridannellianus. 1 ,, 
18. $ rocherianus. i aed 7 Along Bay, Tonkin. 
1), 33 copusianus. 1 
20. A brossodianus. hens 
ales . achardianus. Lea 3) 
22. 2 marcolinus. tL eae, Kechen, Tonkin. 
23. 5 gendrelianus. 1 ee Keso, Tonkin. 
24. a berthetianus. Ue Keso, Tonkin. 


OC. morianus | was unable to find in the collection. 


HEUDE’S CHINESE MAMMALS. 21 


A look at the map of China will show that the places where 
the above specimens were collected can be arranged in five areas, 
more or less distant and separated from one another, ranging 
from the coastal regions to the border of Thibet. 

These are :— 

(a) Chekiang Province. 

(6) Kuang-tung (Canton) Province. 

(c) Tonkin. 

(Zz) North-eastern Ssuchuan and South Shensi. 
(e) West Ssuchuan and Mcupin. 


Whether the last two are really separable is doubtful, for they 
consist of much the same sort of country, 2. e. high, precipitous, 
and well-forested mountains. 

In examining the skulls, however, I found that they could be 
divided up according to the above divisions. 

I first examined the large series of fine skulls from the Chen- 
kou-ting district of North-eastern Ssuchuan, close to the borders 
of Shensi and Hupei. They were all collected by M. Farges 
during the year 1894. I found it impossible to separate them 
one from the other, the whole series showing certain well-defined 
characteristics. Most noticeable of these was the uniformly great 
length of the horns, which sloped sharply back, were well curved, 
and were closely set, the tips having a tendency to point inward. 
Though the bases of the horns were annulated and there were 
the usual narrow longitudinal grooves, yet these were not very 
pronounced, and the horns had a smooth shiny appearance. 

Turning to the skins corresponding with these skulls, I was 
at first struck by the great variation in colour, but on a closer 
examination I found that this was due entirely to the quantity of 
under-wool present. Thus some pelts were of a grey-buff colour, 
which was due toa great abundance of under-wool, through which 
long coarse black hairs protruded, while other pelts were entirely 
black, having no under-wool at all, the hairs being coarse and 
short. There were all stages between these two extremes. It 
is obvious from this that the varying colours as described by 
Heude, and upon which he bases to a large extent his classifi- 
cation, are due entirely to seasonal changes, mid-winter specimens 
being light and mid-summer specimens dark. 

An examination of the manes, which were pretty well developed 
in all the specimens, showed that in nearly all of them white or 
cream hairs were present, in some greatly predominating over the 
black and chestnut hairs, in others scarcely noticeable on account 
of their scarcity. There seems to be little connection between 
the lightness or otherwise of the mane with the lightness of the 
coat generally, and I came to the conclusion, finally, that the 
white mane is a characteristic of age more than of season. At the 
same time it may be a characteristic which varies in individuals. 
However that may be, it seems pretty certain that the presence 
or absence of the white mane is not a specific characteristic. I 
also noticed that the rusty brown of the legs seemed a scmewhat 


ih) MR, A. DE CARLE SOWERBY ON 


variable characteristic, disappearing almost altogether in the 
winter specimens, and being better developed in some summer 
specimens than in others. 

A comparison of the series of skulls from Moupin including the 
one from W.Ssuchuan, with the series from Chen-kou-ting at 
once revealed a ‘difference, permanent throughout the series (six 
specimens), in the horns. Those of the Moupin series were much 
more deeply ringed and longitudinally grooved, and at the same 
time were shorter and had a greater tendency to spread. The 
longest horn in the Moupin series measured 9/’ as compared with 
103" in the Chen-kou-ting series. The widest spread occurred 
in the two specimens labelled cornutus from Moupin and West 
Ssuchuan. 

In other respects there were no really specific differences 
between the specimens of the two series. 

After my examination of this large series of skulls and skins, 
finding as [ did that the individuals were subject to considerable 
variation in the predominance of the white hairs of the mane over 
the chestnut and black hairs, and also that the rusty colour on 
the legs is more intense in summer and more inclined to grey- 
ness in the winter, I cannot agree with the late Mr. Lydekker’s 
opinion, expressed in his paper (P. Z. S. 1908, p. 942),.that 
there are two species of Serow, one dark-maned and the other 
light-maned, occupying the same localities in West China, but 
would adhere more to the opinion that there are two species, 
both with a tendency to have the white mane, one occupying 
Western and North-western Ssuchuan, the other a more easterly 
region, 


1 & 2. CaPpRICORNIS MILNE-EDWARDSI David and C. VIDIANUS 
Heude. 

To the former I would give David's name of Capricornis milne- 
edwardsi, and to the latter Heude’s name of vidianus, as it 1s 


under phi name that the first description of the N.E. Ssuchuan 
and Shensi Serows occur (tome in. p. 154 of his Mémoires). 


3. CAPRICORNIS ARGYROCHATES Heude. 


The name argyrochetes cannot be given to the Ssuchuan and 
West China Serows, as it belongs to the Chekiang Serow, which is 
certainly distinct from either of the above. The rusty brown 
of the legs, for instance, is distinctly lighter, while the miane is 
creamy brown, due to a greater prevalence of chestnut haus. It 
is interesting to note, however, that here again the presence of 
the light mane is not a permanent character, for a mounted head 
in the collection from Chao-hing (Shao-shing-fu, Chekiang) showed 
no trace of a hght mane. This specimen had horns like the 
Chen-kou-ting ones, but smaller and with a wider spread. The 
name argyrochotes seems to have been given to the Chekiang 
da in 1888 by Heude, the description appearing in tome 11. 
p. 228 of his Mém. cone. |’ Hist. de l’Emp. Chin. 


HEUDE’S CHINESE MAMMALS. yee 


4, CAPRICORNIS COLLASINUS Heude. 


Heude’s species collasinus is represented in the collection by a 
fragment of a skull with one very long horn (9#"') attached, which 
bears a resemblance to those from Chen-kou-ting. Heude had 
the complete skull reconstructed im a drawing which he has 
reproduced on pl. xl. figs. 5-8 of tome iv. of his Mémoires. The 
geographical position of this Serow would almost preclude its 
being placed with the Chen-kou-ting specimens, and, as the horn 
differs markedly from those of the specimens from Tonkin, it 
would seem advisable to let the name remain till material for 
better comparison is forthcoming. ‘The description, if it may be 
called such, appears on p. 211, tome iv. of Mém. etc. 


5. CAPRICORNIS ROCHERIANUS Heude. 


Finally, there remain the large series of skulls from Along Bay 
(Baie de Halong) and Keso in Tonkin to be dealt with. With 
the exception of the specimen labelled C. berthetianus from Keso, 
all the skulls in this series are decidedly smaller than those 
of the Ssuchuan series. They are also proportionately deeper, 
including the berthetianus specimen. The horns of all the skulls 
are smaller and shorter than any of the Ssuchuan or other 
Chinese skulls. Added to these skull and horn differences is the 
fact that the colour of the coat differs markedly from the other 
species, it being black, inclined to blue-black, with a distinctly 
brown face (more so than in the other species) and creamy-white 
legs. Besides this, there is an entire absence of any white or 
cream mane, so that the animal is certainly distinct from the 
other Chinese forms. Nevertheless, there 1s nothing in the series 
to justify Heude’s division of it into ten species. The name 
rocherianus is the first which has a description, and as there seems 
_to be no real difference between the others, it should be applied 
to the whole series. The skull labelled thus is typical of the 
others, and was collected as far back as 1887 by M. Rocher ™*. 
The rest of the names for the Tonkin specimens should be 
considered invalid. 

Thus, Heude’s twenty-four species and David’s one may be 
reduced to the following five :— 


1. Capricornis argyrochetes Heude. The Province of 
Chekiang, 8.E. China. 

oie ss vidianus Heude. ‘The region of N.E. 
Ssuchuan and §. Shensi, Central China. 


Dh ce milne-edwardsi David. W.Ssuchuan, N.W. 
Ssuchuan and 8.W. Kansu, W. China. 

4, ef collasinus Heude. Kuang-tung Province, 
ne Muka. 

by « rocherianus Heude. Tonkin, S.W. China. 


* Mém. conc. Hist. Nat. de ’Emp. Chin. tome ii. p. 225, pl. xxx. 


24 MR. A. DE CARLE SOWERBY ON 


Genus NEMORH EDUS *. 


There are some thirty-two (32) specimens of Goral in the 
collection, representing, according to Heude, seventeen (17) 
species as follows :— 


1. Kemas cinereus. 2 specimens. Moupin. 

2. a OG USCUS: 4 3 “A 

3. »  pinchonianus. specimen. W. Ssuchuan. 

4. »  wanthodeiros. 1 + 6 

5. 4, fargesianus. 6 specimens. Chen-kou-ting. 

6. »  curvicornis. 2 5 on 55 

iG »  initialis. 3 aa a as 

8. >  todinus. 1 specimen. 5 > 

9. » versicolor. 2 specimens. a; 3 
10. 5 niger. 1 specimen. i ig 
ees ae lcanws. 2 specimens. Mountains of Yu-ho, Shensi. 
12. »  vidianus. 2 - ° S. Shensi, 


8 “4s . 
and Si-ngan-fu. 


iby »  tantozatianus. 1 specimen. N.W. Houpei (Hupei). 

14. »  henryanus. 1 5 Yi-chang (I-chang). 

15% »  aldrigeanus. 1 1 oh ” 

16. » arnouxianus. 1 ie Ki-hsien, Chekiang. 

17. 4° raddeanus: 1 55 Oussouri (Ussuri), Manchuria. 


The general areas from which the above specimens were 
obtained are separable in much the same way as those from which 
the Serows were collected, viz, :— 


(a) Chekiang Province, 

(b) Yichang, on the Yang-tze, Central Hupei. 
(c) N.E. Ssuchuan, 8. Shensi, and N,E, Hupei. 
(7) Moupin and W. Ssuchuan. 

(e) Manchuria (Northern and Western), 


Besides these areas, Gorals are also to be found in Chihli and 
North Shansi on the Mongolian frontier, in neither of which nor 
in Manchuria are Serows known to occur. 

It will be seen that by far the largest number (15) of the 
specimens in the collection come from Chen-kou-ting in N.E. 
Ssuchuan. A careful examination showed that they could not be 
separated one from the other—at least, on the evidence of their 
skulls. As regards the coloration, which, as with the Serows, 
seems to have been one of the chief points in Heude’s classi- 
fication, it must be noted that there is considerable seasonal 
variation in the pelts of Gorals. I could not get hold of the skins 
of Heude’s specimens, however, so am not in a position to say 
definitely whether he did or did not fall into the mistake of con- 
sidering seasonal differences as specific ones, but an examination 
of the skulls themselves and also his coloured plates, together 
with my own experience in the field regarding the extreme 
variability of Gorals’ pelts during the seasons, lead me to suspect 
that he did. 


* By Heude this genus was named Aemas. 


"Ne a 


HEUDE’S CHINESE MAMMALS. 25 


1. NEMORHADUS NIGER Heude * 


The skulls of the Chen-kou-ting series in their thin smallish 
horns and general shape and size agreed very closely with the 
specimens labelled griseus, of which there were four in the 
collection from Moupin (N. ‘E. Ssuchuan on the Thibetan border). 

The specimens from Shensi and N.W. Hupei also resembled 
the Chen-kou-ting specimens so closely that I should not hesitate 
to place them together. Whether these and the Chen-kou-ting 
specimens are separable from grisews 1s an open question, though 
I feel inclined to allow it on the grounds of their geographical 
distribution—adopting for them the name of niger as the first one 
of a Goral from this general district having a regular description 
annexed. It is worth noting that the specimens labelled vidianus, 
from Shensi, have unusually long horns, the longest measurement 
being 7'’, I have a single horn from the same district which 
measures 74” in length. The length of this horn is doubtless 
only a local variation, and certainly fails to justify the creation 
of a distinct species. 


29, NEMORHEDUS GRISEUS Milne-Edwards. 


The two specimens from W. Ssuchuan, labelled pinchonianus 
and xanthodeiros respectively, were so like the skulls of grisews as 
to be indistinguishable, and, as they came from practically the 
same district, should, I think, be included in that species. 


3. NEMORH2DUS CINEREUS Milne-Edwards. 


Very different were the two skulls representing cinereus from 
all the other skulls. They were heavier and larger with a wider 
forehead, while the horns were narrower and more closely set 
together. 


4. NEMORHZDUS HENRYANUS Heude ft, 


The two specimens marked henryanus and aldrigeanus from 
Yi-chang closely resembled each other, but differed from the 
Ssuchuan species in being wider across the cheek-bones, with 
wider palate, and distinctly larger eye-sockets. 

It should be noticed that the N.W. Hupei specimen fanto- 
zatianus had a slightly wider palate than the Chen-kou-ting 
specimens, but it was only in this that it resembled those from 
Yi-chang. 

As the first description of the I-chang Goral appears under the 
name of henryanus, that name is the valid one. 


5. NEMORHZDUS ARNOUXIANUS Heude ?%. 


The specimen from Chekiang, arnouxianus, is undoubtedly a 
distinct species, its habitat, which is separated by a wide stretch 


* Mém. conc. Hist. Nat. de Emp. Chin. tome ii. p. 241, pl. xxxviii. fig. 12, 
and pl. xxxix. figs. 2-6. 

ug Mém. cone. ?Hist. Nat. de ?Emp. Chin. tome ii. p. 244, pl. xxxiv. 

t~ Mém. conc. Hist. Nat. de l?Emp. Chin. tome ii. p. 239. 


26 ON HEUDE’S CHINESE MAMMALS. 


of country (apparently uninhabited by Gorals) from the habitats 
of the other species, alone warranting this. A good description is 
given by Heude in tome 11. p. 239. 


6. NEMORHZDUS RADDEANUS Heude®*. 


The Goral from the Ussuri is undoubtedly a valid species. The 
single skull in the collection, though of an immature specimen, was 
slightly larger than any of the other skulls. Specimens which I 
myself have collected in Manchuria (I-mien-po, N. Kiri) show a 
marked difference in colour from others collected by me in Chihli 
and North Shansi, which are undoubtedly referable to Milne- 
Kdwards’s caudatus. ‘The Manchurian specimens are larger and 
greyer than caudatus, and also have a long white tail instead of 
black as in caudatus. The median-dorsal line is broad, dark, and 
well-defined, having a tendency to continue down the face. 


7. NEMoRHADUS CAUDATUS Milne-Edwards. 


That caudatus, of which the Sikawei Museum has no repre- 
sentative, is a good species needs no proof. It inhabits high 
Than regions which are cut off by wide intereouste 
spaces, uninhabited by any form of Goral, from the Shensi, 
Ssuchuan, and Hupei species. 


Thus, it may be claimed that there are the following seven 
valid species of Goral in China and Manchuria :— 


1. Nemorhedus cinereus M.-Edw. Moupin. 


2. i griseus M.-Edw. Moupin and W. 
Ssuchuan. 

3. - niger Heude. N.E. Ssuchaun, 8. Shensi, 

: and N.W. Hupei. 

Ay eager a henryanus Heude. I-chang, Central 
Hupei. 

D. as arnouxianus Heude. Chekiang. 

6. ne caudatus M.-Edw. N. Shansi and 
Chihli. 

kg vaddeanus Heude. Manchuria. 


%* Mém. conc. l’Hist. Nat. de Emp. Chin. tome 1. p. 240, pl. xxxv. 


ON DEATHS IN THE SOCIETY’S GARDENS, DiTf 


3. Report on the Deaths which occurred in the Zoological 
Gardens during 1916, together with a List of the Blood- 
Parasites found during the Year. By H..G. PLimmer, 
F.R.S., F.Z.8., Professor of Comparative Pathology 
in the Imperial College of Science and Technology, 
London, and Pathologist to the Society. 

Received and Read February 6, 1917. | 


INDEX. Page 
eit LOC We eter ecceastithes ada nnls 9 Gtoelute dawieasor eee, 27 
WSOC EN eATOSIECS#: ceo ccc kN Goes iebos eens eee 3] 
INIOMOSa Meee ene ete sateen Cen Ramat 33 


On January Ist, 1916, there were 690 mammals, 1989 birds, 

and 426 reptiles 1 ae Zoological Gardens: during the year 
1916, 165 mammals, 545 birds, and 199 reptiles were admitted, 

pints A total for the year of 855 mammals, 2534 birds, and 
625 reptiles. 

During 1916, 242 mammals, 594 birds, and 179 reptiles have 
died: that is, a percentage of 28: 3. 108 mammals: 23°4 for birds, 
and 28°6 for reptiles: the average percentages for the last five 
years being 27, 23°3, 31°2 respectively. 

Out of the total deaths for the year, 1019 in all, 269 occurred 
in animals which had been less than six months in the Gardens,. 
that is a little more than a fourth of the deaths. It has been 
found that after six months’ residence in the Gardens the death- 
vate of the animals falls rapidly ; so it is assumed that by this 
time the new arrivals have got used to their new environment, 
or have died from any diseases of a parasitic kind which they 
may have brought with them. Of these 269 animals 60 were 
mammals, 132 were birds, and 77 were reptiles; and, if these 
be deducted from their respective totals, the death-rate per- 
centage will come out as 21:2 for mainmals, 18-1 for birds, and 
16-3 for reptiles, the average percentages foe the last five years 
being 17-3, 15°6, 13°8 respectively. 

The fol Wane ing Tables show in outline the facts which have been 
ascertained. Table IT. summarizes the actual causes of death in the 
three groups specified. Amphibia are included under Reptiles. 


TaBLeE [.—Analysis of the Causes of Death. 


| Reference 
Diseases. /Mammals.| Birds. Reptiles.) to Notes 
| following. 


| 1. Mierobic or Parasitic | | 


Diseases. | | i 
PIETER CUNOSIS” 808 acute ces Sos ates 16 (| 1 2 
MMM OG STS ches. cess Ses cee evet| 12 29 «| 6 3 
PMEMUMOMIA, 6 o5 cia vice) sa vselansses| 30 538 | 48 4 
OD UICSMINTA oo cere cea ese casies 5 awn 1 5) 

JANOS OLEISISio vat hg OR a 2 ae 1 
Syrah ode) 00s | jean ea en eee Ce 2 Aa 
Peritonivises aes 2 1 


28 PROF, H. G. PLIMMER ON 


TABLE [.—Analysis of the Causes of Death (continwed). 


Reference 
| Diseases. Mammals.| Birds. | Reptiles.) to Notes 
| following. 
1. Microbie or Parasitic 
| Diseases (cont.). | | 
Cystitis eee: Bono aap 1 ee a 1 
Leucocytozoa infection... | oe Ly: Alanis 
Acavi infection 01M. pend 3 i a 6 
Syngamus infection ...............) ae a er 
Hilariasis: Naa et Wis. te asec 1 ae aS 
Aiscaridosisesa...cccctsaa) caer 2 ee toe) 1 o 
2. Diseases of Respiratory | 
Organs. | 
AMtelectasisne sw eect ene 5 ) 
Bronchitisescncdspcsc.a ct oe 8 | 
| Broncho- -_pheumonia . AOR et a a | 42 eae fee he Maat 7 8 
| Roveestion of STnER iy eek 3 hy | 8 | 
| Pleuritis 2228 03. Re enter iL oe |) ese 
3. Diseases of t the Heart. | 
Degeneration of muscle ......... 1 Ie 
| 4. Diseases of the Liver. | 
| Hepatitis RAPP e inet des 558 Lm as) 2 
| Cirrhosis ccc -k he eonte betes ae 4 2 ah) ses 
| Fatty degeneration Ae ss | 3 | 1 
| 5. Diseases of the Mee) | ne | 
Tract. | 
Gastro-enteritis) uscjietce ..2 4 2 2 
Gastric ulceration cue. 2 ee se | 
ji) Ainteritis Weide see ena. cee. | 22 190 14 9 | 
| Gastritis .. : ees, one af 3 
| Intestinal obstruction ............ pe | 1 vee | 
Intussusception Peet 2 vee 
Prolapse of rectum <...45-41.....- | it | 
| 
| 6. Diseases of Urinary and | 
Generative Organs. | 
Nephritis, ceoica) <. ce eee | 34 45 5 10 
Cystic kidneys 3.5...5..c0 eee e 1 25 
| Salip ng iGisc.ke.cee sc tes eee: | “i 1 3 | 
| Extroversion of uterus. ......... | 1 _ nae | 
| 7. Various. | 
| Caremomiais: sa.cecaie umes 3 es bas 11 | 
SAT COMA hu. tec eee 2 1 12 
Tbeukheemual “eee eee 1 13 
| Caries of spine ... otnee | 1 
| Spinalkcuryature: j97.005. 9a 1 | 
Hamorrhage...00.e oy eee | a 1 as | 
| Injuries discovered aa os 1 - 14 | 
MOTCOME, canna ieee ete Neer 


} cas. = 
Besides those tabulated above, 
25 mammals, 85 birds, 7 reptiles, were killed by order or 
by companions, 
1 bird, 37 reptiles, died from malnutrition 
or starvation, 


6 mammals, 22 birds, 38 reptiles were too decomposed 
for examination, 


1 bird was preserved unopened, 
completing the total. 


DEATHS IN THE SOCIELY’S GARDENS. 29 


In Table I. an analysis is made of the immediate causes of 
death, but in nearly every case the animals were found to be 
suffering from other lesions as well. Table IL. summarizes these 
other diseases from which the animals were suffering ; and if 
this Table be taken in conjunction with Table I., a much more 
accurate estimate of the amount of disease in the Gardens will be 
arrived at. 


Taste IT.—Secondary Diseases found in the animals tabulated 
in Table I. 


: | Reference 
| Diseases. Mammals. | Birds. Reptiles.| to Notes 
| | | following. 
MINE COMP (alk sec abies ae siciesruandeoes 1 vee | wes 
ALU COSIS aoc a eRe re ae ee ee ee oan) 2 
| PPE UTI 65555 an dacs s cencascie sense: 2 6 
MBC GICADUULIS: 82a: cheeses s-ntsanees cen dnge 1 aes 
WRBRCEICOMIGIS occ veces ove cacmestensedese 5 Z| 3 
DAMP SC OSH MT he oe Ng siocs\s iba S wedi wae des 2 }; 
Je. HPPLN0 SE a ee ee 33 | | 
JEUNE UIGISS al Sen eet ae eee eee ea ih ree ewsee | 
NVOrMA WMPECEION. ........00c00000 oeeeuiens 1 3 3 
PMMA tie cae cog ee sltwns ns vinaan stn Suviess a 4 | | 
|, UNUCEIEN ietiag Seen ee 1 Dee ASP aye 15 | 
F HBMOPTEPATINES 2.262 .....c cece ene an on Ve 4 
WINEMGOCYLOZOM: 0. 0:58: 0cae cosa iene cveass - Ls Ae 3 
CHORTLE ee ne ee ee seh, tl 3 16 
= — | | 
PSEOMOMMIS Wh 2 0. pdt sateen tsa niesnds 6 eee 
Broncho-pneumonia .................. 11 tee aN as 
Congestion of lungs. .................. 20 162"). 16 
CHGema-OF TUNES 1.. 5.6 .cccseceacec set ae i 103 | 8 
@ollapsed LUNES) ii... ccs tee exes cee ees 1 on ee 
Degeneration of heart. ............... 4, Ls i 
Hypertrophy 0 of heart 1 
Hepatitis .. i Pee cath sien ah es Lea eee 
PeriCArOitiS cos cc-s-.--sscccsceseess ses eat Se Rely Back Ly 
MGweroMma Of AOLbA ....6..6. 1. ces cee 3 4 
PAMMNENUENS Mss sc ie ciate cc's vtedioe et » bc 1 ae 18 
Fatty pee yo Of lIVer ccs... ls 41. 5 
Cirrhosis of liver.. 3 | 1 
Cholecystitis 1 | 
CASING Rent tone Grae. shecsinieks 3 1 
Gastric ulceration ........ teat nade 9 “te 
GaSELO-EMbCEILIS’ ©..25.. can scheeeve aces 5 4 
Enteritis. ...... ee SAN ae Kt Sh 28 90 14 
| Intestinal obstruction ............... se il 
IN QR NTT TE) a Nr 36 78 a 4 
MC ystic tidy S..< 0... ..aceeee--.s0h0-- 00 ul 2 
PSL NIT Nae oe a 3 2 ons 
PS CINES Pst c6 os asciec ob sale w Saheb aon as Hey a | 
Angioma, ........ 1 
| Lardaceous disease .. 1 
MCtAIMEME LOBES! fears. does sesee- 1 
Rickets 2 
| Urzemia 1 7m 19 
_ Injuries il 5 


30 PROF. H. G. PLIMMER ON 


Table IIT. shows, in still further detail, the distribution of 
the immediate causes of death, shown in Table I. , amongst the 
varlous large orders of mammals, 


Tasie Ili.—The Distribution of Diseases causing Death 
amongst the principal Orders of Mammals. 


a s ; Ben be Zs 
Diseases. | = S | = | = ees 2 
| (= oA | D | =) = e 
bce Bol 1 gs) es 
| yb Oo | a | eae 
| of a pe al eee 
Mulberculosisy ove sox ese tate eee naa ee iL 2 2 
MyCOSIS. sited. Sate Sait Winger. aeeneeet el aged al 6 4, 
Pnennvoniae. 2) cs asso in neat erten cs hele te AN San) 1 2 
[@eptl Cooma. te tees. 4! ear eee eye | > cena eee SRL Re 1 
IMDS CESS Sati: S50. Fe cmaeces ante eee. ane Sash pal: 
Empyema apes 2 | 
Peritonitis [eas SS 1 | 
Cystitis Peta aR ee eet ete ee | eee AL : | 
HATIASISi eS eetont. 2. een ae tae cere i; Sean ame 1 | 
(AS CATICOSIS Wen ence ete atte smeee | 2 | 
| | % | ~ 
Abeleehasiay, #0 cen eee tant cee Lies a a 1 
ie LOD CRIUIS Metis aten em te woe 2 2 3 eae 
| Broneho- pneumonia Er ee eet eos). tied = Ok Ot | aes 4 
| Congestion of lungs wad tes aasless tau sone e| oes es 2 iL alee 
Pleuritis cee Be 3 Ge eee Peer roe ae ‘dee el 
Degeneration of heart peer cee so = Uy | 
t'Cirshe@sisoF livers ais. etree: cee |e sce ee Secale aaa 2 
‘Gastro-emtermbis: et. cee 3 Cae eer | | 
Gastriciulcetatione.-41 + se eee lan Sa] ae d) ilweate 12 Pe 
Hin teritisran: ..e.cke = eee ee ea 8 2 | 19 | 923) See 
i Intussusception ...7--5.. else ee eee: | 2 ee i ecaecmeal 
i: Prolapsexol rect... eee ae | ook pat yr cae ee eee 1 
| Nephritis... : Mercere sot epenet fame be-fam lK €) 44 § 2 1 
| Extroversion of uterus .. eas = /uc<) [roan aon gece | 
| Carcinoma Pf Se | 
| Sarcoma 5A Sere Cihk sae : ee ee 
Tsewkshvemiia 4s 89 2 re eal nse 1 | 
Cariesvot: Spimennvs- masse |) peal | | 
Curvature’ ofspine 2)... 2.0: ‘i | 
\ 


The following, Table IV., shows the number of Deaths from 
those Diseases of the greatest numerical importance for the last 
four years: the total number of animals of each class is placed 
above, so that a proper comparison can be made. 


a at —, 


| 
‘ 


| 


| 
| 
| 
| 


| 


| 
) 


DEATHS IN THE SOCIETY 8 GARDENS. 3 
TABLE IV. 


Mammals. 


Meat wee con ess 19138. 1914. 1915. 1916. 


Total number of mammals ... 1272 1261 1613 855 
Tuberculosis 31 | 12 6 | 16 
_ Mycosis Sine MR ee carp eer See 8 | 6 | 10 12 
PAM CUMMOMIG: ... ..0cee cc csc eh neces von cee 34 | 53 48 30 | 
PMPMO MULE Se). cd oi discs ce stone ede oes 6 Tet 10 8 | 
Broncho-pneumonia ............-..--- 25 24 34 42 | 
Congestion of lungs 14 | 14, 13 3 | 
_ Gastro-enteritis ... Dern eee 7 | 16 Ay 4, 
MOMENI RC 5. ee Cecio e sdlivse ceca cahe oe 33 | 33 25 29 
IN [DUDES GO Qe ean a a 90 | 66 40 34 
Birds. 

Total number of birds ......... 3518 3610 3240 2534 
MMVETOMIOSIS ? 2. co.cc. cde dee ccs devceeeee 104 iGis) 60 vl 
ISIN OPUS EMR AG oe cers ccilceae cuccses 1s 75 | 88 | 32 29 
JEST ACY iii er 89 | 118 | 70 53 

mwoneestiom OF lungs .i.....60c0ce-c-0s 98 133 130 78 

| SUS WG. WOES er 148 | 169 150 190 

JIN| ) OCU ETS I ae te a 135 129 81 | 45 
Reptiles. 

Total number of reptiles ...... 1169 1045 608 625 
Unsere mVOsts) coos. ele se asads oan ce 6 A | 2 1 
LUO SICE et eevee ae | 1 10 | 6 6 
PC eMMNONIA ee eee oe. Cota esse | 138 69 | 3d 48 
Congestion of WINES. ....5.... 205.0000 13 | 19 11 | 8 
LE STO IEY ll ee a ae ee re a 15 | 17 8 | 14 | 


BLoop-PARASITES, 


During the year the blood of every animal mentioned in the 


foregoing lists has been examined, with the result that parasites 
have been found in 24 cases, in 13 species for the first time. 


They have been distributed as follows :— 


Hilarie. In 3 mammals; in all for the first time. 
In 3 birds; in all for the first time. 

( Hemoproteus danilewskyt. In 5 birds; in 3 species 
iota: Z for the first time. 

\ Plasmodium precox. In 1 bird for the first 

time. 
Leucocytozoa. In 3 birds; in 2 species for the first time. 
Hemogregarines. In 9 reptiles; in 3 species for the first 
time. 

Intestinal organisms, In 1 reptile. 


ou PROF. H. G. PLIMMER ON 


The particulars of the blood-parasites are as follows :— 


Filarice. 
Mammals. 

Found for the first time in: HaBItTat, TYPE. 
Bay Lynx (Felis rufa) ........2......4...... Mexico, Long. 
Hamster (Cricetulus obscurus) ............ Mongolia. Long, thick. 
Grison (Girison allamandi).................. Argentina. Long. 

Birds. 


Scarlet Cardinal (Cardinalis pheniceus). Venezuela. 


Found for the first time in: 


Black-headed Sugar-Bird (Chlorophanes Brazil. Short, thick. 
spiza). 
Gay’s Finch (Phrygilus gayi) ............. Chili. Long, no capsule. 
Malaria. 
birds. 


Heemoproteus danilewskyi. 


Paradise Whydah Bird (Vidua para- W. Africa. 
disea). 


Shama (Cittocincla macrura) ...........: India. 


Found for the first time in: 


Rose Finch (Propasser rhodochrous) .... India. 
Yellow-backed Lory (Lorius jflavo- Batchiam. 
_ palliatus). 

Scarlet Cardinal (Cardinalis phaniceus). Venezuela. 


Plasmodium preecox. 
Found for the first time in: 


Pitta (Pitta nove-quine@).................. Aru Islands, 
& 
Leucocytozoa. 
Birds. 
Scops Owl (Scops git) ...............0....... Captured at sea. 


Found for the first time in : 


Rose Finch (Propasser rhodoehrous) ... India. 


Carrion Crow (Coreus ecorone) ............_Kurope. 
Haemogregaries. 
Reptiles. 
Common Boa (Boa constrictor)............. S. America, 
King Snake (Coronella getula) ............. N. America. 
Banded-tailed Tree-Suake (Leptophis Europe. 
viperinus). 


Indian Python (Python molurus) 
Eyed Lizard (Lacerta ocellata) ............ Europe. 


DEATHS IN THE SOCIETY'S GARDENS. 3a 


Found for the first time in: iS iy Gena ic 
Annulated Snake (Leptodira annulata). S. America. + Stout. 
2 Burrowing Boas (Hryx thebaicus) ...... Gold Coast. Ordinary size. 
Ocellated Bladder-frog (Leptodactylus Argentina. Long, thick. 
ocellatus). 


Intestinal Organisms. 
Leptiles. 


Viperine Snake (Tropidonotus viperinus). Europe. Amoeba. 


NOTES ON THE FOREGOING. 


1. The total incidence of microbic and parasitic diseases causing 
death in the Gardens for 1916 is 8°5 per cent. in mamunals, 
6°] per cent. in birds, and 9:4 per cent. in reptiles. If those 
cases which have not lived six months in the Gardens be ex- 
cluded, these percentages will be greatly reduced. 

2. The numbers of deaths from tuberculosis amongst the 
mammals and birds show a relative increase, the incidence being 
1-8 per cent. in mammals and 2°8 per cent. in birds. Amongst 
the mammals dying from tuberculosis, 16 in all, 7 were Monkeys, 
and of these 5 had been pet animals; of the remaining 9 animals 
4 had been pets. ‘These figures seem to me to indicate quite 
plainly that pet animals should not be accepted by the Society, 
or only after the strictest quarantine. An Orang Utan which 
had been 84 years in the Gardens had chronic tubercle of 
human type, which he might very well have brought with him. 
The most remarkable case this year was in an Ibex which died 
when about a fortnight old with tubercle of liver, spleen, and 
mesenteric glands. There is a relative increase this year in the 
number of deaths from tubercle amongst the birds, and in a 
much larger relative number were the lesions generalized, namely 
in 36, indicating a more severe type of the disease. The one 
reptile was an Alligator in which the disease was of human 
type, and was no doubt caused by the habit of certain visitors of 
spitting at the animals, to which I have often called attention in 
previous reports. 

3. As usual I have grouped all the diseases caused by moulds 
under mycosis. In the mammals six of the cases were of the 
ordinary type, associated with abscesses; and in six, tumours 
(mycetomata) in various organs were present. Ina sheep the 
growth began in the cavities of the nose and spread into the 
adjacent bone-spaces. There is a slight relative increase amongst 
the birds, but there are now less than half the number of cases 
there were a few years ago. In one Parrot it was caused by 
Aspergillus nger. Three of the six reptiles died with mycotic 
tumours in various parts. 

4. The incidence of pneumonia remains about the same as 
last year amongst the mammals and birds. In one bird it was 


Proc. Zoot. Soc.—1917, No. ITI. 3 


34 PROF, H. G. PLIMMER ON 


due to trematodes and in three reptiles to worms, otherwise it 
was pneumococcal. . There has been a slight increase amongst 
the reptiles. 

5. The septicemias were due to abscesses in two cases, to 
extensive ulceration in two, and in one to a decomposing feetus. 

6. This rare infection of the lungs and air-sacs of a Pheasant 
from Mongoha has been noticed only once before in the Gardens, 
in» Hangnest from La Plata. The mite was a Cytoleichus. 

7. In these cases of nematode worm infection the lesions pro- 
duced were very different; in a Lynx they caused a pseudo- 
tubercle of the liver, ina Puma they caused perforation of the 
stomach-wall and peritonitis, and in a Toad they produced a 
pneumonia. 

8. There has been relatively a small increase in the diseases 
of the respiratory organs amongst the mammals. They are, of 
course, largely Acpended on Poa and 26 of the cases 
of broncho-pneumonia occurred in the last two months of the 
year, during bad external conditicns. Among the birds there 
has been a considerable relative decrease in the deaths from 
congestion of the lungs. 

9. There has been a considerable increase in the number of 
cases of enteritis amongst the birds, but it has been of a milder 
type. 54 of the bird cases were hemorrhagic; in 4 worms were 
the cause, and in 3 foreign bodies, and 2 were due to ameebe ; 
during the last three months of the year 28 Parrots have died 
from that specific kind of enteritis called Psittacosis. Of the 
cases in mammals, 3 have been hemorrhagic, 4 associated with 
ulceration of the intestines, and 1 was caused by foreign bodies. 
The term enteritis covers all inflammatory diseases of “the intes- 
tines, and as these are due to so many different causes it may be 
worth while to mention them. In the Gardens I have found 
five different varieties, which are due to the following causes: 
1. Errors in feeding. 2. Foreign bodies. 3. Worms. 4. Bac- 
teria, 5. Protozoal organisms. Of these causes the first and the 
fourth are the most important. i 

10. There has been a considerable relative decrease in the 
number of cases of nephritis. 'This word, hike enteritis, covers 
inflammations of the kidney which are due to various causes. 
Nephritis can be divided, for purposes of classification, into acute 
and chronic; the former being due to infection, or to exposure, 
and the latter either following the acute disease, or being due to 
degenerative changes and associated with car dio-vascular changes, 
with old- -age eliamece or the artificial old-age changes induced by 
captivity. The absence of proper space for exercise must render 
animals more sensitive to temperature changes, and especially to 
draughts. 16 of the mammals had acute nephritis, and 18 
chronic; only 1 bird had acute nephritis, all the others being 
chronic ; in all the 5 reptiles it was: chronic. 

11. The three cases of cancer occurred in the liver, spleen, and 
adjacent glands of an old Bear, in the liver and adjacent organs 


~ dee 


DEATHS IN THE SOCIETY’S GARDENS, 35 


and glands of a Cavy, and in the neck of a Wolf. The case of 
the Wolf is of interest because his father died here in 1914, and 
his mother in 1915, of the same disease in the same situation. 
A sister is alive and well. 

12. A Coypu died from an angiosarcoma of the liver; it had 
also an adenoma of the kidneys, and is of interest as being the 
third case of similar tumour of the liver, and the fifth of similar 
tumour of the kidneys in Coypus dying in the Gardens during 
the last 9 years. 

13. Of lymphatic variety in a Polecat. This is the second 
case of this rare disease which has occurred in 9 years. 

14. The injury was a ruptured heart in a Flamingo, and is 
mentioned because it is the fifth Flamingo which has met with 
the same rare accident. 

15. Further details of these blood-parasites will be found under 
the section ‘ Blood-Parasites,” p. 32. Under the term malaria 
are grouped cases due to Hemoproteus danilewskyt and to 
Plasmodium preecox. 

16. ‘These were due to a large Saccharomyces. 

17. These cases in birds were due to the deposition of masses 
of crystals (of guanin, probably) in both visceral and parietat 
pericardium. 

18. The only aneurism found this year was in a Civet, of 
fusiform variety. 

19. This rare complication of kidney disease occurred in a 
Mandrill which had acute upon old nephritis. 


2 
‘ 
* 
i 
= 4 
4 
4 
— 
t ew 
\ 
i o 
of 
os 


THE MOUTH-PARTS OF THE PALAMONID PRAWNS. 37 


4. On the Structure and Function of the Mouth-parts of 
the Paleemonid Prawns. By L. A. Borrapatze, M.A., 
F.Z.8., Lecturer in Zoology in the University of Cam- 
bridge, Fellow, Dean, and Lecturer of Selwyn College. 


[Received September 19, 1916: Read February 6, 1917 | 
(Text-figures 1-51.) 


INDEX. Page 
Mint ROG ilOMy eee maeen cla. Cet tke detest ete ueteeviden LO: 
Morphology of the crustacean limb ................... 37 
Morphology of the mouth-parts of Palemonide ...... 62 
Physiology of the mouth-parts of Paleemonide ....... 69 
ile 


In the course of an investigation of the commensal prawns of 
the subfamily Pontoniine, it became necessary for me to compare 
their mouth-parts with those of the free-living Palamoniine, in 
order to discover whether there existed between these groups 
any difference, in the organs in question, which might correspond 
with the difference in the diet of the animals. For the most part 
I have taken as representative of the Palemoninze the Common 
Prawn (Leander serratus), in which I have studied in some 
detail the structure and arrangement of the mouth-parts, and 
endeavoured, by the observation of living specimens, to find out 
how the organs in question are used. Surprisingly little trace 
has appeared of such structural differences between the sub- 
families as I was looking for, but my observations have suggested, 
with regard to the morphology and functions of the parts around 
the mouth of Palemonide, certain reflections and conclusions 
which form the subject of this communication. 


II. 


1. The morphology of the jaws of Malacostraca is still in a 
good deal of confusion, and there is much disagreement as to the 
relation of the parts of each of them to those of other crustacean 
limbs, and even as to the names to be applied to certain of their 
processes or “lacinize ”*, A great part of this difficulty arises 
from the fact that no theory as to the primary form of limb of 
which all the appendages of Crustacea are modified representatives 
has met with general acceptance. This thorny question is not 


* T have not, in this summary article, made any reference to the literature of the 
subject. The contributions ot Beecher, Boas, Claus, Coutiére, Hansen, Huxley, 
Lankester, Packard, Thiele, and others to our knowledge of homology of the limbs 
of Crustacea are well known to those who are interested in the subject. The reader 
will readily gather in what respects my views differ from or agree with those 
expressed by each of them, and it is needless to emphasize the fact that all present 
speculation must be based upon the foundation they have laid. 


38 MR. L. A. BORRADAILE ON THE 


only raised by the jaws of Malacostraca, but receives from the 
study of some of them—notably from that of the maxilla— 
considerable elucidation. The following, briefly stated, are the 
considerations in regard to it which have influenced the 
morphological suggestions put forward in the present paper. 

2. It is not to be assumed without question that a prototype 
can be found from which all the limbs of all Crustacea can be 
derived by modification. The mere fact that appendages stand 
in the same position on the bodies of two or more segmented 
animals, or are members of the same meristic series In one 
animal, affords no ground for the assumption that there exists 
a common plan which underlies the arrangement of the parts of 
each of them. Moreover, even when there are resemblances be- 
tween them, such a conception as that of a common type of them 
is of no use to the zoologist unless the plan of the type does 
not merely exist in the imagination of the observer, but has or at 
some time had an objective existence as a factor in development. 
Resemblances between two limbs may be due either solely to ex- 
trinsic causes—that is, to influences from without the organisms, 
which have brought it about that different developmental pro- 
cesses result in similar structures in the two cases,—or also to 
intrinsic causes—that is, to the operation of identical develop- 
mental factors. Only in the latter case has a common plan for 
the two limbs an objective existence. Conceivably such a plan 
may not be due to community of ancestry in all instances. When 
two appendages closely resemble one another—as do, for instance, 
the various antenniform limbs of Arthropoda—it is possible, 
and sometimes probable, that there exist in the mechanism of 
development facilities for establishing such organs, and that 
these have come into play independently more than once, forming 
appendages upon the same plan. This principle, however, is of 
limited application. Even in the cases which suggest it, it can 
never be applied save to actual features which exist in all the 
limbs under comparison. ‘To assume the existence of a plan of 
which some features are not realized in each limb is quite unjusti- 
fied, except on the hypothesis of the common inheritance both of a 
type of limb and, in the case of serial resemblance, of community 
of type between the members of the series of limbs*. To look 
for traces of a common type in structures in which it is so little 
obvious as in the appendages which appear to correspond in 
different Crustacea would be absurd, unless there were grounds 
for believing that their possessors were descended from a common 
ancestor; and if we are also to find community of type between 
the several limbs of each individual crustacean, then such com- 
munity must have existed among the appendages of the ancestor. 


* The resemblances here classed as due to extrinsic causes are those which are 
generally classed under the head of Analogy. Those which are due to common 
descent are instances of Homology. Those which are due to independent operations 
of the same developmental facilities (if such there be) form a third class not strictly 
the same as either Analogy or Homology. 


MOUTH-PARTS OF THE PALA MONID PRAWNS. 39 


Yor if (as might well be, by bomeosis) it were independently 
established in any members of the subphylum, then the altered 
limbs would lose what they had in common with the corresponding 
limbs of other Crustacea. 

3. In point of fact, however, there can be no doubt that the 
Crustacea are monophyletic, and it is very highly probable that 
their ancestor possessed a complete series of similar limbs. ‘The 
widest gaps in the crustacean system are those which separate 
the Copepoda and the Cirrivedia from the rest of the subphylum, 
but it is quite impossible even in these cases to entertain the 
suggestion of an independent origin. The occurrence of the 

nauplius would by itself negative this. That in the common 
ancestor of the group all the limbs, with the possible exception 
of the antennules, conformed to one type may be gathered with 
some confidence from the conditions in Branchiopodaand Trilobita. 
The Branchiopoda are certainly the most primitive of existing 
Crustacea. They alone possess in a simple form—allowance being 
made for certain obvious specializations, such as the degeneration 
of the mouth-parts—all the elements of the organization of every 
other class of the subphylum, and their suggestive resemblances 
to the Annelida are the only indications of affinity with other 
phyla shown by any recent members of the group. These facts 
fairly entitle them to be regarded as indicating broadly the 
ancestral features of the Crustacea, It is needless to dwell upou 
the fact that their trunk-limbs are all of one type, and, although 
that type becomes unrecognizable in their maxille and maxillules 
its presence in these limbs of other groups (Ostracoda, Malaco- 
straca, and Copepoda) supplies the lacking evidence here. In the 
Trilobita, which are without doubt related to the forbears of the 
Branchiopoda, a single type of limb extends throughout the 
body, from the antenne backwards. It is not possible to avoid 
attributing to the ancestor of the Crustacea that similarity of 
appendages which characterizes these groups. This, of course, 
is not to say that the limbs of the first crustacean resembled 
closely either those of the Trilobita or those of the Branchiopoda. 
The form of the primitive crustacean appendage can only he 
conjectured with probability after comparison of all those which 

may be assumed to be derived from it. But, whatever it may 
have been, the foregoing considerations justify the belief that it 
existed, a that from it can be derived all the post-antennular 
limbs of every member of the subphylum. 

4, The primitive limb of the Crustacea must have been of the 
kind to which the terms “leaf-like” and “ phyllopod” are 
applied—that is, flattened, lobed, and feebly, if at all, jointed 
(text-fig. 2). This is a priore likely in view of the structure of 
the parapodia of Annelida, and seems established by the way in 
which the phyllopod limb is distributed among the Crustacea, 
occurring as it does throughout the trunk of Branchiopoda 
(text-figs. 3-5), on the ible and presumably less mature, 
segments of Z'riarthrus (text-fig. 6), in Leptostraca (on the thorax, 


AQ MR. L. A. BORRADAILE ON THE 


text-fig. 8), and in the hinder part of the head of all groups. 
Whether it was biramous, like most parapodia, the limbs of the 
Trilobita *, and many of those of the higher Crustacea, is another 
question. ‘In the majority of biramous crustacean limbs the 


Text-figure |. Text-figure 2, 
a 

I\E p 
a 3 
= ZB 
Zac ee 


a 


V.C. Cp. 
PREP) nom 
Parapodium of Hunice maxima, Hypothetical limb of 
atter Ehlers. proto-phyllopod. 


For lettering see p. 71. 


Text-figure 3. 


Tenth thoracic limb of Apus sp. 


For lettering see p. 71. 


protopodite and endopodite form an axis to which the exopodite 
is appended, while the trunk-limb of the Branchiopoda is so 
clearly uniramous that there is discussion as to which of its 


* But see footnote to p. 48. 


MOUTH-PARTS OF THE PALZMONID PRAWNS. 41 


Text-figure 4. Text-figure 5, 
7 (cap) 


Wil 4 ' ee 


Yen” Ze 

=e. 
GC a 

2 \\ = 2a, S 
ee 2 
ig a 4 
Thoracic limb of Chiro- First thoracic limb of Limnadia 
cephalus sp. lenticularis, after Sars. 


For lettering see p. 71. 


Text-figure 6. Text-figure 7. 


Anterior pygidial limb of One of the anterior 
Triarthrus becki, drawn thoracic limbs of the 
from a model by Beecher. same, 


For lettering see p. 71. 


42 MR. L. A. BORRADAILE ON THE 
processes represents the exopodite. Probably the primitive 
crustacean appendage (text-fig. 2) resembled that of the Branchio- 


poda in being uniramous. On its outer side this limb must have 


Text-figure 8. Text-figure 9. 


Theracie limb of Nebalia sp. Outline of axis of thoracic limb of 
old female of the same. 
For lettering see p. 71. 


Text-figure 10. 


Thoracic limb of Paranebalia longipes, after Sars, joints added 
after Thiele. 


For lettering see p. 71. 


borne near its base several epipodites. There is no necessity to 
suppose that these were present to the full number of the gills 


MOUTH-PARTS OF THE PALZAMONID PRAWNS. 


43 


Text-figure 11. 


and other epipoditic structures that may be found on a single 
segment in various Malacostraea *. 


It is highly probable that 


Text-figure 12. 


Maxilliped of Calanus sp. 
For lettering see p. 71. 


Maxilliped of Anaspides, after Calman. 
Text-figure 13. 


For lettering see p. 71. 


Text-figure 14. 


\ > 


Maxilla of Calanus sp. 


Maxilla of Nebalia sp. 
For lettering see p. 71. 


either branching and subsequent division, or meristic repetition 
has taken place here. 


But it is not unlikely that the presence 
* See Ann. Mag. Nat. Hist. ser. 7, vol. xix. p. 462 (1907). 


44 MR. L. A. BORRADAILE ON THE 


of three epipodites (the epipodite proper and two “ pro-epipo- 
dites”), as, for instance, in Chirocephalus (text-fig. 4), is a primitive 
feature. Beyond the epipodite stood a longer process, the 
flabellum, which may have been slender, like those of Concho- 
straca (text-fig. 5) and the exopodite of Paranebalia (text-fig. 10), 
but is perhaps more likely to have been broad, like that of Apus 


Text-figure 15, Text-figure 16. 
*) gee 
2 ye 
LZ ¢ 
Shy 
J 7 by lez 
eX Y2 
ap is 
AB 
—3 
== 
Text-figure 17. Text-figure 18. 
Text-fig. 15. Maxilla of Cerataspis, after Boas. 
f 16. Larval maxilla of Peneus, after Claus. 
i 17. Maxilla of Anisocaris, after Ortmann. 
3 18. Larval maxilla of Palemonetes, after Boas. 


For lettering see p. 71. 


' (text-fig. 3) and the exopodites of most Leptostraca (text-fig. 8) 
and of the decapod maxilla (text-figs. 15-18, and 47). It pro- 
bably extended both distally and proximally from its attachment, 
like the conchostracan and notostracan flabella and the maxillary 
expodite of Decapoda. The axis of the limb ended in an apical 
lobe, and on the inner-edge stood a row of endites. It does not 


MOUTH-PARTS OF THE PALEZMONID PRAWNS. 45 


follow, from the fact that these, with the member resembling 
them which probably represents the apical lobe, number six in 
many Branchiopoda, that the primitive limb possessed only that 
quota. In certain cases the series is more numerous. ‘Thus in 
Anostraca it has probably seven members *, the maxille of certain 
decapod larve (text-figs. 16, 17) certainly possess eight, and the 
same limb of Cerataspis (text fig. 15) bears nine. If, as seems 
likely, the first member is missing in the maxilla of Calanus (text- 
fig. 13), there is evidence here also that the series may consist of 
eight endites and an apical lobe. It may be that the ancestral 
crustacean possessed even more endites, but there is some reason 
for believing that it had eight only, since, as will be shown later,, 
that number allows an arrangement which corresponds with the 
segmentation of the thoracic limb of Malacostraca, and with what 
was probably the primitive structure of the maxilla both in 
Decapoda and in Copepoda. If this view be adopted, there may 
be recognized in the primitive crustacean limb nine successive 
regions—eight represented by the endites and a ninth consisting 
- of the apical lobe. These regions would be potential segments, 
since jointing would certainly take place between the endites rather 
than across them, as, in point of fact, it is seen to have done 
in Triarthrus (text-fig. 6), in the larval maxilla of Penewus (text- 
fig. 16), and in Notostraca (text-fig. 3). In the latter there may 
be found, immediately behind the attachment of each endite, a 
distinct articulation. In the case of the distal members of the 
series, the articulation extends only as far as the longitudinal scle- 
rite by which the part of the axis that bears the subapical lobe, 
flabellum, and epipodite is separated from the rest. Just behind 
the flabellum, however, a very sharp jointing extends right 
across the limb, and another less marked articulation runs from 
the base of the epipodite to the inner margin, which it reaches 
just above the attachment of the blunt-ended proximal endite. 
Yet another articulation, starting from that just mentioned, 
joins the inner edge of the limb behind the base of the first of 
the large, pointed “ondites, thus cutting off a roughly triangular 
segment which in Apus, but hardly in Lepidurus, projects as a 
small lobe like an endite 7. 

The proximal endite—the gnathobase —of the primitive limb 
may be judged from the evidence afforded by the Branchiopoda 
and Triarthrus to have differed in shape from the others, been 
directed towards the mid-ventral line of the body, and served for 
the manipulation of food. Probably it had a stout, blunt end, 
set with rows of strong, short bristles (text-fig. 27). The 
remaining endites were, most likely, subsimilar to one another. 

It is natural to attempt to derive this limb from the parapo- 
dium of an Annelidancestor. If the limb was uniramous, we must 
seek for analogies among uniramous parapodia, such, for instance, 
as those of the Sy Nlide or the Eunicide. In the latter family 


* See footnote to p. 48, 
7 See footnote, p. 48, 


46 MR. Le. A. BORRADAILE ON THE 


(text-fig. 1), the arrangement of the parts is strikingly suggestive 
of those of the phyllopod limb, the so-called ‘“ dorsal cirrus ”— 
really the notopodium, whose small size and position upon the . 
base of the large straight neuropodium make the parapodium 
uniramous—standing for the flabellum, the gill for the epipodite, 
and the short, broad, ventral cirrus for the gnathobase. Needless 
to say, no more than analogy can be claimed for this likeness. 
The point of origin of the Arthropoda from their worm-like 
ancestors is not to be settled merely by a general and hypothetical 
resemblance of the limbs. 


Text-figure 19. 


iy iy BA 


Maxilla of Lysiosquilla maculata. 


For lettering see p. 71.! 


6. It has already been pointed out (p. 39) that the presence 
upon the hinder part of the head of Malacostraca, Ostracoda, and 
Copepoda of appendages not greatly modified from the assumed 
primitive type (text-figs. 13-21) justifies the surmise that in the 
original crustacean the series of similar limbs extended further 
forward than in Branchiopoda. Probably it resembled that of 
the Trilobita in including all the appendages behind the anten- 
nules, but comparison of the maxilla of the Decapoda with the 
pygidial limbs of 7riarthrus suggests that there was less unlike- 
ness between the extreme members than in that genus. Hach 
appendage was no doubt capable of being used for swimming, for 


MOUTH-PARTS OF THE PALAMONID PRAWNS. AT 
the manipulation of food, and for respiration. In all recent 
Crustacea, however, the specialization of certain of the limbs for 


each of these functions, and for a fourth, contact with large 
bodies, sueh as prey, or the substratum, has caused them to 


Text-figure 20. 


EX~Lf 
PREP? Ep? £ EL 


Text-figure 21. 


Text-fig. 20. Maxillule of Calanus sp. 
3 21. Maxillule of Cypris sp. 


For lettering see p. 71. 


depart in various directions from the type. ‘Those which were 
specially adapted for swimming became biramous*, those for 


* This change must have taken place more than once. It is already established, 
for instance, in the antennze and larval mandibles of various Branchiopoda, which 
show no trace of it in their other limbs, and it is well developed upon the abdomen 
of the Nebalia, while it is still but incipient in the thorax. The Conchostraca 
(text-fig. 5), whose trunk-limbs are almost biramous, show how easily that condition 
can arise by an alteration in the proportions of the parts of the limb. 


48 MR. L. A. BORRADAILE ON THE 


mastication developed the gnathobase with or without some of the 
other endites, those for respiration the epipodites, and those for 
contact with large objects the main axis. Generally speaking, the 
limbs which in the adult are adapted to other functions than 
swimming are in the larva (and were perhaps at one time through- 
out life) natatory, and retain in their later condition traces of the 
biramous plan, so that they may be regarded as belonging to the 
biramous type. In becoming biramous, the limb has lost its leaf- 
like character. This has in all cases befallen the antenne and 
mandibles, which, precociously adapted in the Nauplius to swim- 
ming, never exhibit the primitive configuration. On the other 
hand, the maxille almost invariably retain a good deal of resem- 
blance to the phyllopod prototype, probably because their position 
makes it impossible for them to be of much use either as jaws or 
for swimming. The trunk-limbs of Branchiopoda are phyllopod, 
and show in some detail the features which it is necessary to 
attribute to those of the ancestral crustacean *; those of the 
other groups are biramous 7, with the exception of the thoracic 
appendages of the Leptostraca, which exhibit various degrees of 
transition from the phyllopod to the biramous condition. 


* The lobes of the trunk-limbs of Branchiopoda (text-figs. 3-5) are exceedingly 
difficult to homologize. It seems, for instance, quite possible that the Anostraca 
have no flabellum, and that the structure known by that name which is jointed to 
the end of their limbs corresponds to the so-called last (sixth) “endite” of Apus and 
Limnadia. If so, however, an additional endite must be intercalated into the series 
in Anostraca or excalated from it in Notostraca, for in the former group there are 
six of these lobes before the terminal structure in question, but in the latter only 
five. The facts of meristic variation would explain this discrepancy without 
difficulty, but the actual solution is probably indicated by the presence in Apus, 
though barely in Lepidurus, of what looks hke a vestigial endite, standing in the 
gap between the first and that which has been regarfled as the second of the series. 
This little lobe bears no bristles, but the section of the axis opposite it is defined by 
lines of soft cuticle, such as those which mark out the segments of the other endites. 
If there be here a true endite, the series in Apus corresponds with that of Anostraca. 
Lepidurus must then be considered to have lost the second endite (though not its 
segment), and in Limnadia it has perhaps fused with the third. It is true that 
this hypothesis is open to the objection that it brings the epipodite opposite the 
third endite, but such a displacement presents no great difficulty in view of the 
vestigial nature of the supposed second segment, which does not reach the outer 
edge of the limb, and of the fact that the epipodite is attached not wholly to the 
third segment, but astride of the articulation between it and the portion of the limb 
proximal to it. Again, the little process known as the “‘subapical lobe,” present in 
Apus and Branchipus, but barely or not at all represented in some other genera, as 
Chirocephalus and Limnadia, may or may not represent the true end of the axis 
of the limb. If it does not, then the apex must be the so-called last endite. This 
(if the above conjecture with regard to the flabellum of Anostraca be true) is always 
an organ sui generis, separated by a more or less distinct joint from the rext of the 
limb. In either case, the condition of the apical lobe in the Branchipoda is 
abnormal as compared with that which it shows in the larval maxilla of the 
Decapoda and in Tviarthrus, which in this respect probably more nearly represent 
the ancestral Crustacea. To judge from the thoracic limbs of Nebalia (text-tig. 9), 
whose end-joint differs strongly from the rest and is much more sharply articulated, 
the appendages upon the trunk of the phyllopod ancestor of the Malacostraca must 
have resembled those of the Branchiopoda in respect to the apical lobe. 

+ The limbs of the Trilobita (text-figs. 6, 7) present an interesting problem in 
that, while they are undoubtedly biramous in the sense of consisting of two subequal 
rami upon a common stem, their “ exopodites ” are inserted in a different position 
from those of recent Crustacea or from the flabella of the Branchiopoda. The 
“endopodite”’ of the pygidial limb (in Triarthrus) is clearly comparable with the 


MOUTH-PARTS OF THE PAL-AAMONID PRAWNS, 49 


7. The relation of the parts of the biramous limb to those of 
its leaf-like forerunner may be elucidated by a study of actual 
phyllopod appendages. It would seem that the flabellum is 
represented by the exopodite, and the main axis, ending in the 
apical lobe, by the protopodite and endopodite. In regard to the 
exopodite, the evidence of the Leptostraca, in which it is flattened 
and bears just the same relation to the other parts of the limb as 
the flabellum, is too strong to be set aside without more con- 
vineing reasons than have been adduced. Theories which 
disregard this consideration depend upon the precarious support 
afforded by a comparison of the arrangement in various cases of 
the endites. These structures, however, are very variable and 
difficult to homologize, and in particular those of the maxilla, 
which in adult reptant Decapoda appear to suggest that the fifth 
and sixth endites have become endopodite and exopodite, have in 
more primitive members of the same group a quite different 
aspect, which supports strongly the theory stated above. The 
larvee of. various Carides, as, for instance, that of Palemonetes 
(text-fig. 18), show at the base of the endopodite of the maxilla, 
in the region of the ischipodite, a distinct fifth endite. In other 
cases, as in Anisocaris (text-fig. 17) and Pencus (text-fig. 16), 


axis of the phyllopod limb, and in shape more nearly resembles the larval maxilla 
of a Decapod than the trunk-limb of a Branchiopod. It is remarkable only for the 
comparatively unimportant facts that the region of the third and fourth endites, 
instead of forming a single joint (the basipodite), is divided by an articulation int) 
two, each of which bears one endite, and that beyond the fourth endite the limb has 
but four segments, instead of the five which the maxilla of Cerataspis (text-fig. 15) 
indicates as the full complement of the corresponding region of the decapod limb. 
It does not seem impossible that an additional segment may eventually be 
discovered at the end of this appendage. The conditions are none too clear in 
Beecher’s model. ‘The “‘exopodite,’ however, stands upon the segment of the 
second endite, not upon that of the third and fourth. Probably this indicates that 
the structure here called the exopodite is not homologous with that to which the 
same name is applied in recent Crustacea, but is a modified epipodite. In that 
connection it is interesting to note that Beecher attributes to it a respiratory 
function. If the foregoing theory be correct, the thoracic limbs of Trilobita present 
a very remarkable analogy to those of Decapoda, consisting as they do of a sub- 
cylindrical, seven-jointed axis with a complex respiratory structure borne on the 
outside near the base. Unlike the podobranchs of the Decapoda, however, the 
epipodite of the Trilobita “contrives a double debt to pay” as gill and “ exopodite.’ 

The foregoing interpretation of the limb of Tviarthrus receives very interesting 
support from the arrangement of the parts of the mesosomatic appendages of 
Limulus. Here the axis bears on its outer side three broad exites, separated from 
it and from one another by sutures to which correspond notches on the edge of the 
limb. (The first exite of the gill-bearing limbs is, upon its anterior face, divided 
into about a dozen strips by less-marked sutures, of which only two remain in the 
genital operculum.) That part of the axis which bears the first and second exites is 
unjointed, but the third is attached to a well-marked segment, and beyond this two 
free joints form the apex of the limb. ‘The first and second exites are crossed 
obl quely upon their hinder face by the gill-lamelle, It is pretty clear that they 
are epipodites, and a comparison at once suggests itself between them and the very 
differently shaped “exopodite” which bears gill-lamelle in Triarthrus. The third 
exite in Limulus, sharply distinct from the second, is no doubt the exopodite. 

The supposed biramous mouth-parts of Insecta are not so in reality, and I take 
this opportunity of retracting the view I haye expressed on that point in my 
‘Manual of Zoology’ (London, 1912). The lacinia and galea are, I now believe, 
the third and fourth endités, and the palp is the endopodite. 


Proc. Zoou, Soc,—19]17, No. LV. 4 


50 MR. L. A. BORRADAILE ON THE 


this is succeeded by a sixth and a seventh, while in Cerataspis 
(text-fig. 15) the endopodite bears four endites, making eight 
altogether, besides the apical lobe in the position of the dactylo- 
podite. In the last two cases it is clear that the endopodite 
represents the whole distal part of the limb, and not, as has been 
supposed, the fifth or sixth endite only. 

For a detailed comparison of the jointed stem formed by the 
pr otopodite and endopodite of the biramous limb with the endite- 
bearing axis of the phyllopod appendage, it is necessary to find 
some fixed point of correspondence between members of the two 
series. It would be natural to look for this at the base of the limb, 
but the development of Pencwus and other Decapoda, during which 
the basal joint of the appendages of the thorax recedes into the 
side of the body, is a warning against the use of such a method. 
On the other hand, the differences in the total number of endites 
in different limbs makes it useless to start a comparison from the 
distal end. ‘The epipodites vary in number, and their homologies 
are doubtful, so that their points of attachment afford no reliable 
clue. There remains the flabellum-exopodite. It is unfortunate 
that the uncertainty which exists in regard to the homologies of 
the endites of Branchiopoda makes even the seat of the flabellum 
a doubtful point in that group, but, as will be seen, what can 
be stated on this point is not without its uses in confirming the 
evidence of other classes. In the Anostraca (text-fig. 4) the 
flabellum is probably absent. In Conchostraca (text-fig. 5) it 
stands opposite the junction of the second and third endites of a 
series of six, but, quite possibly, these two members represent the 
third and fourth of the full series, the second, which is present 
in the Anostraca, being in Notostraca vestigial and in Concho- 
straca either lost or fused with the third to form a single 
elongate lobe. In Notostraca (text-fig. 3), where the whole axis 
of the limb is more or less strongly segmented, the flabellum 
stands on the segment of what may be the third or the fourth 
endite. In the maxilla of the Malacostraca (text-figs. 14-18, and 
47) the situation is clear. Here the exopodite arises opposite the 
third and fourth endites, upon a region of the limb which may be 
undelimited, as in most Decapoda, but which is often marked off 
more or less completely to form a segment, as in Leptostraca 
(text-fig. 14), Mysidacea, Cumacea, Kuphausiacea, and the decapod 
larva Anisocaris (text-fig. 17). Where there is no exopodite, 
as in Isopoda, Amphipoda, and Stomatopoda * (text-fig. 19), a 
segment with two endites is found in the same position on the 
maxilla as on that of the orders in which the limb bears an 
exopodite. The maxilla of Calanus (text-fig. 13) agrees remark- 
ably in structure with that of Malacostraca. Though it has no 

* In the maxilla of the Stomatopoda, the basipodite is represented upon the 
outer side of the limb by a single sclerite, but the segment of the third endite is 
differentiated trom that of the fourth by the possession of a sclerite of its own upon 
its hinder face. The structure of this rather puzzling limb is elucidated in text- 


fig. 19. _It is interesting to note that the maxillary gland opens on the second 
joint, like the anteunary “eland of Mysidacea and Amphipoda. 


MOUTH-PARTS OF THE PALE MONID PRAWNS. 51 


exopodite, there may be found in its proximal half a double 
segment bearing two endites. At first sight, the endites upon 
this segment appear to be the second and third, but the length 
of the segment which precedes it, and the position of the endite 
upon the latter at its distal end, suggest at once that that is 
also a double segment, that here, as in Carides, the first endite 
has been lost, and that the second double segment therefore bears 
the third and fourth endites and is the basipodite. This 
impression is strengthened by the fact that beyond the segment 
with two endites there are five simple segments, as there are 
beyond the endites which represent the basipodite of Cerataspis 
(text-fig. 15), which, since in that respect it agrees with the 
thoracic limbs of Malacostraca, may be taken as indicating the 
original number of segments in this region in the malacostracan 
maxilla, In the maxillule of Calanus (text-fig. 20) the first 
endite is carried upon a fairly distinct segment, but the second, 
third, and fourth lie upon a region which is undivided save by 
an incomplete constriction, which marks off the distal portion of 


Text-figure 22, Text-figure 23. 


a 


Maxillule of Lepas sp. Mandible of Lepidurus sp. 


it, where the fourth endite and the exopodite are borne. The 
maxilliped of Calanus (text-fig. 11) has a very short basal 
segment bearing the small first endite, a long second segment 
bearing the second, third, aud fourth endites, and six distal 
segments, which is one more than might be expected. Meristic 
rearrangement has probably been at work in this portion of the 
limb*. The maxillule of Cypris (text-fig. 21), which curiously 
simulates the maxilla of Carides, seems, like the latter appendage, 
to have lost the first endite. If that be the case, the exopodite 
here stands opposite the third and fourth endites. The thoracic 
limbs of Nebalia (text-figs. 8, 9), though they present no endites, 


* An increase beyond the normal of the number of joints in a crustacean limb 
may take place in three ways: (1) by the annulation of the unsegmented apex of 
either branch of the limb to form a “ flagellum,” as in antenne, thoracic exopodites, 
and pleopods of Malacostraca, ete. Three faint annulations upon the apical 
segment of the maxilla of Calanus are probably of this nature ; (2) by subdivision 
of a segment, as in the earpopodite of various Carides; (3) by resegmentation of a 
limb or of part of it, as probably in the maxilliped of Calanus. Fe 


52 MR. L. A. BORRADAILE ON THE 


are instructive in regard to the segmentation of the phyllopod 
limb. Here the number of the segments which are formed in the 
endopodite varies with the limb and with the age and sex of the 
individual. Nearly always the four distal segments can be made 
out, and often a fifth is to be found proximal to them, leaving a 
long basipodite, which is always clearly marked off from the coxo- 
podite. Across the basipodite, distal to the exopodite, there 


Text-figure 24. 


\ WAV AN 
i | ! yy ap 
AY { ye 
WW LLG 
ex 
bas a 
(3 +4) 2 
COX ¥., 
proxi. 4 
OO, 
ss \ en bts Vie 
— |_ Ces = 
di) a : 
: RR 4 ? 
Ge ges. & 


Text-figure 25. 


Text-fig. 24. Mandible of Calanus sp. 
5s 25. Mandible of Cypris sp. 


For lettering see p. 71. 


oceasionally appears an additional joint, the faintest of the series, 
indicating the double nature of the segment *. 


* It is perfectly true that most of these segments are without muscles, but from 
that it does not follow that their evidence may be disregarded. Probably they are 
vestigial, haviag lost their endites,and not, as where the limb becomes subcylindrical, 
acquired a value of their own. The division of the basipodite probably oveurs also 
in Anaspides (text-tig. 12), where, in the anterior thoracic limbs, the endopodite 
appears to contain six joints, but is flexed between the third and fourth of these, 
not between the second and third as in the Eucarida. It seems likely that the first 


MOUTH-PARTS OF THE PALZ MONID PRAWNS, 53 


It appears, then, that the flabellum-expodite, which in the 
unjointed limb arises opposite the third and fourth endites, is, 
when the appendage becomes jointed, generally borne by a double 
segment, representing those of both the lobes in question, but if, 
as occasionally happens, the two components of this segment 
become distinct, the exopodite may be found either upon the 
proximal of them (Vebalia etc.) or upon the distal (maxillule of 
Calanus, Notostraca?). Thus in the axis of the phyllopod limb 
the region of the third and fourth endites corresponds to the 
basipodite of the biramous limb, that of the first two endites to 
the rest of the protopodite, and all that part which les beyond 
the fourth endite to the endopodite. The doubleness of the basi- 
podite makes necessary certain terms for the distinctton of its 
parts. Its two components may be known as the probasipodite 
and metabasipodite, and when either of these alone acts as the 
basipodite by bearing the exopodite, it is a hemibasipodite. In 
contradistinction to this the complete double joint may be called 
the symbasipodite. 


ia hae PRLS V7 b< 
Text-figure 26. Text-figure 27. 


rane Talis 
(( 
\\ 


Mandible of Lepas sp. Mandible of Nebalia sp. 
For lettering see p. 71. 


In the proximal part of the branchiopod limb, the most per- 
sistent of the epipodites stands opposite the second endite. The 
segment to which these structures belong is the coxopodite. it 
other epipodites (pro-epipodites) be present, they stand in the 
region of the gnathobase. This region is the so-called “precoxa, 
or “ pleuropodite,” which may or may not have originally existed 
as a free joint in every biramous limb, but has now nearly always 


segment after that which bears the exopodite is here not the ischiopodite, bus 
the second division of the basipodite, or, as it may be called, the metabasipodite. 
If this be so, the question arises whether the basipodite be not divided in the 
Pericarida, whose thoracic endopodites also are flexed between the apparent third 
and fourth joints. But in that case their apical joint must represent the fused 
propodite and dactylopodite. With Calman, I am unable to regard the so-called 
“ stylopodite ” as more than an enlarged, terminal spine. If it be one of the primary 
members of the axis of the limb, it must represent the apical lobe, and the total 
number of joints in the axis becomes ten, as in the maxilliped of Calanus. 


54 MR. L. A. BORRADAILE ON THE 


disappeared, either by fusion with the trunk or with the second 
joint, or perhaps sometimes by excalation. 

The fifth endite and those distal to it belong to that part of the 
limb which stands beyond the attachment of the flabellum, and 
represent the endopodite of the biramous limb. The maxille of 
Cerataspis and larval Natantia (text-figs. 15-18) seem to show 
that each of these endites is borne upon a portion of the limb 
which represents a single joint of the biramous appendage, and 
that the apical lobe is an unsegmented distal region of varying 
extent, corresponding to the dactylopodite and any adjacent 
segment or segments not represented by an endite. 


“ 


Text-figure 28. 


Guathobase of thoracic limb of Lepidurus sp. 


It is fair to assume that all these relations existed in the 
primitive crustacean appendage, and that the latter gave rise to 
the biramous limbs by a transformation in which the axis of the 
limb became jointed in the way indicated, the endites in great part 
or altogether disappeared, and the flabellum approximated in shape 
to the distal part of the axis and came to stand side by side with 
it at the end of the third (or, if the precoxa were not separate, 
the second) joint of the limb. 

8. With the origina] phyllopod limb, thus reconstructed, the 
jaws of Malacostraca may be compared as follows:—In the 


MOUTH-PARTS OF THE PALZMONID PRAWNS, 5D 


mandible, homologues must be found for the molar process, the 
incisor process, and the palp. The molar process pretty clearly 
represents the mandible of the Branchiopoda, and through that 
the gnathobase of the phyllopod limb. The grinding-surface of 
the mandible of Chirocephalus (text-fig. 30) is covered by two 
comma-shaped patches of fine ridges, each ridge consisting of a 
row of tubercles, which perhaps represent the rows of stout 
bristles upon the end of the gnathobases of the trunk-limbs of 
Notostraca (text-fig. 28). In places the ridges bear some hairs. 
The commas are reversed, so as to fit against one another, and 
one patch is narrower then the other and composed of coarser 
ridges. In view of its structure, position, and function, there 
can be little doubt that this organ is a specialized gnathobase. 


Text-figure 29. Text-figure 30. 


End view of maxillule of Lepas sp. End view of mandible of Chirocephalus sp. 


Two sets of ridges, very similar to those on the mandible of 
Chirocephalus, may be recognized at the end of the molar process 
in Nebalia (text-fig. 31), though here one is of less extent than 
the other, and seems largely to have been replaced by rows of 
delicate hairs. Fine ridges, as may be seen in the figures given 
by various authors, exist also upon the molar processes of various 
Malacostraca (Anaspides, Gammarus, etc.). In the Decapoda a 
crescentic or comma-shaped set of serrated ridges is often present 
(text-figs. 32, 43), with or without patches of hairs and granules, 
though they tend to disappear, and the molar surface is often 
broken up into a small number of large lobes. The ridges vary 
greatly in width, degree of salience, and serration, but are often 


56- MR. L. A. BORRADAILE ON THE 


much like those of Vebalia (text-figs. 38-43). Clearly, we have 
here the mandible of Chirocephalus in process of further evolution. 
The homology of the incisor process is more puzzling. No trace 
of it can be found in the Entomostraca. In Lepidurus (text- 
fig. 33) the end of the mandible, instead of being oval as in 
Chirocephalus, is elongate, tapering to one end, and slightly 
curved. It is crossed by deep ridges, few in number, and marked 
each by a few coarse tubercles. The narrow end stands away 
from the rest, bears three or four teeth instead of the ridges on 
the wider part, and somewhat suggests an incipient incisor 
process, but this is at the hinder end of the organ, whereas the 
incisor process of Malacostraca is anterior. It seems likely that 


Text-figure 31. 


End view of mandible of Nebalia sp. 


For lettering see p. 71. 


the biting-surface of the mandible of Notostraca represents one 
of the two sets of ridges found in the same position in Chiro- 
cephalus. In Calanus (text-fig. 35) the condition is not dissimilar. 
The biting surface is narrow, elongate, tapering towards the 
ends, especially towards the hinder end, and crossed by coarse 
ridges, variously tuberculate. At the anterior end stands a 
structure which at first sight a little recalls the incisor process, 
but in an end view of the mandible this is seen to be only the 
first ridge, removed a little from the rest and connected with 
them by a flange. The mandible of Cypris (text-fig. 34) is of 
the same type, but the anterior ridge is less outstanding and not 


MOUTH-PARTS OF THE PALAMONID PRAWNS. 7 
flanged to the next, the ridges are more regular, the valleys 
between them bear spines, and at the hinder end here is not, as 
in Calanus, a single long, fringed spine, but a pair. Probably 
the biting-surfaces of the mandibles of Calanus and Cypris are 
homologous with one another and with that of Lepedurus. The 
mandible of Zepas (text-figs. 26, 36) is a very different structure, 
thin, with a sharp, toothed edge, and in side view very similar 
to the incisor processes of Pere Decapoda. It is crossed near 
the base by an articulation, and it seems possible that its toothed 
edge represents, not the gnathobase, but the second endite. On 
the other hand, the maxillule of the barnacle (text-figs. 22, 29), 
which appears fae be built on the same plan, is of a fain width at 
the end, and crossed by short ridges, each bearing a row of five 
spines, so that this limb forms a sort of ti Seinon from the 
ridged condition of Cypris to that of Lepas. Unless the mandible 

of the barnacles be homologous with the incisor process, the latter 


Text-figure 32 


End view of mandible of Desmocaris. 


For lettering see p. 71. 


is a structure peculiar to the Malacostraca, and not to be found 
even in rudiment upon the mandible of any other crustacean. 
Two modes of origin can be suggested for it. It may be regarded 
either as an outgrowth from the gnathobase or as representing 
the second endite. On the whole, the latter is the more plausible 
view. From its first appearance in WVebalia (text-figs. 27, 31) 
this process has a knife-edge, quite unlike the ridged surface of 
the molar process and its homologues, although it often comes to 
hear a row of teeth. As has been shown, there is no trace in 
phylogeny of its origin from the proximal part of the limb, and, 
though in ontogeny (as, for instance, in that of Penwus or the 
Stomatopoda) it and the molar process may originate from the 
same simple precursor, this is no more than the “differentiati ion, 
in the course of development, of the rudiment of the appendage, 
and happens also in the case of the palp. If this view be correct, 


58 MR. L. A. BORRADAILE ON 'THE 


the basal joint of the mandible of Malacostraca corresponds to 
the fused precoxa and coxopodite. Those of Branchiopoda 
represent the precoxa only, the rest of the limb having dis- 
appeared. The basal segment of Copepoda, again, corresponds 
to no more than the precoxa, the coxopodite being represented 


Text-figure 33. Text-figure 34. 


Text-figure 35. Text-figure 36. 


Text-fig. 33. End view of mandible of Lepidurus sp. 
” 34. End view of mandible of Cypris sp. 
3 385. End view of mandible of Calanus sp. 
» 986. End view of mandible of Lepas sp. 


by the little segment between that which bears the biting-edge 
and that upon which stand the exopodite and endopodite in 
Calanus (text-fig. 24). In Ostracoda (text-fig. 25), it would seem, 
the coxopodite must either have been excalated altogether or 


MOUTH-PARTS OF THE PALEHMONID PRAWNS. 59 


have fused with one of the adjoining segments. The mandibular 
palp represents the remainder of the axis of the limb. Its first 
segment is undoubtedly the basipodite, though there is nothing 
to show whether it is a’complete symbasipodite. Often a joint 
divides the remainder into two parts, but these cannot be regarded 
as corresponding with any of the normal segments of the 
endopodite *. 

The mandible of the Crustacea is an exceedingly complicated, 
varied, and interesting organ, presenting many problems and 
worthy of a great deal more attention than it has received. 


Text-figure 37, Text-figure 39. 


Text-figure 38. 


Text-fig. 37. Left mandible of Leander serratus. 
9 38. End view of molar process of the same limb. 
A 39. End view of right mandible of Leander serratus. 


For lettering see p. 71. 


The laciniz of the malacostracan maxillule (text-fig. 46) repre- 
sent the first and third endites. This is suggested, though hardly 
proved, by the condition of the limb in Webalia. Hansen has 
shown that in various other genera the lacinie belong to the first 


* G. W. Smith (Q. J. M.S. liii., iii. p. 506, 1909) describes a mandibular palp 
in Paranaspides which would be quite unique among such structures in the 
Malacostraca, having four joints and an exopodite. JI am unable to confirm this. 
All the specimens in the British Museum have three joints and no trace of 
exopodite. 


60 MR. L. A. BORRADAILE ON THE 


and third segments. In many Decapoda, as in Leander (text- 
fig. 46) the same fact is clear upon careful examination. I shall 
call the processes in question the inner and outer lacinie. The 
outwardly-directed lobe which is often present upon the first 
segment has been regarded by various authors as the exopodite, 
and the conclusion drawn that the part of the limb proximal to 
the basipodite is lacking. It is hard to see upon what evidence 
this view can be based. The exite is much more easily inter- 
preted as a proepipodite. It is represented in Leander by a hump 
upon the base of the first endite. 


Text-figure 40. Text-figure 41. 


ay 2 SS 

GRAS Were 
re Nh 

¢ ) \\’ - 

a RR o 


Text-figure 42. Text-figure 43. 
Text-fig. 40. End view of left mandible of Conchodytes tridacne. 
4 41. End view of left molar process of Periclimenes spiniferus. 
a 42. Oblique view of end of right molar process of the same species. 
43. Same view of right wolar process of Saron marmoratus. 


The two cleft lobes usually known as the “lacini#” of the 
maxilla (text-figs. 14-17) are each equivalent not, as has some- 
times been supposed, to one of the elementary segments of the 
limb, but to two, and thus they together represent the first four 
endites of the primitive limb. This interpretation is strongly 
suggested by such appendages as the maxille of larval Penzeides 
(text-fig. 16) and especially by that of Cerataspis (text-fig. 15), 
and is not contradicted by the fact that articulations are often 


MOUTH-PARTS OF THE PALZMONID PRAWNS. 61 


not developed between the first and second or between the third 
and fourth segments—as, for instance, in Anisocaris (text-fig. 17), 
which is an instructive case of this condition. ‘The first endite is 
frequently absent, whether or not its segment be present as a 
separate entity. In the Peracarida, the number of endites is 
generally reduced in this way to three (or fewer, if another of 
them be absent), though the Mysidacea at first appear to form an 
exception. In JM; Wee however, closer examination reveals a con- 
dition which may be described as follows. ‘The edge of the second 
segment proximal to its endite is rounded, meets the base of the 
endite in a notch, and bears a row of bristles which is continued 
from the notch across the origin of the endite. The notch forms 
an outline which simulates the cleft lobe formed in Hucarida by 
the first two endites. If this suggestion be correct, the Mysi- 
dacea, liike other Peracarida, must be regarded as having lost the 
first endite of the maxilla. In Leptostraca (text-fig. 14), Anaspi- 
dacea, Stomatopoda (text-fig. 19), and typical members of the 
Eucarida (text-figs. 15-17), it is present. The degree to which 
the first and second, and again the second and third, endites are 
associated to form cleft lobes varies, and is highest in some 
Decapoda. I shall allude to these double structures—the so- 
called ‘“‘lacinie” of the maxilla—as the first and second lobes. 
The first comprises the endites of the precoxa and coxopodite, 
the second those of the basipodite. The fifth endite is often 
represented in the adult by a slight swelling at the base of the 
endopodite. The exopodite (scaphognathite) is a little-modified 
flabellum, and a small rounded lobe proximal to it in Eucarida 
perhaps represents the epipodite. 

In the first maxuliped of Decapoda (text-fig. 48) the third 
and second endites are distinct. The latter is often marked 
by a shght notch, which, however, is not likely to indicate the 
presence of the first endite, the precoxa being probably fused 
with the body in the thoracic segments of this group. Jn 
Anaspides, as is Shown by the history of the development of the 
gill-rudiments (text-fig. 12), the first and second endites are both 
present, and the basal joint carries also two epipodites, from 
which it would appear that the precoxa in this genus is fused 
with the coxopodite. The Peracarida probably agree with 
Anaspides in this respect, as they have two epipodites (oostegite 
and gill) upon the basal joints of the thoracic limbs. Endites 
may be present upon the endopodite of the maxilliped in the 
Peracarida, as in Mysis (and perhaps also in Gammarus, but in 
the latter there is some doubt about the identity of the apparent 
ischiopodite which bears the endite in question: see footnote to 
p- 53). Knobs bearing bristles which sometimes appear upon 
the coxopodites of the second and third maxilliped of prawns 
(text-figs. 49, 50) may represent the second endite, and a similar 
knob in the sternal region at the base of the second maxilliped 
may be a vestige of the gnathobase. 


62 MR. L. A. BORRADAILE ON THE 


III. 


1. The mandible of the Palemonide (text-fig. 37) is deeply 
cleft into two diverging processes, both directed obliquely towards 
the median plane of the body. One of these—the incisor process 
—is a thin structure, more or less ribbon-like in the Pontoniinz 
and Desmocaris (text-figs. 32, 40), but shorter and broader in 
Leander serratus (text-fig. 37). It trends, at its base, downwards, 
but curves inwards and at the same time twists its outer edge 
forwards, so that, while at its base it is nearly vertical, with its 
width transverse to the body, at its free end it is nearly horizontal, 
with its width longitudinal to the body. The other—the molar 
process —is stout and subrectangular in section, and slants dorsally, 
to end somewhat obliquely truncated on the median plane. In 
the Paleemonine a delicate palp, usually three-jointed, stands on 
the anterior side of the limb, at the base of and just dorsal to the 
incisor process, along whose outer edge it curves towards the 
middle line of the body. The only Pontoniine which possess a 
mandibular palp are Urocaridella and Paleemonella. In these it 
is two-jointed. 

The incisor process of Pontoniinz usually ends in three teeth, 
the midmost of which is shorter than the others, but there are 
sometimes more. Thus in Coralliocaris japonica there are four 
on one mandible and five on the other, and in Conchodytes 
tridacne (text-fig. 40) there are on one side five, nearly equal, 
and on the other six. In Leander serratus (text-fig. 45) there 
are two large teeth, with on the right mandible one, and on the 
left two, smaller intermediate teeth. In Desmocaris (text-fig. 32) 
there are four teeth on the right and five on the left mandible, 
those at the outer ends of the row being rather larger than the 
others. In each case the arrangement is not such that the teeth 
of the two sides can closely interlock. The molar process of 
most Palemonide (text-figs. 38-42) ends in a roughly square 
concave surface, around which is an incomplete wall composed of 
from four to six projecting lobes. Some of these have crescentie 
or horseshoe-shaped rims, with their open sides towards the 
middle of the process. Others are completely rimmed, but raised 
more on the outer side than on the inner. In Leander serratus 
(text-figs. 38, 39) there are four sharply distinct lobes. The lobes 
differ a good deal in shape, and there is only a general corre- 
spondence between those of the mandibles of the two sides, So 
far as this correspondence goes, it is not the mirror-likeness 
usually found in paired structures, but the two arrangements are 
reversed, so that there is a rough sort of interlocking. In 
L. serratus the lobes are only roughened in places. In the 
Pontoniinz one lobe, and part of the rest of the surface, bears a 
fur of bristles or is roughened by tubercles. This, I believe, is 
the last remains of the clothing of bristles found on the end of 
the molar processes of other Carides, such as the Alpheide. In 
the primitive Palemonid Desmocaris (text-fig. 32) the process 


MOUTH-PARTS OF THE PALZMONID PRAWNS, 63 


ends in asemicircular surface, which bears a comma-shaped patch 
of serrated ridges. Sollaud, who called attention to the difference 
between the molar surfaces of Desmocaris and those of other 
Palemonide *, pointed out that these ridges also occur in the 
Acanthephyride, and regarded them as a primitive feature. In 
this he is probably right, since, as 1 have shown, a similar 
arrangement is found in Nebalia and certain Branchiopoda. In 
Amphibeteus and some Hippolytide (text-fig. 43) part of the 
surface 1s covered with such ridges and part with bristles, and 
lobes of various shapes are appearing. We.have here probably a 
transition from the ridged condition to that which is found, for 
instance, in Pontoniine. 


Text-figure 44. 


Text-figure 45. 


Text-fig. 44.—Ventral view of mouth-parts of Leander serratus after removal 
of maxillipeds, maxillw, and left maxillule. 
Text-fig. 45.—The same after removal of maxillules and paragnatha. 


For lettering see p. 71. 


2. The mandibles lie (text-figs. 44, 45) in a chamber enclosed 
between the lips, the hood-like labrum standing in front of them, 
and the large bilobed metastoma behind, while the swollen bases 
of the mandibles themselves close in the chamber at the sides. 


* C,R. Ac, Sci. clii. p. 913 (1911). 


64 MR. L. A. BORRADAILE ON THE 


There are two openings to the lip-chamber—a narrow median 
shit between the lobes (paragnatha) of the metastoma and a wider 
transverse gap between the metastoma and the labrum. The 
incisor processes close the transverse opening, meeting in the 
middle line. The molar processes meet deeper in the chamber, 
just under the opening of the gullet. 

3. In the maxillule (text-fig. 46), the inner lacinia, which is 
usually the narrower of the two, curves towards the outer and 
is provided with relatively feeble bristles. The outer lacinia has 
very stout, yellow bristles and is probably the only structure, 
other than the mandible, which is capable of tearing the food. 
The endopodite is bifid at the tip, the proximal branch, which 
possibly represents an endite, being curved in a short spiral, the 
distal branch nearly straight. The maxillules stand close against 
the paragnatha (text-fig. 44), the lacinie of each opposed to those 
of the other across the median line, but not quite meeting them. 


Text-figure 46. 


Maxillule of Leander serratus. 


For lettering see p. 71. 


The inner lacinizw are just behind the cleft of the metastoma, 
the outer stand beside the cleft. The endopodite is directed 
outwards and hooks its curved process around the outer edge of 
the paragnathum, which has a notch to receive it. Probably 
this gives a purchase for the action of the limb. 

4, In the maxilla (text-fig. 47), the first double lobe has 
disappeared and is represented only by a slight swelling of the 
edge of the limb. The second lobe is of good length, but tends 
to become simple in the Pontoniine. In this respect it is very 
variable, the maxille of the two sides sometimes differing in an 
individual. From the frequent occurrence of reduction in the 
lobes of this hmb in the Carides, it would seem that they are 
struetures of no great physiological importance. The endopodite 
is of simple shape, tapering distally, and the exopodite (scapho- 
gnathite) varies in shape and width with the gill-chamber, and 


65 


MOUTH=PARTS OF THE PALZMONID PRAWNS. 


thus with the habit of body. The long bristles which fringe the 
scaphognathite are feathered, which probably makes the organ 


more efficient in sweeping the gill-chamber. 
Text-figure 47, 


Zijyp 1 
Maxilla of Leander serratus. 
For lettering see p. 71. 


Text-figure 48. 


CX 


= 


First maxilliped of Leander serratus. 


For lettering see p. 71. 


5. In the first maxilliped (text-fig. 48), two endites—the 
second and third—are nearly always recognizable, and usually 
: 5 


Proc. Zoou. Soc.—1917, No. V. 


66 MR. Le A. BORRADAILE ON THE 


separated by a very distinct notch. The notch in the proximal 
endite, though less distinct, can often be made out. The endo- 
podite is simple with a stout bristle near the top, and often 
indications of a joint a little further down. At the tip of the 
exopodite a few small joints may be present and at its base on 
the outer side is a fringed lobe (lobe a of Boas) which varies in 
width with the body and has perhaps some function in regard to 
the current which the scaphognathite sets up in the gill-chamber. 
Its bristles are feathered like those of the scaphognathite. The 
epipodite varies much in size, and its outer border is usually 
notched. . 

6. In the second maxilliped (text-fig. 49), the main axis 
consists of six joints, the basipodite and ischiopodite being fused. 


Text-figure 49. 


AN 


sé 
ZA SS, 
SJ 


Cox 


Second maxilliped of Leander serratus. 
For lettering see p. 71. 


The last two joints are bent strongly backwards on the inner 
side of the limb, so as to lie parallel with the ischiopodite, and, 
owing to the backward (morphologically forward) growth of a 
process of the propodite on the outer (morphologically inner) side 
of the dactylopodite, the latter comes to lie along the median side 
of the former, instead of at its apex. Thus these two joints are 
compacted into a firm plate, which presents a long median 
margin, fringed with bristles, against its fellow of the opposite 
side. This is, of course, the typical caridean condition of the 
limb. The exopodite is obscurely annulate in the greater part of 
its length, and bears at the end a comparatively small number 
of true joints. The coxopodite carries on the outer side a simple 
epipodite (mastigobranch), at the base of which may be a gill 
(Leander, Urocaridella) or the vestige of one. On the median side 


MOUTH-PARTS OF THE PALZ.MONID PRAWNS. 67 
of the coxopodite is a knob, which bears bristles, and may represent 
an endite. A sternal swelling which is sometimes present at the 
base of the coxopodite may represent a precoxal endite. 

7. The main axis of the third. maxilliped (text-fig. 50) consists 
of four or five joints (five in Leander and in Urocaridella, four in 


Text-figure 50. 


ST 


isc.mer 


Dw 


Third maxilliped of Leander serratus. 
For lettering see p. 71. 


most Pontoniine), the propodite being always fused with the 
dactylopodite and the ischiopodite with the meropodite, and often 
also the basipodite with the ischiomeropodite, in which case the 
junction is generally marked by a notch. The coxopodite bears 


on the outside a small rounded epipodite, and often on the inside 
rym 
D 


ra 


68 MR. L. A. BORRADAILE ON THE 


a bristly knob which is perhaps an endite. The exopodite is 
obseurely annulate, and at its end there are usually a few longer 
segments which are sometimes true joints, but in other cases 
appear to be marked merely by a change in the width of the 
organ and the attachment of bristles. ‘I'he ischiomeropodite is 
almost always more or less curved, with the concave side towards 
the middle line of the body. It is ribbon-like and shows in the 
Pontoniine a tendency to widen. The curving of this joint 
brings the last two joints near to those of the fellow limb, so 


Text-figure 51. 


ey 


> 
S. 


~ 
7 


| Ventral view of mouth-region of Leander serratus, all mouth-parts . 
being in place. 


that, while the ischiomeropodites le at the sides of the mouth 
with a wide gap between them in which the second maxillipeds 
are exposed, the distal parts of the limbs lie side by side in front 
of the mouth-region. <A further complexity in the arrangement - 
of the parts of the limb is brought about by the fact that the 
ischiomeropodites are twisted, so that the flat surface of the 
appendage, which in its distal part is in a horizontal plane, is in 
the proximal part in a plane between the horizontal and the 


MOUTH-PARIS OF THE PALAZMONID PRAWNS. 69 


vertical. This arrangement, combined with the presence of a 
fringe of bristles along the inner edge of the limb, has the effect 
of forming a kind of basket below the mouth-region, walled in at 
the sides by the ischiomeropodites and by the bristles which 
project downwards and inwards from their edges. In Leander * 
there are bristles borne in a diagonal row along the ventral 
‘surface of the ischiomeropodite, and also upon ‘ridges of the 
coxopodite, basipodite, and ischiopodite of the first leg (text- 
fig. 51), which complete the basket behind and below, but these 
are less well developed or absent in Pontoniine. In front of the 
mouth, the distal parts of the third maxillipeds, with their 
bristles, afford a surface, horizontally placed below the antennal 
region while the appendages are outstretched, which by bending 
can be brought ventrally under the mouth so as to complete its 
enclosure anteriorly. The last joint often possesses along its 
inner side a thick brush composed of tufts of hairs more close-set 
than the bristles of the rest of the limb. This arrangement, 
which is particularly well developed in Leander, has probably 
some special function, but I have not been able to discover what 
that may be. 


IV. 


1. It is not an easy matter to induce Leander to feed at a given 
moment, and still less so to observe what it 1s doing while it feeds. 
The animal will not take food if it is not hungry, if it is languid 
owing to lack of aeration of the water, or if it is suffering from 
shock, though sometimes it will feed surprisingly soon after 
violent operations, such as the removal of limbs. I have tried to 
observe the action of its jaws by means,of a mirror, but without 
much success. The best method is to fasten the prawn upon its 
back in a shallow vessel of sea water by means of plasticine. It 
will often feed quite freely in this position, and its jaws can easily 
be reached with a needle. When it is feeding, small particles of 
food may be seized by the chelipeds of either pair, and by them 
conveyed to the mouth, where they are generally received by the 
second maxillipeds, though sometimes they appear to be placed 
directly in charge of more dorsally placed structures, probably 
the maxillules. A large morsel occasionally appears to be steadied 
by the legs of the second pair, while those of the first tear off 
fragments and carry them to the jaws, but it is more often placed 
as a whole within the grasp of the second maxillipeds, which 
hold it in place while pieces are torn off it by deeper-lying organs, 
probably in the main by the incisor processes. In handling bulky 
masses of food, the chelipedsare assisted by the third maxillipeds, 
which bend back their last two joints for this purpose. The third 


* In this genus, in which the bristles of the third maxilliped are best developed, 
there may be made out three bands along the limb—an inner, middle, and outer, 
perhaps corresponding to the marginal, submarginal, and lateral of the four series 
found by Claus in Nebalia (text-fig. 8, p. 42). Each band consists of a succession 
of little transverse rows. Towards the ends of the appendage the bands converge 
and become merged. 


70 MR. L. Ae BORRADAILE ON THE 


-maxillipeds are aiso capable by the same action of scooping up 
food and unaided conveying it to the second maxillipeds, between 
which they sometimes thrust it with their tips. During these 
processes the basket which has been mentioned seems to serve 
the purpose of keeping the food under control till it has been 
seized by the second maxillipeds. These are very important 
organs, and play an indispensable part in passing food to the 
mandibles. ‘The animal can still feed if the legs and third 
maxillipeds have been removed, but if all the other organs be 
left and the second maxillipeds cut away it is apparently incapable 
of taking food. The second maxillipeds have three principal 
movements. In one, the broad flaps in which they end open 
downwards like a pair of doors, and with their stout fringes 
gather up the food; in another they rotate in the horizontal plane 
to and from the middle line of the body and thus narrow or 
widen the gap between them; in the third the bent distal part of 
the limbs tends to straighten so as to brush forward any object 
which lies between them. Frequently these movements are © 
combined. Once the food is past the portals formed by the 
second maxillipeds its course is hard to trace, but the following 
seems to be its fate. Ifit be small in quantity and finely divided, 
or very soft, it is abandoned to the action of the maxillules, by 
whose strong, fringed laciniz it is swept forwards and probably 
caused to enter the mouth through the slit between the para- 
gnatha. The laciniz can be moved separately, and the difference 
between them, in shape and in the kind of bristles they bear, 
probably corresponds to some difference in function. If the food 
be bulky or tough, the second maxillipeds assist the maxillules in 
brushing it forwards towards the incisor processes, The action 
of these latter is not so much a cutting as a process of tucking 
the food into the lip-chamber by first backing outwards and then 
moving inwards and rotating upwards. No doubt, during this 
the food generally undergoes some tearing, and when the mass 
of it is large, pieces have to be torn from it before they can be 
swallowed. The palp does not appear to take any mechanical 
part in the process of feeding. If it has a sensory function this 
is probably not of great importance, for the organ is present and 
absent in closely related genera in many cases among Carides. 
Finally, to enter the gullet, the food must pass between the 
molar processes and doubtless be pounded by them as it goes. 
Their concave ends are usually found to be clogged with a pasty 
matter. They must do their work very quickly, for the move- 
ment of the mandibles, as judged by that of the incisor processes, 
ceases very soon after the food leaves the latter. How swallowing 
takes place is not clear. Parker and Mocquard suggest that the | 
food of Decapod crustaceans is caused to pass up the gullet by 
suction from the crop (stomach), but, as i have shown elsewhere *, 
the case of the land hermit-crabs of the genus Cenobita throws 


* Gardiner’s ‘ Fauna of the Maldives,’ vol. i. p. 79 (1901). 


MOUTH-PARTS OF THE PALA MONID PRAWNS. 71 
doubt upon this explanation. It may be that the constrictor 
muscles of the cesophagus conduct the process. 

2. The first maxillipeds and the maxille probably take no very 
prominent part in manipulating the food. The feeble lobes of 
the maxille are in incessant movement to and from the middle 
line as they are carried inwards and outwards by the action of 
the scaphognathite. .It seems not unlikely that their sole 
function is to regulate the motions of the latter. The large 
lacinia of the first maxilliped is a rather weak structure, with 
slender silky bristles, and is not strongly moved during feeding. 
Probably, by covering the lobes of the maxilla, it prevents them 
from being clogged by the food. 

The part played by the paragnatha seems to be a passive one. 

The labrum undergoes active movements, whose function is 
probably to aid in keeping the food under the action of the 
incisor processes. 

3. The exopodites of the maxillipeds are in constant rapid 
motion, setting up by their activity a strong current forwards 
from the mouth. No doubt, this assists in carrying away the 
exhausted water from the gill-chambers and the excreta of the 
green glands poured out at the base of the antenne. But it has 
also a significance in the feeding process. From time to time 
particles are rejected by the second maxillipeds, which kick them 
violently forwards, the distal parts of the third maxillipeds at 
the same time straightening so as to admit them to the outgoing 
stream, by which they are swept away. 


Explanation of Lettering of the Text-figures. 


te: endites or the primary segments | z., incisor process of mandible. 
which correspond to them. | i.1., inner lacinia of maxillule of 
9, additional segment in the max- Malacostraca. 
illiped of Calanus. ise.mir., 1schio-meropodite. 
17., 21., first and second lobes of maxilla | l., labrum. 
of Decapoda. L.’, side lobe of labrum. 
1lg., first leg of Leander. l.r., lateral row of sete. 
lmexp., 2mxp., 3 mxp., first, second, and m., molar process of mandible. 
third maxillipeds of Leander. m.yr., marginal row of sete. 
ap., apical lobe. mer., meropodite. 
a.r., third or additional row of setzx m., notch on paragnathum to 
on thoracic limb of Nebalia. receive endopodite of max- 
ax., axis of parapodium. illule. 
bas., basipodite. o.l., outer lacinia of maxillule of 
b.is., basi-ischiopodite. Malacostraca. 
b.m., base of mandible. p., paragnathum. 
car., et p.m., palp of mandible. 
cox., coxopodite. pr.cx. pYecoxa. 
dac., dactylopodite. pr.ep., proepipodite. 
d.c., dorsal cirrus. pro., propodite. 
en., endopodite. pro.dac., pro-dactylopodite. 
ep., epipodite. yr, 7.,r.’, ridges of segments of first leg 
ex., exopodite. of Leander, 
ji., flabellum. s.7., second side row of sete. 
g., gill of polychaete worm. s., subapical lobe. 
gl., papilla for opening of maxillary v.., ventral cirrus. 
gland of Stomatopoda. x., so-called ‘‘exopodite” of 
g’.; groove across paragnathum of Triarthrus. 


Leander. 


A 


Os 


ON THE SCOLEX OF A CESTODE. c 


5. On the Scolex in the Cestode Genus Duthiersia, and on 
the Species of that Genus. By Frank EK. Bepparp, 
MPAG 1 Sce:,cR RS... F.Z.8: 


[Received December 15, 1916: Read February 6, 1917. | 


(Text-figures 1-5.) 


INDEX. Page 
Duchiersia fimbriata Wiese... .0....cceieeeceies 79 
Dea paisa: VemierY ..c.5ichccke hives cdeuetemenaiy (SO 


The genus Duthiersia is fairly: well known to us from the 
investigations of authors whose several contributions are quoted 
later *, but there still remain a few points to which attention 
has not yet been directed, or concerning which there is up to the 
present some difference of opinion. 

Some little time since I had the opportunity of examining 
living examples of the genus from the Nilotic Monitor, which 
ene bled me to ascertain a structural feature which has escaped 
the attention of my predecessors —very probably because their 
investigations were made upon preserved material only. The 
scolices of several examples were in active movement, and from 
the apex of the scolex was seen to protrude a finger-like process 
which explored the surroundings. A more careful study of these 
living worms showed that the apex of the scolex is occupied by a 
circular pit, quite small like that of many species of /chthyotenia, 
which is apparently eversible. This pit lies between the upper 
extremities of the dorsal and ventral bothria, on a patch of 
integument which is not invaded by the bothria. ‘The area in 
question is more extensive than the pit which occupies its centre. 
Transverse sections confirmed the existence of this structure, 
which has not yet been described in the genus Duthiersia. The 
pit is so small that it only appeared in two sections of one series 
which I prepared, and only in five of another (thinner) series. 
And as these are naturally the very first sections of the series 
and very small in area, the apical pit might be easily missed, and 
possibly has been. 

The two series of sections referred to were transverse. I have 
also found the apical pit in horizontal sections through the scolex. 
In all of these it appears as a mere pit; certain special structures 
(text- fig. 1, s) were to be observed in the shape of delicate filaments 
arising from the margin of the pit, possibly of a sensory nature. 
As to the protrusion of the entire apex of the papilla, I believe it 
to consist of the tissues surrounding the pit as it was too large to 
be a mere eversion of that orifice. But possibly the pit was also 
everted, a fact of which I am not able to speak with certainty. 
The apical pit appears to me to be of a sensory nature, and thus 


* Page 75, footnotes. 


74 DR. F. E. BEDDARD ON THE 


perhaps does not bear any relation to the rostellum of other 
Cestodes. Related genera belonging to the same division of the 
Cestodes (i.e. Pseudophyllidea) throw no light upon this question. 
It is true that an apical depression has been described in other 
genera; but where this has been carefully investigated it would 
appear to be produced simply by an apical fusion—or nearly com- 
plete fusion—of the lateral bothria. Thus, in Bothriomonus*, the 
presence of a dividing septum shows that the apical vertical slit 
is merely the abbreviated remains of the two bothria. ‘Were the 
septum absent the homology of the depression in question might 
be more doubtful; I therefore believe this apical sensory (?) organ 
to be new to the Pseudophyllidea. 


Text-figure 1. 


A transverse section through the apex of the scolex of Duthiersia fimbriata. 


ce, cuticle; s, apical sense-organ. 


While I found this structure in examples of Duthiersia from 
Monitor niloticus, 1 examined other specimens of Duthiersia in 
vain. 

Two series of transverse sections of the scolex of specimens 
from Monitor bengalensis showed absolutely no trace of the organ. 
As these were much larger-scolices, the probability of my having 
failed to recognise the organ is thereby reduced. I have in fact 
little doubt that the apical pit is in those specimens quite un- 
developed. A comparison in other ways between the specimens 
from the two species of Monitors showed plainly that we have 
here to deal with two undoubtedly distinct species of Duthiersia. 


* Cholodkovsky, Annuaire Mus. Zool. de l’Acad. Imp. Sci. Petrograd, xix. 1914, 
p. 520, figs. 6, 7. 


SCOLEX OF A CESTODE. 7) 


This is not a novel conclusion; but it is not accepted by the. 
majority of recent writers, Perrier*, the original describer of: 
the genus, found differences in examples from different species of 
Momitor and recognised two species, viz. Duthiersia expansa from 
Eastern species of Monitor, and D. elegans from African. Perrier 
was perfectly right, and the majority of his successors are quite 
wrong. ‘The confusion of two distinct species 1s connected with 
various assertions with regard to the form of the bothria in 
this genus. It will be necessary to clear up this confusion. In 
D. expansa the form of the scolex is more spear-shaped than in 
the other species, as is plainly shown in Perriev’s figures. The 
bothrium on each side is closed posteriorly and opens again by a 
minute pore closely adpressed to the commencing strobila, thus 
producing a tube-shaped bothrium open widely in front and by 
but a narrow orifice posteriorly; this funnel-like arrangement 
has been justly compared by many to the tube-like bothria of 
Bothridium (Solenophorus). According to Perrier the same orifice 
exists posteriorly in D. elegans, but at some distance laterally 
from the fusion of the bothria with the strobila. Monticelli and 
Crety 7, who examined only examples of Duthiersia from an Indian 
Monitor, confirmed the existence of the posterior pore in that 
worm ; and, inferring its existence also in examples from JJonitor 
miloticus from Perrier’s statements, united both these worms into 
one species under the name of Duthiersia fimbriata; this name 
was given by Diesing{ to what he regarded as a species of 
Solenophorus, though tabulated as ‘‘ species inquirenda.” Diesing 
made his observations upon Perrier’s “species” D. elegans. Just 
previously to the memoir of Monticelli and Crety, Lithe$ took 
the opposite view and denied the posterior orifice of the bothrium, 
but agreed with the first mentioned authors in regarding the 
Cestodes from all species of J/onitor as belonging to one species 
only, namely (of course) D. fimbriata. This view is accepted 
by Braun |) in Bronn’s ‘ Thierreich,’ who, in defining the genus 
Duthiersia, described the hinder region of the bothrium as ‘“ nicht 
perforirt,” the italics being his own. It is true that in earlier 
numbers of the same volume Braun accepted Perrier’s statements 
and even used his figures, but later altered his opinion by reason 
of Liihe’s observations. Still later Shipley | re-asserted the 
existence of a posterior opening of the bothria in specimens from 
Monitor salvator and M. bengalensis, as did Southwell ** ‘ In 
Varanus spp.” The latter regards as synonyms both of Perriex’s 
species. Klaptocz 7, however, in 1906 again definitely denied the 
existence of the posterior orifice in the bothria of Duthiersia from 


* Arch. de Zool. Expér. ii. 1873, p. 349. 

+ Mem. R. Ace. Sci. Torino, (2) xli. 1891, p. 381. 

t Sitzungsb. Wien. Ak. xiii. 1854, p. 589. 

§ Verh. Deutsch. Zool. Ges. 1899. p. 48. 

|| Klassen u. Ordn. des Thierreichs, Vermes, Bd. iv. Abth. 1B. p. 1689. 
{| Spolia Zey]. i. 1903, p. 47. 
** Rec. Indian Mus. ix. pt. v. 1913, p. 281. 
tt Sitzungsb. Wien. Ak. cxv. 1906, p. 133. 


76 DR. F, E. BEDDARD ON THE 


Monitor niloticus. The latest statement known to me is that of 
Cholodkovsky *, who has defined Duthiersia by (inter alia) the 
fact that the “ Bothridia have the appearance of a funnel with 
blind narrow ends formed behind.” This definition presumably 
implies the existence of only one species of the genus. 

‘The above brief résume shows that, while Perrier examined 
examples of Duthiersia from both the Nilotic and the Indian 
species of Monitors and asserted the existence of the posterior 
orifice of the funnel formed by the partial coalescence of the two 
lips of the bothrium in both of these, subsequent observers based 
their results upon the personal examination of one only of the two 
alleged species; and also shows that there is a consensus of opinion 
that no posterior orifice exists in examples from Monitor wiloticus, 
while it is asserted to exist in examples from Indian Monitors. 
These observers finally appear, on the whole, to have concluded that 
their own observations, though made upon one set of individuals 
‘only (whether from Africa or India), applied to the others ex- 
amined by their fellow-workers, and that the genus Duthiersia 
was definitely to be characterised by the possession or non- 
possession of these orifices according to each observer’s own 
discovery of fact. I have made myself an examination, as already 
stated, of examples of Duthiersia from both African and Indian 
species of Monitor, and I cannot see why the obvious differences 
pointed out by Perrier nave not been universally accepted. To 
these I have some fresh observations to add which, as I think, 
entirely justify the position taken by Perrier. 

Perrier’s figures show the great difference in the general form 
of the scolex in the two species, which is, moreover, much larger 
in D. expansa than in D. fimbriata (as we must, of course, cail 
Perrier’s D. elegans). I have already described the apical pit in 
D. fimbriata, which is not to be found in D. expansa, and I agree 
with other observers that the posterior aperture of the bothria 
does not exist in D. fimbriata. I have examined several series of 
sections both transverse and longitudinal, and can find no trace 
of this orifice. In D. expansa, on the other hand, it is exceedingly 
obvious though very minute. It lies closely adpressed to the 
commencing strobila to which the posterior end of the folds 
forming the bothrium are attached, instead of, as in D. fimbriata, 
turning upwards to be attached at a point much higher up.and 
within the shelter of the lateral folds forming the bothrium. At 
the point of opening of the orifice the fused bothrial folds project 
laterally as a papilla upon the side of the strobila ; but the actual 
orifice is not upon the apex of this papilla, but upon its inner 
side. ‘There are other differences between the scolices of these 
two species of Duthiersia. In D. expansa, in tracing a series of 
sections from the strobila region forwards, the axis of the scolex 
is more sharply defined than in the other species. This is seen 
in transverse sections to be due to the fact that the flaps of 


* Trav. Soc. Imp. Nat. Petrograd, xlv. 1914, p. 62. 


SCOLEX OF A CESTODE. 77 


tissue which form the walls of the bothria do not unite at their 
attachment to the axis, but leave a space between their inner 
terminations. In D. fimbriata, on the other hand, the two walls 


Text-figure 2. 


. 4, 


Transverse sections through scolex of (left-hand figure) Duthiersia fimbriata 
and (right-hand figure) D. erpansa. 


w, walls of bothria; e, junction of these with the axis of the scofex. 


of the bothrium practically meet at their insertion (text-fig. 2, ¢). 
Thus, in both transverse and horizontal sections the axis assumes 


78 DR. F. E. BEDDARD ON THE 


a greater distinctness in D. expansa. This is also due to the fact 
that in the last-mentioned species the axis of the scolex 1s formed 
from the medulla only (text-fig. 3, a), while in D. fimbriata the 
axis 18 apparently formed from both medulla and cortical layer. 
The point of difference is further emphasised by the more modified 


Text-figure 3. 


,a 


Longitudinal section through scolex of Duthiersia expansa. 


a, axis of scolex continuous with medulla only of strobila region (m) ; c, cavity 
of bothrial groove displayed here and there; o posterior orifice of this 
cavity. 

@ 
structure of the axis in D. expansa, where it presents the appear- 
ance of a more clearly defined network, the spaces being largely 
quadrilateral in outline. There is not this plain histological 


ss anal 


SCOLEX OF A CESTODE. 79 


differentiation in J), fimbriata. Finally, in transverse sections 
through the bothrial canal up to its point of opening on to the 
side of the strobila, the same restriction of the bothrium to the 
cortical layer is to be seen very plainly; this is due to the fact 
that here the medulla is marked off from the cortex by a thick 
layer of longitudinal muscles which is itself sharply marked off 
both internally and externally. The tube of the bothrium 
traverses the cortical layer (text-fig. 4, d), only pushing back but 
not in any way taking up or invading these longitudinal muscles. 


Text-figure 4. 


cl m 


V 
Transverse section through posterior extremity of scolex of Duthiersia expansa. 


d-and v, dorsal and ventral bothria forming a narrow canal in the cortical layer ; 
the external orifice of v is shown; m, longitudinal muscular layer dividing 
the cortex from the medulla; 2, nerve-cord. 


Another difference between the scolices of the two species 
affects the water-vascular system. In both the scolex is per- 
meated by a network of these tubes which is very obvious in 
sections both transverse and longitudinal. J am not able to give 
a detailed account of the course of these vessels in the scolex, but 
it is quite clear that the number of tubes is much greater in the 
smaller species D. fimbriata, and that they are here of a smaller 
size than in the larger species D. expansa. We may now sum- 
marise the characters of the two species as follows :— 


Genus DuTHIERSIA, 


(1) D. Fimpriata Diesing. 


Solenophorus fimbriatus Diesing, SB. Ak. Wien, 1854, p. 589. 
Duthiersia elegans Perrier, Arch. Zool. Exp. 1873, p- 360. 


Scolex smaller ; bothria opening by continuous antero-lateral 


80 DR. F. E. BEDDARD ON THE 


groove only; apical pit at extremity of scolex; water-vascular 
system of scolex an abundant network of small tubes. 
Hab. Monitor niloticus, 


(2) D. expansa Perrier, 

Duthiersia expansa Perrier, Arch. Zool. Exp. 1873, p. 359. 

Scolex larger ; bothria opening by continuous antero-lateral 
groove and by separate posteriorly sitwated pore, being thus funnel- 
shaped ; apical pit not present ; water-vascular system of scolex a 
less abundant network of larger tubes. 

Hab. Monitor bengalensis and other Indian forms. 

It is quite possible that were these two species found in quite 
different hosts (7. e. of different genera or families) they would be 
placed in separate genera. ‘The differences of the scolex are 
obviously large and important as these differences go among the 
Pseudophyllidea. I do not, however, attempt this separation. 


In conclusion I desire to draw attention to a few minutie in 
the structure of the scolex of Duthiersia which have not been 
dwelt upon by those who have already studied the structure of 
this genus. The strobila near to the scolex is somewhat hour- 
glass-shaped in section, having a dorsal and ventral depression, 
and thus a bulging at the two sides; this is more marked in 
D. fimbriata than in the larger species. The medulla is separated 
from the cortical layer by a sharply marked band of longitudinal 
muscles which become frayed out and thus end—as a distinct and 
circumscribed layer—at the junction with the scolex. This layer 
is the same in both species. A transverse layer lying within this 
is to be seen in longitudinal section, but does not form a con- 
tinuous coating of muscular fibre: there is simply a slender 
bundle of fibres at the posterior end of each segment. This layer 
escaped my attention in D. expansa, where it cannot at any rate 
be so obvious as in the other Duthiersia. This state of affairs 
contrasts with what obtains in Solenophorus, believed to be closely 
allied to Duthiersia. In the former the longitudinal layer is very 
much thicker and with more scattered and at the same time 
larger fibres, and the extent of the medulla is reduced. In trans- 
verse sections the strobila of Solenophorus contrasts with that of 
Duthiersia by its stouter form and oval to circular outline. This 
thickening of the muscular layer in Solenophorus is, no doubt, 
connected with the strong muscular supply of the walls of the 
bothrial tubes in this genus. But in Duthiersia, in transverse 
section, a thinnish layer of fibres is seen to extend along the 
projecting walls of the bothria and represents the constricting 
muscles seen in Solenophorus, though diminished in importance. 
Within the bothrial tubes of Solenophorus the hypodermic cells 
(subcuticular layer) are covered by a structureless stained (by re- 
agents) and slightly opaque cuticle, outside of which is a clearer 
but still rather granular yellowish cuticle of chitinous appear- 
ance, of which the outermost layer is stained by reagents. In 


SCOLEX OF A CESTODE. 8] 


Duthiersia the layer which is thrown off most externally by the 
outermost layer in the bothrial groove is quite different. It is much 
deeper and greatly stained by reagents. It presents (text-fig. 5) 
the appearance of closely approximated plates, thinner towards 


Text-figure 5. 


Upper figure a section through a portion of the wall of the bothrial tube of 
Bothridium (Solenophorus) megacephala. 


Lower figure a similar section of Duthiersia fimbriata. 


1, membrane immediately lining bothrial groove and secreted by (e) epithelial lining : 
between the two lies another membrane, shown as a darker line. The difference 
of the outermost of the two membranes in Duthiersia and Bothridium 1s 
clearly shown. 


Proc. Zoou, Soc.—1917, No. VI. 6 


82 ON THE SCOLEX OF A CESTODE., 


the outside but reinforced by thicker bars. When the walls of 
the bothrium are closed upon each other there is absolute contact 
between the cuticular layers of the two sides. The appearance is 
totally different from what is to be seen in Solenophorus, and thus 
presents a striking difference between the two genera. It should 
be added that in Duthersia, as in Solenophorus, a second layer 
lies within the outer cuticular layer just described which is 
precisely like that of Solenophorus. 


ON BRITISH CISSID BEETLES. 83 


6.—1. The Coleoptera of the Family Cissidee found in 
Britain, with Descriptions of two new Species.— 
2. A new Species of the Coleopteran genus Cryptor- 
rhynchus Iiger. By C. J.C. Poon, Assistant Curator 
Caird Insect House *. 


[ Received Febraary 6, 1917: Read February 20, 1917.) 


INDEX. Page 
1. The Coleoptera of the Family Cisside found in Britain... 83 
Introduction «........ Se eee Sons Ser re Nn eaae of?) 
Table of Genera and Buetiks Gates 84 
Notes on Characters and Diccvation: en Description 
OEENE WAS POClOS a hr tacits come demt emi ates ada er earn ee OO 
2. A new Species of the Coleopteran Genus Cryptorrhynchus 
Introduction. 


It is well known that since at least the time of that eminent 
French entomologist the Abbé Latreille (1806), the Order 
Coleoptera has been divided into major groups superior to 
families, which have been based principally on affinities in the 
form and character of the antenne, or on the number of tarsal 
joints. 

These groups constituted, as they have been, differently by 
different authorities and known by different names, present 
anomalies difficult to reconcile or explain. 

The family Cisside Mellie (1848), which forms the subject of 
these notes, is in its morphology one of the most anomalous of 
these constituents. 

It was included by the British authority Marsham (1802) in 
the genus Ptinus L., and by Stephens (1839) in the family 
Bostrichide Leach, and close to Anobium F. 

Thus the family maintained its position in the works of sub- 
sequent British authorities paca the Teredilia or Serricornia, 
of which such genera formed part. 

In the latest general Kuropean list (Heyden, Reitter, and 
Weise, 1906), the Cissidee have been placed after the Myceto- 
phagide among the Clavicornia, and as that arrangement has 
been followed by,the most recent list of the British Coleoptera 
(Newbery and Sharp, 1915), L propose to adopt it here as well as 
the specific nomenclature of the family there used. 

The Cissidee are fungivorous, and may be found in various kinds 
of Boleti and Polypori on old trees and logs. 

Very few species are attached to any particular kind of fungus. 

A piece of Boletus from Godalming once produced no less than 


* Communicated by the SECRETARY. 


‘ , 6 


84 MR. C. J. C. POOL ON 


seven species, representing the four gone Crs, Ennearthron, 
Octotemnus, and Rhopalodontus. 

At first one experiences some difficulty in naming specimens 
owing to the confusion of species and the presence of many im- 
pertect or immature examples in the older collections available 
for study and comparison. 

Colour variation is a constant source of difficulty owing to the 
fact that it might be due either to simple aberration or to different 
degrees of maturity of the specimens. 

The colour of the clubs of the antenne, sometimes mentioned 
as a guide to the identification of species closely related, must 
not be accepted as a reliable character without allowance for 
variation. I have studied great numbers of living specimens, 
from emergence from the pupa to undoubted maturity, and the 
results of my investigations enable me to say in which of 
the species this character is regular, or the reverse. 

Any uncertainty or difficulty relating to immaturity may be 
avoided by breeding and keeping the insects alive for several 
months. 

The breeding-process is very simple, and consists of gathering 
infested fungi, which may be kept in carefully labelled tins or 
jars in a cool situation, where the insects may develop under 
healthy conditions. <A little damp blotting-paper or peat will 
supply all the necessary moisture, care being taken that sufficient 
ventilation is provided to prevent mildew. 

Most of the species are long-lived, and may be found in the 
larval or adult state at any season. Pupation appears to be 
unusual in winter, but sometimes occurs in sheltered situations. 

In the formation of the following table of the genus Cis, for 
which I ciaim little more than the merits of simplicity, I have 
to some extent grouped the species, as I have found them con- 
founded in collections. For instance, the series of C. micans in 
the National Collection at one time consisted of four different 
species, viz. :—C. nicans, C. setiger, C. boleti (small and imma- 
ture), and C. hispidus (immature): 

The general characters of shape and size will enable the student 
without figures or reference-collectious to decide to which of the 
five groups his specimens belong before attempting the more 
difficult task of specific identification. 


Table of Genera and Species. 
Genus Cis Latreille. 
Antenne 10-jointed. Anterior coxe transverse, round-oval ; 
tibize not (or very rarely) dilated at apex. 
Group 1.— Form short.. Almost glabrous. L. 13-13 mm. 


lineatocribratus Mell. Elytra strongly punctured in rows. 
Never black. Clubs of antenne always pale. 


BRITISH CISSID BEETLES, 85 


C. nitidus Herbst. Colour pale brown to black. Clubs of 
antenne always dark. Anterior angles of thorax 
produced, 

C. jacquemarti Mell. Colour pale brown. to black. Clubs of 
antenne always pale. Anterior angles of thorax not 
produced. 


Group 2.—Form short. Plainly pubescent. L. 13-2 mm. 


C. bilamellatus Wood ov Fowler ?=minutus Blackburn. Upper 
surface dull. Clubs of antenne dark. 

C. lineatosetosus, sp. n. Upper surface shiny. Clubs of antenne 
pale. 


Group 3.—Oblong. Rather broad. L. 2-4 mm. 


C. boleti Scop. Thorax uneven, irregular impressions on dise, 
base not bordered. Clubs of antenne pale testaceous or 
black. 

C. setiger Mell. =villosulus Marsh. Impressions on thorax more 
or less obsolete, base bordered. Clubs of antenne pale 
testaceous or black. 

C. micans F. Never quite black. Clubs of antenne always 
dark. 

C. hispidus Payk. Mature specimens always black. Clubs of 
antenne always pale red. 

C. bidentatus Ol. Dull. Scanty pubescence. 


Group 4.—Elongate. Large. L. 24-22 mm. 


C.alni Gyll. Shiny. Finely punctured. Very scantily pub- 
escent. Clubs of antenne pale or dark. Tibiz long and 
slender. 

C. latifrons, sp. n. Not so shiny. Coarsely punctured. Dis- 
tinctly pubescent. Head very broad. Clubs of antenne 
pale. Tibi short and stout. 

C. punctulatus Gyll. Dull. More coarsely punctured and with 
longer pubescence. Clubs of antenne pale or dark. 


Group 5.—Elongate. Size smaller. —L. 1-23 mm. 


C. oblongus Mell.=pygmeus Marsh. Black. Legs ved or partly 
e ved and black. Clubs of antenne variabie, pale or dark. 
Pubescence usually reddish, rarely yellow or white. 
Females sometimes much above average size of this 
| group. 

C. vestitus Mell. First ventral segment of abdomen of male 
bearing a small umbilicate depression in the middle 
(Newbery). Colour variable, but rarely black. Legs 
ferruginous, never red or black and red. Clubs of 
antenne pale or black. Upper surface dull. 


86 MR. C. J. Ce POOL ON 


C. festivus Panz. First ventral segment of abdomen of male 
rugose all over, but with no umbilicate depression in 
the middle (Newbery). Clubs of antenne always"pale. 
Upper surface shiny. 

C. castaneus Mell.= fuscatus Mell. Unicolorous castaneous, 
Depressed and dull. » Clubs of antenne never black. 


Genus Ruopautopontus Mellie. 


Antenne 10-jointed. Anterior coxe shorter (more or less 
conical); tibie dilated and denticulate at apex. 


R. perforatus Gyll. Size larger. Dull unicolorous. Pale or dark 
brown. Clubs of antennz never black. L. 17-2 mm. 

kh. fronticornis Panz. Size smaller. Dull black, with pale 
testaceous legs. Clubs of antenne sometimes black. 
L. 1-14 mm. 


Genus ENNEARTHRON Mellie. 


Antenne 9-jointed. 


E. affine Gyll. Colour black. Clubs of antenne alway dark. 
L, 1-13 mm. 

EF. cornutum Gyll. Colour brown. Clubs of antenne usually 
dark, rarely pale. L. 1-1? mm. 


Genus Ocroremnus Mellié. (Ovrophius Redtenbacher.) 


Antenne 8-jointed. 


O. glabriculus Gyll. Smooth, shiny. The number of joints of 
the antenne will distinguish it from Cis jacquemarti in 
Group 1. L. 1-14 mm. 


Notes on Characters and Distribution, with Description of 
New Species. 


Genus Cis Latreille. 


C. LINEATOCRIBRATUS Mell. . 
There is little or no variation noticeable amongst mature 
specimens. 


Previously only recorded from Scotland and Cumberland (Day). 
The latter record (Brit. Col., Fow. & Don. vol. vi. p. 281) refers 
to a single specimen taken from fungus on birch at Gt. Salkeld 
by Mr. H. Britten. Abundant in a large detached brown fungus 
at Mark Ash, New Forest, Sept. 1913 (Pool). Near Brockenhurst 
(Dr. Sharp), Denny Wood (Bedwell). 


BRITISH CISSID BEETLES. 87 


C. nrtipus Herbst. 

Common and widely distributed throughout Britain. 
~ Treland, common in Dublin and Belfast districts. 

It is found in Scotland in company with C. jacquemarti Mell., 
and is sometimes confounded with that species in collections. 


C. JACQUEMARTI Mell. 


Previously only recorded from Scotland, where it has been 
taken freely at Rannoch (Turner, Beare, and Denisthorpe), 
Garve in Ross-shire (Dr. Joy). Near Brockenhurst, 1915 
(Dr. Sharp), in company with C. nitidus Herbst. 

Not recorded from Ireland. 


C. BILAMELLATUS Wood = bilamellatus Fowler (Europ. List 
Heyden, Reitter & Weise, 1906) = minutus Blackburn. 

There is some considerable variation in size and development 
of the males. Small specimens occur without the upright plates 
on the thorax and clypeus, which might easily be mistaken for 
females or for members of another species. 

Additional localities : Orpington, Kent (Pool), Richmond Park 
(Donisthorpe & Perkins), Highgate (Janson). 

It has occurred at Port Lincoln, South Australia, and was 
named OC. miuutus by Blackburn, with whose type I have 
compared British specimens. 

Exceeding abundant in Kent and Surrey, but is probably an 
introduction like the following species. 


C. LINEATOSETOSUS, Sp. n. 


Short and broad, unicolorous testaceous, shiny. Head smooth, 
finely punctured and pubescent. Thorax finely punctured and 
pubescent, slightly narrowed in front, front margin with two 
indistinct teeth which merge with two other more distinct teeth 
on the clypeus when viewed from behind. Elytra twice as long 
as thorax, broad, closely punctured, especially near the scutellum, 
and clothed with ten straight rows of erect sete. Legs and 
antenne entirely pale testaceous. 

Length 14 mm. 

This insect resembles the small undeveloped males of C. bila- 
mellatus, which bear only slight traces of the plates on the thorax 
and clypeus. It is not an indigenous species, but because of its 
long residence in London and the possibility of its having become 
established in our parks or woods, it is desirable that its origin 
should be recorded with these notes on species found in Britain. 

Several specimens are in British collections, which I have 
traced to the following source :— 

‘* 86. In a fungus from the South Sea Islands that had been 
many years in Mus. Brit. (alive). From W. Carruthers, Esq., 
Sept. 1866.” 

I am indebted to Mr. J. N. Halbert of the National Museum, 


88 MR. C. J. GC. POOL ON 


Dublin, for this extract from Dr. McNab's notebook. It is of 
special interest as showing how (C. bilamellatus might have been 
imported from South Australia. 

My specimen, given to me by Mr. O. E. Janson from his 
father’s collection, bears a label with the following inscription :— 
“From Fungus in British Museum. Dr. McNab.” : 


C. BOLETI Scop. 

Exceedingly variable in size, colour, and punctuation of mature 
‘specimens. 

A specimen from Sandown, which lived for five months, is 
entirely pale testaceous. : 

Pale examples oecur with black clubs on the antenne. Dark 
brown or black specimens may be found with either dark or 
pale clubs. 

Rev. W. W. Fowler (Col. Brit. Is. p. 206, vol. iv.) says :— 

“The Cis rugulosus of Mellié, which was introduced into our 
lists by My. Crotch, appears to be only a variety of C. boleti in 
which the rugose punctation of the elytra is more apparent and 
the larger punctures more or less obsolete ; as intermediate varia- 
tions occur it can hardly be regarded as a stable variety, much 
less as a species.” 

My series of this and other species of Cis contains some 
remarkable aberrations, which I have refrained from naming 
as I consider the latter part of the remarks quoted might apply 
equally well to any of them. 

Common in fungus on logs, stumps, ete., throughout the 


Kingdom. 
C. sErrIGER Mell. = villosudiws Marsh. 


Probably the most variable species of the genus in size and 
colour. 

I possess specimens displaying the following combinations :— 
Body black with black clubs of antennew; dark brown with pale 
clubs ; pale testaceous with black clubs; and unicolorous 
testaceous. 

Recorded from Lancashire, Cheshire, and Suffolk. Common 
in Southern Counties in ecmpany with C, boleti. Exceedingly 
abundant in the [sle of Wight. 

Not recorded from Scotland or Ireland. 


C. MICANS F. 


No variation observable amongst mature specimens. 

Recent records of -localities:—Oxford and Chatham districts 
(Walker), Newbury (Harwood), Cumberland (Britten), New 
Forest (Dr. Sharp), Burnham Beeches, Bucks, and Fittleworth, 
Sussex (W. E. Sharp), Godalming, Surrey, bred in abundance 
from Loletus, which was also inhabited by C. boleti and C. setigey 
(Pool), Penarth (Tomlin). 

Not recorded from Scotland or Iveland.., 


BRITISH CISSID BEETLES. 89 


C. nisprpus Payk. 


Pubescence usually bright reddish, but I have seen specimens 
with yellow or white pubescence. 

Common and widely distributed throughout England. 

J have never seen it in the Isle of Wight. 
. Taken in Scotland at Nethy Bridge (Beate) and Garve ih Ross- 
shire (Dr. Joy). 

Ireland: Maryborough (W. E. Sharp). 


C, BIDENTATUS Ol. 

This species is sometimes abundant in a large fungus on old 
elms, which also produces Dacne rujfifrons and Mycetophagus 
multipunctatus. 

Common and widely distributed thr preter England. 

Scotland: Rannoch, in an old Polyporus on birch, also in- 
habited by C. nitidus, C. gacquemarti, and Lolitophagus reticulatus 
L. (Donisthorpe). 

Iveland: Armagh and Dublin. 

I have not seen it in the Isle of Wight. 


C. penratus Mell. 


Mr. Donisthorpe has withdrawn this species from the British 
list (Ent. Record, vol. xxviii. p. 155, 1916). 


©. ALNI Gyll. 


The specimen recorded as C. dentatus Mell. (Ent. Record, 
vol. xix. p. 136, 1907) is a curious aberration of C. alni with 
abnormally coarse punctation of the thorax. 

There appears to have been some interruption of the pigment 
which gives the insect a strange greenish opaque appearance, 
only the seutellum having the normal dark brown colour. 

Whilst withdrawing C. dentatus from our list, Mr. Donisthorpe 
proposes the name of var. mitfordi for this aberration of C. alii. 
For reasons already stated in connection with C. boleti, I am not 
following Mr. Donisthorpe in this direction. 

This species appears to be common in most districts where 
elders are growing.’ It occurs under the bark of dead elder- 
stems upon which the curious black ‘“Jew’s ear” fungus is 
growing. It has been recorded in fungus on oak in Dunham 
Park, Manchester, and Professor Beare tells me he has taken it 
plentifully i in fungus on dead birch boughs at Nethy Bridge in 
Scotland. 

Calbourne, I. of Wight (Morley), Sandown (Mitford). 

There appears to be only one Irish record, from Mote Park, 
Roscommon. 


C. LATIFRONS, Sp. n. 


Black, elongate, parallel-sided, clothed with white outstanding 
pubescence, which, when examined under a lens, is very distinct 


90 me 
on the sides of the thorax and elytra. Head with eyes nearly as 
broad as the thorax, finely punctured and pubescent. Thorax 
and elytra coarsely but evenly punctured. There is an im- 
punctate line behind the middle of thorax equal in width to the 
space occupied by two of the surrounding punctures. Antenne 
testaceous ; clubs slightly darker. Legs red. Tibie short and 
stout. 

Length 14-23 mm. 

This species comes near to C. alni, with which it has been 
confounded in collections. The longer pubescence, coarser 
punctation, as well as its shorter and stouter tibize, will easily 
distinguish it from that species. 

I have recently taken two specimens from rotten beech to 
which is attached a small brown fungus containing Cis larve, 
collected with other fungi at Lyndhurst in September 1916. 
There is a specimen in the Power Collection from the New 
Forest, and another in the Waterhouse Collection taken in the’ 
same locality by Mr. Kemp. Another example without data is 
in the collection of Mr. H. Willoughby Ellis. It was along 
with some common New Forest species, so is probably from 
that district. 


C. POOL ON 


C. puncruLatus Gyll. 


This is now the only undoubted British species of Cis which 
has not been recorded from the south of England. Orton Woods, 
nr. Carlisle (Day & Britten). 

Scotland: Braemar, Aviemore, Nethy Bridge, Balmuto, Ran- 
noch and Peebles. Professor Beare tells me the species lives in a 
white fungus which develops between the wood and the bark of 
dead Scotch fir. 

Not recorded from Ireland. 


C. optoneus Mell.=pygmeus Marsh. 


Forest Hill, Dulwich, Horsell, Coombe Wood, Tonbridge, 
Windsor, Reading ; Knowle, Warwickshire. 

I have bred it in profusion from fungi gathered from oak posts, 
at Sandown and Brading, I. of Wight. 

Chatham (J. J. Walker), Oxford (Collins), North Holt, Mid- 
dlesex (W. E. Sharp), Brockenhurst (Dr. Sharp), Enfield and 
Epping Forest (Pool), Harlech (Donisthorpe). Exminster, June 
1909. Abundant on old posts, Bovey (P. de la Garde). 

Not recorded from Scotland or Ireland. 

I have never found it accompanied by any other species of Cis 
and have never seen the imago alive in winter. It is an active 
creature in June and July, when I have-seen it in numbers 
running in hot sunshine upon posts and tree-trunks infested 
with the fungus in which it feeds. 


BRITISH CISSID BEETLES 91 


C. vestirus Mell. 

A variable species, of which I have seen the following 
combinations :— 

Body black, with pale testaceous clubs of antenne, Wallington 
(Power). Black, with black clubs; pale testaceous, with black 
clubs; as well as unicolorous brown and smoky-yellow forms. 
I have never found C. vestitus and C. festivus inhabiting the 
same fungus together. 

Forest Hill, Dulwich; Olton and Sutton Park, Staffs. ; Man- 
chester district; Teesdale; Epping Forest (Beare). Richmond 
Park (Donisthorpe). 

I have bred it in plenty from fungus scraped from the under- 
side of dead oak boughs from Enfield, Edmonton, Brockenhurst, 
and Sandown, I. of Wight. Widely distributed and not un- 
common, but often overlooked or mistaken for C. festivus, with 
which I have seen it confounded in the Power and _ other 
collections. 

Not recorded from Scotland or Ireland. 


CG. FESTIVUS Panz. 


Clubs of antennz always pale. Little or no variation in 
mature specimens. | 

Common and widely distributed in England and in I. of Wight. 
Usually found in fungi on oak posts or branches.. Prof. Beare 
finds it every year with C’. alni, in fungus on birch branches or 
faggots at Nethy Bridge. I have bred it in plenty from fungi 
from New Forest, Godalming, and Portsmouth district, but never 
in company with any other Cis. 

Ireland (Boris); Queenstown (J. J. Walker). 


C. castanEus Mell.=fuscatus Mell. . 


The exchange list of British Coleoptera, Newbery & Sharp, 
1915, shows the type-form of this species as not yet recorded 
from Britain. 

This form actually represents the mature insect, which I have 
bred in great numbers from fungi from Enfield, Epping Forest, 
New Forest, Warlies Park, Essex, and Sherwood Forest. Rich- 
mond Park (Donisthorpe); Symonds Yat, Cardiff Cannock Chase, 
and Liverpool in a flour mill (‘Tomlin). 

Immature specimens were called C. fuscatus by Mellié = 
fuscatus Mell. = var. fuscatus Newbery & Sharp, nec Mell. 


C, ELONGATULUS Gyll. 


First mentioned as British by Dr. Sharp (Ent. Mo. Mag. 
vol. vil. p. 83, 1871), who says Mr. Crotch considers he has 
Scotch examples of this species. 

Dr. Sharp does not mention having seen the specimens. 

There is a single specimen of a Cis in the Crotch Collection at 
the University Museum, Cambridge, standing in the name of 


92 MR. C. J. C. POOL ON 


C. elongatulus. Unfortunately it has lost all of its legs and 
antenne and is without data. It may be one of Mr. Crotch’s 
Scotch examples, but I have failed to trace the others in any 
other collection. 

The only other record of the species is Knowle, Wawickshire 
(Blatch) (Cat. Brit. Is., Fowler, vol. iv. p. 209). This is an error, 
as there is not even a label for elonyatulus in the Blatch cabinet. 

The position of this species in the British List depends entirely 
upon a single imperfect specimen of doubtful origin, and it pro- 
bably remains for collectors in the north to prove if the Scotch 
record is justified. 

Professor 'T. Hudson Beare tells me he has never found any 
specimen in Scotland which in any way agreed with the descrip- 
tion of this species. 

[I am indebted to Mr. H. Willoughby Ellis for allowing me to 
study the Cisside in the Blatch Collection, and to Mr. Hugh Scott 
for the loan of the Crotch specimen for examination. 


_ Genus RHOPALODONTUS Mellie. 


R. PERFoRAtTUS Gyll, 


There are not any recent records of this insect, which has only 
occurred at Rannoch in Perthshire. Bred with Cis nitidus, from 
a hard Boletus found on old birch trees in the Black Forest 
(Foxcroft, 1853-4). The last record was Rannoch (‘Lurner, 
1858). 


R. FRONTICORNIS Panz. 


IT have bred this species in plenty from fungi on elm and 
willow and beech from Epping Forest, Enfield, Cheshunt, 
Godalming, and have taken it in small numbers at Brockenhurst 
and Orpington, Kent. 

Bovey Tracey (P. de la Garde, 1909, fungus on saliows); Oxford 
district (J. J. Walker); Kerne Bridge, Newbury (Tomlin). 

Not recorded from Scotland or Ireland. 


Genus ENNEARTHRON Mellie. 

K. AFFINE Gyll. 

I have taken this insect at Brockenhurst, Sherwood, Epping 
Forest, Sandown, I. of Wight, Millwall Dock in fungus on aspen 
lbg with C. setiger and Dermestes frischi. 

Windsor, Ashdown Forest (W. E. Sharp); Sheppey, Oxford, 
and Chatham (J. J. Walker); Teesdale, Durham (Bagnall). 

‘ Not recorded from Scotland or Treland. 


EK. cornutum Gyll. 


Ditchingham, Norfolk (Beare), Ashdown Forest and Crow- 
thorne (W. E. Sharp), nr. Exeter (Rendel). I have not seen 


A NEW BRITISH BEETLE, 93 


it in the Isle of Wight. Bred in numbers from’various fungi 
from Portsea Island, Horndean, Hants, Sherwood Forest, New 
Forest, Epping Forest, Warlies Park, Essex, Enfield, Orpington, 
Kent (Pool). 

Not recorded from Scotland or Ireland. 


Genus Ocroremnus Mellie. 


O. GLABRICULUS Gyll. 


No variation observable amongst Mature specimens. 
I have seen pale immature specimens in an old collection under 


the name of Cis flavus Kirby. 
Abundant everywhere in Boletus on various trees. 


2. A new Species of the 
Coleupteran Genus Cryptorrhynchus Uliger. 


C. HARRISONI, sp. n. 


Black, rostrum stout, slightly narrowed in centre, almost 
glabrous shiny and red in front; centre sparingly and_ base 
thickly clothed with elongate white scales. Antenne red and 
shiny. Thorax rounded at sides, narrowed at apex, coarsely 
and closely punctured, sparingly clothed with white scales. 
Scutellum dull black, finely punctured. Elytra densely clothed 
at base and apex with white and yellow scales, and more 
sparingly with similar seales in the centre and marked with 
straight rows of large shallow punctures, smooth between rows, 
very finely punctured near suture. Legs dark reddish, clothed 
with elongate white scales; femora with a distinct tooth; 
tarsi red. 

Leneth 3 mm. 

A single specimen was taken by Mr. F. A. Harrison at Frinton- 
on-Sea, Essex, during the summer of 1915. Unfortunately 
Mr. Harrison does not remember the circumstances of its 
capture. 

It may be an importation, but, like Catharmiocerus maritimus 
and other exceedingly local coast weevils, it might easily have 
been overlooked, especially as the locality appears to have pro- 
vided but few attractions for British collectors. 


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THE SECRETARY ON ADDITIONS TO THE MENAGERIE, 95 


EXHIBITIONS AND NOTICES. 


February 6th, 1917. 


Prof. KE. W. MacBripg, D.Sc., F.R.S., Vice-President, 
in the Chair. 


The Srecrerary read the following report on the Additions 
made to the Society’s Menagerie during the months of November 
and December 1916 :— 


NOVEMBER. 


The registered additions to the Society’s Menagerie during the 
month of November were 60 in number. Of these 50 were 
acquired by presentation, 8 were received on deposit, and 2 by 
purchase. 

The number of departures during the same period, by death 
and removals, was 115. ; 

Amongst the additions special attention may be directed 
UOn— 

2 Grizzly Bears (Ursus horribilis), from North America, 
deposited on November 21st. 


DECEMBER, 


The registered additions to the Society’s Menagerie during the 
month of December were 27 in number. Of these 15 were 
acquired by presentation, and 12 were received on deposit. 

The number of departures during the same period, by deaths 
and removals, was 120. 

Amongst the additions special attention may be directed 
UO} 
1 Sing-Sing Waterbuck (Cobus defassa) 3, from W. Africa, 
deposited on December 8th. 

1 Kashmir Deer (Cervus hanglu) 2, from Kashmir, presented 
by H.G. The Duke of Bedford, K.G. 


Mr. EDWARD GERRARD, on behalf of Mr. Crabb, exhibited a 
mounted specimen of a bird bred and reared in captivity under 
conditions in which it seemed certain that it was a hybrid 
between a male Thrush (Zurdus musicus) and a female Black- 
bird (Merula merula). 


96 MR. C. J. C. POOL ON THE 


February 20th, 1917. 


Dr. A. Surra Woopwarp, F.R.S., Vice-President, 
in the Chair. 


The Srecrerary read the following report on the Additions 
made to the Society’s Menagerie during the month of January 
1917 :— 

The registered additions to the Society's Menagerie during the 
month of January were 118 in number. Of these 49 were 
acquired by presentation, 58 were received on deposit, 5 by 
purchase, 4 in exchange, and 2 were born in the Gardens. 

The number of departures during the same period, by death 
and removals, was 128. 

Amongst the additions special attention may be directed 
tO— 

1 White-bellied Hedgehog (Frinaceus albiventris), new to the 
Collection, from Dakar, Senegal, presented by Mrs. C. H. A. 
Reaney on January 20th. 

1 Greater Double-collared Sunbird (Cinnyris afer), new to the 
Collection, from South Africa, presented by Alfred Ezra, F.Z.5., 
on January 4th. ‘ 

1 Three-lined Snake (Zvimerorhinus triteniatus), new to the 
Collection, from Nakuro, British East Africa, presented by R. 
Holmes on January 19th. 

12 Beautiful Tree-Frogs (Hyla pulchella), new to the Collection, 
from Cordova, Argentine, presented by W. A. Smithers, C.M.Z.S.. 
on January 29th. 


—_—- —— 


Mr. D. Seru-Smiru, F.Z.8., Curator of Birds, exhibited the 
trachea from a male Anseranas semipalmata, showing its extra- 
ordinary convolution, which had been described by Latham so 
long ago as 1797. The exhibitor stated that neither in this 
species nor in Phonygama keraudreni, another species with a 
greatly developed convoluted trachea, did the birds produce any 
great volume of sound,,such as might be expected from so 
elaborate an arrangement. 


Notes from the Caird Insect House. 


Mr. C. J. C. Poon, Assistant Curator of Insects, read the 
following notes upon species bred and exhibited :—_ 


LEPIDOPTERA. 


The Magpie Moth (Abraxas grossulariata). 


Vast numbers of larvee were collected in North London district 


CAIRD INSECT HOUSE, 97 | 


and turned out upon Huonymus plants. This species is a good 
exhibit as it is visible in all stages. 

A long series of varieties have been preserved and are now 
exhibited. he larve display considerable variation of colour, 
some of them being almost completely black. I was advised to 
isolate these melanic forms with the idea of producing some 
unusually dark imagos. I selected 73 melanic larvae, but the 
results were disappointing, and I found the adults merely, 
varied like those bred from unselected larve. 


ODONATA. 


Dragen-flies in the earlier stages have done well in the tanks. 
The following species have been bred :—#schna grandis, 4. jun- 
cea, Brachytron pratense, Libellula quadrimaculata, Orthetrwm 
cerulescens, and Agrion puella. Specimens have been preserved, 
together with the nymph skms from which they emerged. 

We have no difficulty in rearing these creatures, as they feed 
upon other insects bred in the tanks; our trouble begins with 
the emergence of the Dragon-fly, which is a very fastidious 
feeder and will only take insects in active flight in hot 
sunshine. 

The right conditions are obtainable only in an outdoor en- 
closure, which we hope to provide later, to exhibit these creatures 
during their natural adult life, which varies from one to three 
months. 


COLEOPTERA. 


Teratological specimens. 
Tetropium gabrieli. 


Some larch logs were received from Sutton Park, Staffordshire, 
infested with Jarvee of this species. 

One adult specimen, which emerged im the house, has mal- 
formed antenne, the right member of the pair baving an 
additional branch consisting of five joints. 


Donacia linearis. 


Specimens were collected at Waltham Abbey and exhibited on 
reeds in a shallow tank. . 

One specimen has ten instead of eleven joints to the antenne, 
the eighth joint on each side being almost equal in length to 
joints 8 and 9 of a normal specimen. 


Bananas as Food for beetles. 


The problem of providing suitable food for various species of 
Coleoptera, whose ordinary feeding-habits are as different as 


Proc. Zoot. Soc.—1917, No. VII. a 


98 MR. C. J. C. POOL ON THE 


those of the lion and the lamb, has been solved in the Insect 
House by using the banana. 

The following list of beetles and their ordinary foods, from 
which, under natural conditions, some of these insects are rarely, 
if ever, found very far distant, will form an interesting record of 
species which during the years 1915-1916 have been found to 
thrive partly or wholly upon this fruit :— 


Voracious ground-beetles. 


Carabus violaceus. 


BS nemoralis. 

glabratus. 

as auratus. 
catenulatus. 


9? 
Calosoma inguisitor. 
Pierostichus vulgaris, 
ater. 

S madidus. 
Harpalus ruficornis. 


99 


Aquatic species. 


Hydrophilus piceus. 


Wood-boring species. 


Dorcus parallelopipedus. 


Saperda carcharias. 


Clytus mysticus. 

» arietes. 
Aromia moschata. 
fylobius abietis. 
fTylesinus crenatus. 
Scolytus destructor. 

>» tntricatus, 
EKlater pomone. 


Dung-feeding species. 


Geotrupes spiniger. 
stercorarwus. 


29 
# sylvaticus. 
‘; typhaus. 


Plant-feeding species. 
Chrysomelu fastwosa. 
grams. 


hyperice. 


9? 


9) 


Ordinary Food. 


Usually other insects. 


” ” ” 
” ” ” 
” ” ” 
” ” ” 
9 re) oe) 
’ ” ” 
” ” ” 
” ” ” 
oe) ” ” 


General scavenger, vegetable or animal 
matter, 


Moist rotten wood (comes to sugar). 

Foliage and bark of young stems of 
poplar. 

Hawthorn blossom. 


85 

Flowers ; sap and foliage of willow. 

Fir bark. 

Ash bark. 

Elm bark. 

Oak bark. 

Flowers in summer (larva in decayed 
beech). 


Horse-dung or decayed fungi. 


Rabbit-dung. 


Leaves of sallow. 
Leaves of sallow and wild mint. : 
Leaves and flowers of hypericum. 


CAIRD INSECT HOUSE. 99 


Adult Insects which do not feed. 


It is well known that many species of Lepidoptera, of which 
the Atlas Moth of India is a good example, are incapable of 
feeding in the adult state. The larva, when full fed, having 
enclosed itself in a protective cocoon and changed to the pupa, 
has provided itself with sufficient nourishment to sustain it over 
the very often far distant period of short, 1f energetic, adult life. 

The length of life varies according to temperature, which if 
high will produce great nocturnal activity, quickly exhausting 
the energy of the insect. Prof..H. M. Lefroy says these creatures 
usually become a complete wreck in three nights in the tropics. 
I have frequently kept them alive and in almost perfect con- 
dition for from 10 to 16 days by transferring them to a cooler 
case immediately after emergence from the cocoon. 

Little has ever been written about the feeding-habits of 
Coleoptera belonging to the Longicorn group, often “referred to 
as wood-feeders, which is generally only” correct as regards the 
lavve. 

I have studied these beetles alive, both in the field and in 
captivity, and 16 is my opinion that in their feeding-habits as 
adults we must regard them under two headings, viz., those 
which feed upon flowers or foliage of various plants and trees 
and those which, like the Atlas Moths, do not feed at all. 

Those which feed may be seen in active flight or settled upon 
their food-plants in hot sunshine 

The following list includes the nocturnal species teste@in the 
Insect House, and which I have failed to attract with any of 
the foods which appeal to the diurnal species :— 


Prionus coriarius. 
Asemum striatum. 
Tetropium gabrieli. 
Criocephalus polonicus. 


” rUsticus. 


Haperiments with living Specimens of the Sexton Beetles 
Necrophorus humator (black) and N. ruspator (ved and black) . 


Offered as food to the following animals in the small Mammal 
house —— 


Egyptian Mongoose.—Seized and devoured both species without 
hesitation. 


8S. American Coati.—Lost sight of the first beetle, but after- 
wards sniffed and followed its trail right across the cage. It 
then picked up the beetle.in its mouth, and, after spitting it out 
several tines, finally devoured it, 


100 MR. R. TIT. POCOCK ON THE WORK OF 


e 


The White Marmoset.—This little animal never vefuses an 
inset, but there was some hesitation about attacking these foul- 
smelling beetles. However, its love of insect-fool overcame 


everything else, and after aen sniffing and spitting two were 
devour ed. 


The Meercats.—Smelt, but would not touch, either species. 
T often feed these animals with such species as Dytiscws mar- 
ginalis, Carabus violaczus, and Cleotrupes sylvaticus. 


Crpuchin Monkeys.—'These animals seized both species readily. 
One Pea monkey satisfied herself with eating some of the 
numerous parasitic mites from the body of the beetle. A male 
monkey ate three beetles, and then stole and devoured the one 
held by the female. 


. 


———— ———— = 


March 6th, 1917. 


Dr. S. F. Harmer, Sc.D., F.R.S., Vice-President, 
in the Chair. 


Work of the Beavers in the Soctety’s Gardens. 


Mr. R. I. Pococs, F.B.S., F.Z.S., Curator of Mammals, gave 
an exhibition “heise by lamter n- des to show the work a the 
Beavew® in the old enclosure in the Gardens, and remarked :— 

“ Until the middle of 1913 the pair of Canadian Beavers had 
shown no special activities of any kind, apart from felling in 
characteristic manner the tree-trunks erected for that purpose. 
In the summer of 1913, however, a shallow pool appeared in the 
surface-clay of their enclosure. This they proceeded to deepen and 
widen by digging out the bottom and sides. They carried away 
the excavated clay and plastered it in different places, especially 
up against the fence. From the enlarged pool they drove an open 
channel through the soil for about a couple of yards; but this 
they soon roofed over with small logs and straw plastered with 
mud, so that their subsequent operations were concealed from 
view. But from the end of the channel and continuous with it, 
as it afterwards transpired, they dug out a burrow nearly three 
yards in length and passing under the root of a witch elm. The 
first intimation we had of this was the appearance of a hole, 
smaller than a rabbit’s burrow, close to the fence on the side 
remote from the pool where the channel started. ‘This hole they 
built up with tufts of straw and a small log or two, possibly to 
prevent the clay falling in. / 

When I opened ane burrow to satisfy myself, that it was not 
being carried beneath the fence, it appeared to me that the hole 


BEAVERS IN THE SOCIETY’S GARDENS. 101 


at the end was a ventilator. At all events no attempt had been 
male to enlarge it for exit or entry, and the part of the burrow 
it Opened into was a comparatively large chamber, wide enough 
for the beavers to turn about in and deep enough for them to 
immerse themselves under the water. The narrower part of the 


Text-figure 1. 


AY 
DRY 1 
( 
IPORTION G Ses 
y gr ae _H \ 
y gu yNo-n me - E PY ale EN 


Seat 


LANDING 
PLACE., 


Plan of the old enclosure for Beavers, showing the work done by the 
animals. 


burrow was from 13 to 2 ft. wide and about 2 ft. deep from the 
surface of the ground, the soil of its roof being only a few inches 
thick. It was rather more than half full of water, continuous 
with that of the pool at the entrance, which we subsequently 
found proceeded from a burst underground pipe. 


102 MR. R. I. POCOCK ON THE WORK OF 


In the evening of the day on which the burrow was opened 
one of the beavers started to repair the damage done, using 
straw, branches, and mud for the purpose. The task occupied 
him off and on the whole of the day following. He dug up the 
wet clay from the bottom of the burrow and plastered it with his 
paws over the straw and branches which he had previously laid in 
place to roof the open channel. 

From that date until the summer of 1916 the burrow was left 
undisturbed and two litters were born in it. The beavers, how- 
ever, were never idle. Working from the pool, at the entrance of 
the burrow, they dug out a ch: vaael | in an opposite direction from 
the first, but roughly in the same line for a few yards, then 
turned it with a sharp bend towards the artificial pond in the 
centre of the enclosure. This channel was roofed over in the 
same way as the first. In the meantime they had excavated a 
trench up against the concrete wall of this pong on the side where 
the burrow was situated. and the roofed-in channel was continued 
till it communicated with this trench, which was deep enough for 
the beavers to swim in except at one or two places on the margin 
where shallow landing-places were made, 

So long as the beavers were pa with straw, branches, and 
logs, they employed them continually for str enothening the roof 
of the burrow or heightening the roof of the terminal portion up 
against the fence. When the enclosure was done away with in 
1916, this portion where the burrow ended was found to consist 
of a comparatively large chamber partly filled with water; but ats 
the very end towards the trunk of the witch elm above referred 
to, there was a flat dry platform above the level of the water and 
about large enough to accommodate two beavers and their young. 
There was no bedding on this platform, merely a scanty covering 
of wood-chips. This part of the burrow was ventilated through 
chinks in the fence left open by the beavers, and from it a narrow 
passage, ending blindly, ran in the divection of the trench, and 
would probably have been carried through to it, giving the 
burrow a second entrance. 

The main points in the str eal of the burrow are shown in 

the plan of the enclosure (fig. 1, p. 101). The circular central 
area is the brick-house eae by the concrete pond, and the 
mud path is a beaten track along which the beavers used to carry 
the mud to pile up against the fence. 

I have watched the beavers at work night after night, and 
can confirm the observation of others as to their methods and 
activities. They carry mud in their fore-paws tucked up against 
the throat and chin and walk the while flat-footed on ea hind 
feet, with a kind of awkward waddle, the tail trailing behind and 
acting as a support and balance. Mud is laid on with the fore- 
paws to fill up chinks between branches and left to dry. It is 
never smeared over or pressed down with the tail, but sometimes 
it is jammed tightly home with the muzzle. 


. 


BEAVERS IN THE SOCIETY'S GARDENS. 103 


In the warmer months of the year the beavers seem to be 
pervaded with a restless energy, driving them to work with the 
materials at hand, whether there was anything to achieve or not. 
Their industry is certainly wonderful; but I was equally struck 
by the amount of absolutely aimless work they put in. Time and 
again I have seen a beaver, with infinite labour, haul a log out of 
the pond, pull it over the rough ground up to the top of the 
burrow and lay it there; then seizing it again, tug it back to 
the spot he started from. I have never seen any indication of 
intelligent co-operation between two beavers in shitting a log 
too heavy for one. If two get hold of the same log they do not 
work together and haul it in one direction, but tug and push in 
such a way that the efforts of the one counteract those of the 
other.” 


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aera biaded. to « Orked ax pend Ghat priiartss 
instedt Of as it the Selachii, etl graduaily wequiling ese 
approxy nately in the line peels: ed of the parachordals.. VPiy 
chondp6tranium of Phiten ts these earl ly stages has: rag) 
tunatery net been described. [I®hhas, however, been deg 
these stay res Th certain of @he Jelhehi 11, and af sligl| tly | 
in Callor ? ms os ‘ang gg eiees must be mqde-to t ié ‘tii 
s| befor 


Tn PS athias and 


lence probably i infall 
ag achii, the cranial g 


at when t th¢ Seo es 
begins to chondrify Phat mertzoffs (Lad97 figures 
af embryos of these fhe and the enclosed 
notochord project slightly into i hollow .gf the mesocephalic 
flexure (plica ence phali ventri von Kupffer, 1906). The 
trabecule cannot, agtor ling r; aad dowK in the line pr 
longed of gre paracghordn is SUA, _presq igtheNhoormal relations 
to the vefttral syfface of thesaraie’ they at fNxt | 
angles to § thordats end slightly posterior to 
ends. ‘The marked cranial flexufe at this stage al 
agg prosvisceretp@the¥ tla lie anterior: 
arch, and“the palatoquadrete and mangibula, whiph 
either the cartilaginous bar of the mandibular 


ne wh ae 


2 ee a oe ae . iy - ne . : : . Fe 
Stites ERP A with paatts of the SHS ost OUSPOr MrIOVe fat 
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ON THE SKULL OF CHIMERA. 105 


PAPERS. 


7. The Prechordal Portion of the Chondrocranium§ of 
Chimera collie. By Epwarp PueLps ALLIs, Jun., 
E.Z.S 

[ Received December 27, 1916; Read March 20, 1917.) 
(Plates I.-ITI.*) 


INDEX. 
Page 
Structure or Morphology ......5..02..000 fi. cco ses 10D 


The prechordal portion of the chondrocranium of the adult 
Chimera is directed ventro-anteriorly, and not anteriorly as 
Hubrecht’s (1877) figure of this fish would lead one to suppose. 
This is unquestionably due to the trabecule having been first 
Jaid down at a marked angle to the parachordals, and to their 
having retained, to a marked extent, that primitive position, 
instead of, as in the Selachii, later gradually acquiring a position 
approximately in the line prolonged of the parachordals. ‘The 
chondrocranium of Chimera at these early stages has unfor- 
tunately not been described. It has, however, been described at 
these stages in certain of the Selachii, and at slightly later stages 
in Callorhynchus, and reference must be made to the conditions 
in these embryos of these fishes before describing those in the 
adult Chimera. 


SELACHITI. 


In Acanthias and Pristiurus, and hence probably in all of the 
Selachu, the cranial flexure is so great when the neurocranium 
begins to chondrify that, as shown in Sewertzoff’s (1899) figures 
of embryos of these fishes, the parachordals and the enclosed 
notochord project shghtly into the hollow of the mesocephalic 
flexure (plica encephali ventralis, von Kupffer, 1906). The 
trabecule cannot, accordingly, be laid down in the line pro- 
longed of the parachordals, and, preserving their normal relations 
to the ventral surface of the brain, they at first lie at right 
angles to the parachordals and slightly posterior to their anterior 
ends. ‘The marked cranial flexure at this stage also affects the 
position of all the visceral arches that he anterior to the hyal 
arch, and the palatoquadrate and mandibula, which represent 
either the cartilaginous bar of the mandibular arch alone, or 
that bar together with parts of the bars of one or more pre- 
mandibular arches, lie, when first laid down, at right angles to 
the trabecule, and hence in a nearly horizontal instead of a 


* For explanation of the Plates see p. 148. 


Proc. Zoou. Soc.—1917, No. VIII. 8 


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106 MR, E. PHELPS ALLIS, JUN., ON 


vertical position. Because of this, and also. because of the 
relatively much greater development of these bars, they have 
crowded the bars of the more posterior arches posteriorly to 
such an extent that their dorsal ends lie in the auditory and 
spinal regions. ‘The sphenolateral cartilage of either side (ali- 
sphenoid, Sewertzo{) lies, at this stage, in the horizontal plane 
of the parachordals, lateral to the hollow of the plica encephali 
ventralis. The nervi facialis, trigeminus, and ophthalmicus 
profundus all ran outward postero- dorsal to this cartilage, the 
oculomotorius and opticus, and apparently also the trochlearis, 
running outward antero-ventral to it. In slightly older stages 
an ethmoidal cartilage is developed, which begins at a point 
ventral to the lateral edge of the trabecula of its side, and 
ventral to the eyeball, and runs dorso-anteriorly dorsal to the 
nasal sac. 

When the cranial flexure later becomes reduced, there is first 
formed, in Pristiwrus, and hence probably in all the Selachii, a 
pronounced pontial flexure, but this flexure also becomes later 
reduced, and almost entirely disappears in the adult. During 
these changes the trabecule curve forward, or forward and 
upward, following the curved veftral surface of the brain, and 
this change in their direction and position affects the mandibular 
branchial bars, which, retaining their perpendicular relations to 
the trabecule, swing downward and forward, and so acquire a 
position vertically beneath the latter cartilages. These mandi- 
bular bars had apparently already become attached to the hyal 
bars by connective or hgamentous tissues, and the latter bars 
and the bars of the branchial arches had become similarly 
attached to each other both dorsal and ventral to the branchial 
clefts. When the mandibular bars swung downward and 
forward they accordingly pulled on the more posterior bars, but 
as the dorsal ends of the hyaland branchial bars, and the ventral 
ends of the latter bars, had become attached to tissues in the 
region where they lay, they were relatively fixed in position, and 
the middle elements, alone, of each bar could be moved forward, 
the branchial clefts of. course shifting forward with them. This 
gave rise to the sigma form of branchial bar actually found in 
the adults of these fishes. In the Teleostomi, where there is but 
little cranial flexure when the neurocranium begins to chondrify, 
the trabecule are laid down in the line prolonged of the para- 
chordals. The mandibular branchial bars were accordingly there 
laid down approximately in a vertical position, and the bars of 
the more posterior arches, not being pushed poster lorly by them, 
retained their primitive attachment in the cranial region. The 
sigma form of arch was accordingly not impressed upon the bars 
of the branchial arches of these fishes. 

The trabecule, in the older embryos described by Sewertzoff, 
have fused with each other in the median line anterior to the 
pituitary body, and been prolonged, as a wide median plate, to 
the nasal region. There the plate contracts abruptly and is 


THE SKULL OF CHIMARA, 107 


prolonged between the nasal sacs as a narrow rostral stalk in 
Acanthias, but as a wide internasal plate in Pristiurus, the two 
structures evidently being, as they are said by Sewertzoff to be, 
strictly homologous. Anterior to the nasal sacs this stalk or 
plate expands laterally, on either side, and in dcanthias forms 
the rostral plate of Sewertzoft’s descriptions, a median keel- 
shaped ridge projecting ventrally from its ventral surface and 
being continuous with a similar ridge on the ventral surface of 
the rostral stalk. In Pristiurus the rostral plate is said by 
Sewertzoff to be represented in the three bars of the rostral 
basket, the keel of the plate of Acanthias representing the middle 
bar of the basket of Pristiwrus, and the lateral portions of the 
plate representing the lateral bars. This I consider to be an 
erroneous interpretation of the conditions, the rostral plate of 
Acanthias certainly including, with the three bars of the rostral 
basket of Pristiurus, the narrow strip of cartilage that, in the 
latter fish, runs laterally immediately anterior to the nasal sac 
and is called by Sewertzoff the cartilage c. This cartilage c is 
said by Sewertzoff to not yet be developed in the oldest embryos 
of Acanthias examined by him, but to be found in the adult as a 
bar of cartilage that cuts the fenestra olfactoria of embryos into 
two parts, the lateral one being the definitive foramen olfac- 
torium, and the mesial one the basal communication canal of 
Gegenbaur’s (1872) descriptions of the adults of certain of. the 
Selachii. The large perforation of either side of the floor of the 
rostral basket of Pristivrus is accordingly said, in accord with 
Gegenbaur’s earlier conclusion, to be a basal communicating canal 
strictly similar to the one found in the adult Acanthias, but 
greatly enlarged and shifted forward; this large perforation 
of the rostral basket of this fish, and the large opening that 
actually transmits the nervus olfactorius, thus together repre- 
senting the fenestra olfactoria of embryos of Acanthias. In an 
earlier work (Allis, 1913), I came to the conclusion that these 
perforations of the rostral basket of the Carchariidee and 
Scyllide were not basal communicating canals, and that they were 
‘probably ... simply defects in the cartilage (of the rostrum) 
due to the presence of the Jarge number of ampulle found in 
this region in these fishes,” and this conclusion I consider to be 
confirmed by conditions that I have since found in one specimen 
of Acanthias blainvillii. In this specimen there is a basal com- 
municating canal in normal position on each side of the head, 
and on one side of the head the rostral process is perforated by 
an irregular opening certainly strictly homologous to the per- 
forations found in the Carchariide and Scylliide; and there are 
numerous ampulle lying opposite the perforation. 

The cartilage ¢ of Pristiurus thus certainly corresponds to a 
part of the rostral plate of Acanthias, and the two cartilages of 
Pristiwrus, one on either side of the head, can, for convenience, 
he together called the rostral plate, although they represent only a 
part of that plate as found in dcanthias. This plate and its stalk, 

gx 


108 MR. E, PHELPS ALLIS, JUN., ON 


in both these fishes, lie, at this age, in the line prolonged of the 
trabecular plate, and each lateral end of the narrow plate of the 
one (Pristiurus), and each postero-lateral corner of the broad 
plate of the other (Acanthias), turns dorso-posteriorly as a band- 
like process which passes dorsal to the nervus olfactorius and 
dorsal to the nasal sac, and, fusing posteriorly with the anterior 
ends of both the ethmoidal and sphenolateral cartilages, forms 
the dorsal boundary of the opening that transmits the nervus 
olfactorius ; the sphenolateral cartilage quite certainly not taking 
any part in the formation of the actual boundary of the opening. 
This opening will be hereafter called the fenestra olfactoria in 
both these fishes, notwithstanding that it is said by Sewertzoff to 
be later subdivided into a foramen olfactorium and a basal com- 
municating canal in Acanthias, and to represent the definitive 
foramen olfactorium in Pristiurus. 

The nasal capsule of either side, as shown in Sewertzoft’s 
figures, is represented by a curved plate of cartilage which 
encircles the basal portion of the anterior and lateral surfaces of 
the nasal sac and projects ventrally from the ventro-lateral 
surface of the rostral plate. It is completely fused, by its dorsal 
edge, with the rostral plate, but its position, suggests that it is 
primarily an independent cartilage secondarily fused with that 
plate. Parker (1876) shows the nasal capsule in a somewhat 
different condition in Scylliwm, but the figures given by him are 
difficult to comprehend and not readily compared with those 
given by Sewertzoff. The capsule, however, as shown by both 
these authors in these several fishes, lies ventro-lateral to the 
trabecular and rostral cartilages, and where, in these or others 
of the Selachu, the capsules of opposite sides have been pressed 
together so as to meet in the median line and there be separated 
only by a septum nasi, that septum lies ventral to the trabecule, 
as shown in Parker’s sectional views of Seylliwm and in Gegen- 
baur’s sectional view of Mustelus. The trabecule, in each of 
these latter fishes, curve dorsally above the nasal sacs and are 
furnished with a deep median subethmoidal keel which extends 
ventrally between the sacs and is shown, in each case, as largely 
fenestrated in its middle portion. Parker (1876, p. 207) appa- 
rently considered this internasal septum to be an upgrowth of 
the trabecular cartilage, but Parker and Bettany (1877) say that 
the trabecular plate gradually rises to the level of the nasal roof 
and there suddenly narrows and is confluent with the internasal 
septum, this seeming to mean that the septum lies ventral to 
the trabecule, as i consider it todo. This difference in interpre- 
tation, by these two authors, of identical conditions is apparently 
due to Parker having considered the trabecule to be continued 
forward in the ventral edge of the subethmoidal keel, while 
Parker and Bettany consider them to be represented in the 
plate from which that keel descends, that is in the trabecular 
crest of Parker’s descriptions. 

A ventro-Jateral trabecular process, lying antero-ventral to 


THE SKULL OF CHIMERA, 109 


the nasal sac, is shown, slightly developed, in Parker’s (1876) 
figures of embryos of Scylliwm, the process there projecting 
laterally from the ventral edge of the subethmoidal keel. This 
process is called by Parker the cornu trabecule ; but two other 
cartilages on each side of the head are also so designated by him, 
one being the cartilage c of Sewertzoft’s descriptions, and the 
other the lateral bar of the rostral basket. This ventro-lateral 
trabecular process is not shown or described by Sewertzoff even 
in the oldest of his embryos, but that it is not peculiar to fishes 
in which there is a subethmoidal keel is shown by the conditions 
in the adult Chlamydoselachus, where the process is found well 
developed and there forming the anterior portion of the solum 
nasi of my descriptions of that fish (Allis, 1913). 

The beginnings of a process that lies postero-ventral to the 
nasal sac is shown, in Sewertzoff'’s figures, at the ventro-lateral 
corner of the ethmoidal cartilage, and it 1s apparently it alone 
that is later prolonged into the important ventro-latero-posteriorly 
directed process that is currently called the antorbital or ethmo- 
palatine process. This antorbital process is, however, said to be 
formed, in certain other fishes and in higher vertebrates, by an 
outgrowth of the trabecular cartilage. It is therefore possible 
that there are two processes here, one of ethmoidal and the other 
of trabecular origin. If so, they are indistinguishably fused 
with each other in the Selachii, and to avoid confusion I shall 
call the process, whatever its origin, the ventro-lateral antorbital 
process, the entire antorbital process being considered to form 
the anterior wall of the orbit and to include both this ventro- 
lateral process and a more or less developed dorso-lateral 
process found in the adults of most fishes and frequently called 
the antorbital or preorbital process. 

The sphenolateral cartilage has, in the oldest embryos of 
Acanthias described by Sewertzoff, grown forward and fused with 
the ethmoidal cartilage, and its supraorbital ridge, turning 
downward at its anterior end, forms the dorsal portion of the 
lateral edge of the antorbital process, that portion of that edge 
corresponding to the well-developed dorso-lateral antorbital 
process found in certain other fishes. The ramus ophthalmicus 
superficialis trigemini and the nervus ophthalmicus profundus 
both, in younger embryos, run forward through the orbit and 
then dorsal to the nasal sac, passing external to the ethmoidal 
cartilage, between the dorso-lateral and ventro-lateral antorbital 
processes. But when the sphenolateral and ethmoidal cartilages 
fuse, the two nerves and an accompanying vein and artery 
become more or less enclosed between them, either separately or 
together. The nasal vein, which comes from the nasal pit to fall 
into the orbital venous sinus, also either becomes enclosed 
between these two cartilages or is enveloped in the lateral edge 
of the ethmoidal cartilage alone. 

The nasal sac of embryos of the Selachii thus lies between four 
processes, more or less developed, two being derived from the 


fel) MR. E. PHELPS ALLIS, JUN., ON 


trabecule and lying anterior to the sac, and: two derived either 
from the trabecular, ethmoidal, or sphenolateral cartilages and 
lying posterior to the sac; and the nasal eapsule, when it 
develops, also lies between and 1s more or less fused with these 
processes. ‘The dorsal trabecular process and the ethmoidal con- 
stituent of the dorso-lateral antorbital process always fuse with 
each other above the nervus olfactorius, so forming the dorsal 
boundary of the fenestra olfactoria, and they also form, together 
with the sphenolateral eartilage, the lateral wall of the cavum 
preecerebrale of my descriptions (Allis, 1913). The two ventral 
processes may also fuse with each other at their outer ends, so 
forming a perforated plate which lies beneath the nasal sac, the 
perforation of this solum nasi being closed by membrane and 
representing an unchendrified portion of the nasal capsule. 

The ventro-lateral antorbital process and the two prenasal 
processes of the Selachii are found in embryos of Ceratodus, the 
former being called by Greil (1913) the processus anbouicenke and 
the two latter the dorso-lateral and the anterior or lateral trabe- 
eular cornua. No derso-lateral antorbital process is shown in 
Greil’s figures of embryos of this fish, but it is shown by 
Fiirbringer (1904) in a figure of the ade Ceratodus, the process 
there apparently arising as a lateral process from the dorsal end 
of the posterior wall of the nasal capsule, and being called by 
Viirbringer the preorbital process. The septum nasi of Ceratodus 
apparently lies dorsal to the trabecule, as it does in the Teleostei, 
Holostei, and Crossopterygil, the septum in these several orders 
of foen accordingly not being the homologue of the septum in 
the Plagiostomi, unless it be that the internasal prolongation of 
the trabeculae of embryos of the former fishes corresponds to the 
ventral edge of the subethmoidal keel of embryos of the Plagio- 
stomi and not to the rostral stalk and rostral plate, which seems 
quite probable. 


CALLORHYNCHUS. 


In Callorhynchus, as in Acanthias and Pristiwrus, the trabecule 
are first laid down practically at right angles to the parachordals, 
as consideration of Schauinsland’s (1903) figures of a 60 mm. 
embryo of this fish makes plainly evident. This is due, as in the 
Selachu, to the marked cranial flexure at this period : but in 
C ‘allorhynchus the anterior portion of the brain, instead of 
recurving upward and forward as if seeking to reacquire a posi- 
tion in the line of the axis of the body, projects ventro-anteriorly 
ina straight line. Related to this retention, in this position, of 
this anterior portion of the brain, the eyeballs of the adult fish 
lie dorso-anterior to it; but whether it is this position of the 
brain that has forced the eyeballs to pass antero-dorsal to it, or 
the precocious development of the eyeballs that has prevented the 
brain from recurving upward, is not apparent. 

The trabecule fuse to form a wide trabecular plate which, 


THE SKULL OF CHIMARA. 111 


instead of curving gradually upward and forward, as in the 
Selachii, follows the straight line of the ventral surface of the 
brain until it reaches what Schauinsland considers to be the 
anterior end of the cranial cavity. There the trabecular plate, 
which at this age js still procartilaginous in its anterior portion, 
contracts abruptly to a narrow procartilaginous bar which hes at 
a marked angle to the trabecular plate, being directed dorso- 
anteriorly instead of ventro-anteriorly. This narrow procarti- 
laginous bar is called by Schauinsland the “ Septum 1 im vorderen 
Se tL ” but it is evident that it is the strict homo- 
logue of the rostral stalk of the Selachii. The three rostral 
processes of Schauinsland’s descriptions are all shown arising 
from this stalk, the median process from its dorsal end and the 
two lateral processes from its lateral edges at about the middle or 
dorsal third of its length. A short, plate-like subethmoidal keel 
has begun to develop on the external and hence morphologically 
ventral surface of the stalk. 

From either side of the dorsal end of this rostral stalk, a band- 
like process runs laterally and then posteriorly, and, spreading 
dorsally and ventrally, fuses with the procartilaginous anterior 
portion of the ethmoidal cartilage, which cartilage forms the 
antorbital process and the anterior wall of the orbit. This band- 
like process is accordingly a dorso-lateral trabecular process, 
similar to that in the Selachii, and the large opening enclosed 
between it, the rostral stalk, and the ethmoidal and trabecular 
procartilages should be primarily a simple fenestra olfactoria. 
similar to that in the Selachii, but, even in the voungest embryos 
described by Schauinsland, the dorsal portion of the fenestra has 
apparently been already converted into a fenestra nasalis by the 
beginning of the development ef the nasal capsule, 

The development of the nasal capsule is not readily com- 
prehended from the several figures given by Schauinsland. In 
the figures of the 60 mm. embryo “(ligs. 130-131) the capsule 
is not indicated by index-letters, but the lateral edge of the 
anterior opening of the cranial cavity is irregular, the dorsal 
portion projecting anteriorly beyond the ventral portion and 
occupying the place in Sieh. in the 85 mm. embryo (figs. 124— 
126), the nasal capsule is shown. In an embryo said to be older 
than the 85 mm. one, but of which the length is not given, the 
nasal capsule is a dome-shaped protuberance of the dorso-lateral 
portion of the cranial wall immediately posterior to the actual 
anterior opening of the cranial cavity. Ventral to this so-called 
capsule the lateral wall of the cranial cavity runs evenly onward 
to the edge of the anterior opening of the cranial cavity, the 
capsule apparently not extending into this part of the cranial 
wall notwithstanding that the ventral portion of the nasal sac 
must certainly lie internal to it. The nasal sac also quite 
certainly does not project upward into the dome-shaped pro- 
tuberance, that protuberance apparently lying above the nasal 
sac and resembling a dome-shaped swelling that | find, in the 


412 MR. E. PHELPS ALLIS, JUN., ON 


adult A/ustelus, lying posterior to the foramen olfactorium and 
directly above the lobus olfactorius. 

The dome-shaped protuberance of Callorhynchus, the so-called 
nasal capsule, always lies, at all stages of its development, in the 
region between the bases of the median and lateral rostral 
processes, and as the capsule increases in size the bases of the 
vostral processes are correspondingly separated from each other, 
and the anterior opening of the cranial cavity carried corre- 
spondingly forward ; but whether this growth of the cranial wall 
takes place posterior to the fenestra olfactoria of the 60 mm. 
embryo, or is due to growth antevior to that fenestra, cannot be 
told foam the figures. In the one case the nasal sac, which 
certainly lay primarily anterior and external to the cranial 
‘avity, would have been pulled relatively backward into the 
anterior end of that cavity; the fenestra olfactoria of the 60 mm. 
embryo would remain morphologically unchanged, but would 
change in function from a fenestra olfactoria to a fenestra 
nasalis; and the so-called nasal cartilages of Schauinsland’s 
figures would represent detached pieces an an undeveloped nasal 

apsule, In the other case, the nasal capsule would be developed | 
as a direct anterior, but morphologically ventral, prolongation of 
the cranial walls, a fenestra nasalis gradually developing external 
to the fenestra olfactoria of the 60 mm. embryo. The conditions 
in the adult Chimera decidedly favour this latter view, but one 
of Schauinsland’s figures of Callorhynchus equally decidedly 
tavours the former view. The figure in question (fig. 165) gives 
n median vertical sectional view of an embryo older than the 
85 min. one, and shows the brain in place in ‘the cranial cav ity. 
The trabeculae are shown lying in the line prolonged of the 
parachordals, which is not in accord with the figures of other 
embryos both older and younger than this one. The septal 
cartilage is shown as a simple rostral staik, while in the younger, 
85 mm. embryo, it is already an important plate-like structure 
the posterior portion of which corresponds to the rostral stalk 
of the 60 mm. embryo, and the anterior portion to the sub- 
ethmoidal keel of that’ stalk. The rostral stalk of the embryo 
shown in figure 165 projects dorso-posteriorly, instead of, as in 
the other embryos, dorso-anteriorly, and the vential portion 
of the nasal sac is shown extending forward anterior to the level 
of the base of the median rostral process and close to the ventro- 
anterior corner of the ehondrocranium. ‘The nasal sacs, as here 
shown, must accordingly project beyond the anterior openings of 
the oem cavity, if those openings are found in this embryo at 
all in the positions that they have in the other embryos, both 
older and younger, and furthermore, they must he dorsal to the 
trabecule, as they do in Ceratodus and the Teleostei, instead of 
ventral to the trabecule, as they do in the Selachii and as they 
must also in Callorhynchus if the septum nasi of this fish hes 
ventral to the rostral stalk, as shown in Schauinsland’s other 


£3 


THE SKULL OF CHIMERA, sgl 


figures. There would accordingly seem to be some error in this 
figure 165. 

The ethmoidal cartilage, as shown in these embryos of Callo- 
rhynchus, arises from the trabecular plate posterior to the 
fenestra olfactoria of its side, and runs dorsally and then 
anteriorly to meet and fuse with the band-like dorso-lateral 
trabecular process. At its ventral end the ethmoidal cartilage 
gives off a lateral process which, even in the earliest stages shown 
by Schauinsland, has already fused completely with the palato- 
quadrate. The sphenolateral cartilage grows forward dorsal to 
the orbit, and in the 60 mm. embryo, where it is still pro- 
cartilaginous, bifurcates anteriorly, one end passing dorsal and 
the other ventral to the ramus ophthalmicus superficialis tri- 
gemini, which nerve includes the lateralis fibres destined to 
supply the latero-sensory and ampullary organs of the region. 
The ventral one of these two ends of the sphenolateral cartilage 
has already fused, in this embryo, with the ethmoidal carti- 
lage internal to the ramus ophthalmicus superficialis, and the 
dorsal end also later fuses with that cartilage but external to 
the ophthalmicus superficialis, this nerve rae being enclosed in 
a foramen and a dorso-lateral antorbital process being formed. 
This process is called by Schauinsland the preorbital process, and 
both it and the postorbital process are said by him to be 
primarily independent pieces of cartilage. Concomitantly with 
the formation of this process and the related foramen, a root of 
cartilage has been formed over the nervi ophthalmicus super- 
ficialis trigemini and ophthalmicus profundus, enclosing them im 
a median canal called by Schauinsland the ethmoidal canal. This 
canal is evidentiy formed by the coalescence, in the median line, 
of two canals, one on either side of the head, each of these canals 
being formed by the roofing over of the groove that, in the 
Selachii, lodges the corresponding portion of the ramus ophthal- 
micus superficialis trigemini. In certain specimens of Chlamydo- 
selachus I find these grooves partially roofed, either by lips of 
cartilage that project toward each other from the edges of the 
grooves, or by nodules of cartilage suspended in membranous 
strands that stretch across the grooves ; and if this roofing process 
were to be completed and the grooves so formed pressed together 
in the median line, a median canal would be formed the roof of 
which would form the actual roof of the cranium and its floor the 
root of the cranial cavity. 

A remnant of the fenestra precerebralis of my descriptions of 
the Selachii (Allis, 1913) is found, as will be later fully explained, 
in the small median opening marked ¢ in Schauinsland’s figures 
124 and 125, and said by him to be a “Spalte, oa das 
Schideldach von dem vorderen ethmoidalen Teil des Schiidels 
trennt.” The two large openings in the cranial roof that are 
called by Schauinsland the anterior and posterior divisions of the 
primary single ‘ Preefrontallicke,” are not parts of the fenestra 


114 MR, E. PHELPS ALLIS, JUN., ON 


preecerebralis, and hence not parts of the Prefrontalliicke of 
Gegenbaur’s (1872) descriptions of the Selachii. The cavum 
precerebrale, which in the Selachii les directly anterior to the 
fenestra preecerebralis, cannot be represented in any part of 
the ethmoidal canal, for, aside from the origin of that canal as 
above explained, the floor of the canal is not formed by the 
trabecule, while the floor of the cavum precerebrale is. The 
hind end of the ethmoidal canal is shown, in Schauinsland’s 
figure 165, lying -but slightly anterior to the anterior surface of 
the mid-brain. If this figure is correct, the membranous mesial 
walls of the orbits must accordingly, even in the oldest embryos 
considered by Schauinsland, be separated from each other by 
a considerable interval. 

The chondrocranium of embryos of Callorhynchus thus ap- 
parently owes the several points in which it differs from that of 
the Selachii mainly, if not wholly, to the fact that that portion 
of the central nervous system that lies anterior to the plica 
encephali ventralis not only projects ventrally or antero-ventrally 
at the time when the trabecule are laid down, but that, for some 
reason, 1t has continued to lie in that position instead of later 
eradually curving forward or forward and upward. The tra- 
becule still apparently seek to curve upward into the line 
prolonged of the pavachordals, but, because of the interference of 
the overlying brain, this upward curve is found immediately 
anterior to the lobi olfactorii instead of, as in the Selachii, in the 
pituitary region. 

The mandibular branchial bars were doubtless laid down 
primarily at right angles to the trabecule, and hence, as in the 
Selachii, in a nearly “horizontal position, but as the trabecule 
later grow downward and forward instead of, as in the Selachii, 
curving gradually forward or forward and upward, the mandibular 
bars have been distorted. The dorsal ends of the processus 
oticus and basilaris, the latter representing the primitive dorsal 
end of the arch, apparently remain approximately in their 
prunitive positions in relation, respectively, to the lateral wall of 
the otic capsule and the trabecule, but they have been stretched 
out into long cartilages by the marked ventro-anterior growth of 
the trabecule, and the epal (quadrate) and ceratal (mandibuia) 
elements of the arch lie in the region of the antorbital process. 
This will be again referred to when describing the conditions in 
Chimera. Because of this distortion and change in position of 
the mandibular branchial bars, the branchial bars of the more 
posterior arches have also been carried forward and somewhat 
downward without having been previously pushed backward to ' 
the extent that they were in the Selachii, and the dorsal ends 
of the anterior branchial bars are shown lying in the cranial 
region in Schauinsland’s oldest embryos. The sigma form of arch 
has, however, been impressed upon their dorsal ends, but not 
upon their ventral ends. 


THE SKULL OF CHIMARA. Pe 


CHIMERA. 


1. Newrocranium. 


In a six-month embryo of Chimera colliet, Dean (1906, p. 108) 
shows the trabecule projecting ventro-anteriorly at a marked 
angle to the parachordals, which would seem to establish that 
when first chondrified they lay, as they do in Callorhynchus, 
approximately at right angles to the parachordals. In the adult, 
the conditions shown by Dean in this embryo still persist to a 
marked extent, for, as shown in the accompanying figure (PI. I. 
fig. 2), the line of the vertebral column, if pr olonged, “would pass 
approximately across the dorsal edge of the postelinoid wall and 
issue from the cranium somewhat dorsal to the base of the 
median rostral process, the larger part of the prechordal portion 
of the cranium projecting ventro-anteriorly below this line at an 
angle of about 30°. 

Because of this position of the trabecule, the mid-ventral line 
of the chondrocranium of the adult Chimera projects ventro- 
anteriorly, and from the level of the foramen magnum to the 
level of the fenestre nasales it is shghtly curved, the hollow of 
the curve presented ventro- posteriorly, Anterior to the level 
of the fenestre nasales, the mid-ventral line changes abruptly in 
direction, running at ne dorso-anteriorly and then ventro- 
anterior ly and ending at the anterior end of the short beak-hke 
process of the Enon dimmer iniarn, This latter process is morpho- 
logically subnasal in position, as will be later explained, but as it 
has the appearance, in lateral view, of being prenasal, it may be 
ealled the prenasal process. 

Beginning at about the level of the middle of the orbit and 
extending forward to its anterior edge, there is a median, longi- 
tudinal, gash-like groove which lodges the degenerated tissues “of 
the extracranial portion of the’ hypophysis, the groove being 
deepest at its posterior end and gradually vanishing anteriorly. 
This groove represents a persisting remnant of the hypophysial 
fenestra, that fenestra being, in the adult, completely closed 
toward the cranial eavity, as it apparently was even in the 
ehondrocranium of the sixth-month embryo shown by Dean. 
Starting lateral to this grodve, on either side, a pronounced but 
low and rounded ridge runs antero-laterally to the level of the 
ventral end of the antorbital wall, where it turns somewhat 
abruptly antero-ventrally and but shghtly laterally and bears on 
its end an articular facet and an articular head, the former lying 
directly mesial to the latter and both surfaces giving articulation 
to the mandibula. From the anterior edge of the articular facet 
the narrow ventro-lateral edge of the chondrocranium runs 
antero-mesially in a curved line, concave ventrally, till it reaches 
a point slightly anterior to the ventral edge of the fenestra 
nasalis, where it turns anteriorly and becomes the ventral edge 
of the heak-like prenasal process. 


PG MR. E. PHELPS ALLIS, JUN., ON 


Between the mandibular articular facets of opposite sides, the 
ventral surface of the chondrocranium is deeply concave, this 
concavity lying in the ethmoidal region and being traversed by 
the raised median longitudinal ridge of the fused trabecule. 
Anterior to this large subethmoidal depression there is a smaller, 
subnasal one which lies in the hollow of the beak-like prenasal 
process, the two depressions being separated by a V-shaped ridge, 
each arm of the V running antero- -laterally from the projecting 
ventro-posterior corner of the subethmoidal keel. The anterior 
dental plates lie against the dorsal wall of the anterior half of 
the subnasal depression, the posterior dental plates lying against 
the anterior border of the subethmoidal depression. From the 
bottom of the posterior half of the subnasal depression a tough 
pad of tissue arises and projecting ventrally fills the space between 
the dental plates. 

The bottom of the large subethmoidal depression is separated 
by the raised ridge of the fused trabecule into two parts, each 
triangular in shape, and as each depression lies beneath the 
lateral portion of the ethmoidal cartilage it may be called the 
ectethmoidal depression, the two ectethmoidal depressions forming 
the large subethmoidal one. Hach ectethmoidal depression is 
traversed by the nasal vein of its side, this vein, in some spe- 
cimens, lying in a pronounced groove at the bottom of the 
depression. This vein comes from the nasal capsule and the 
regions anterior to it, and is directly continuous, in the nasal 
capsule, with a vein that enters the cavum cranii, accompanying 
the nervus olfactorins, and joins a vein that issues from the 
cranial cavity through the foramen for the nervus vagus. The 
nasal vein, running posteriorly from the nasal capsule, first 
traverses a canal that leads from that capsule through the 
anterior wall of the ectethmoidal depression, then receives a 
branch that has traversed the overlying cartilage through a small 
foramen, and then itself traverses a foramen in the posterior wall 
of the ectethmoidal depression to enter the orbit and fall into the 
orbital venous sinus. The ectethmoidal depression of this fish 
thus corresponds, in general position and in its relations to the 
nasal vein, to the ectethmoidal chamber of my descriptions of 
Chlamydoselachus (Allis, 1913), that chamber being, in certain 
of the Selachii, an open fossa which may, as in Heptanchus, be in 
direct communication with the cranial cavity through the basal 
communicating canal of Gegenbauer’s descriptions. There is, in 
Chimera, no indication of this latter canal, but it is perhaps 
worthy of note that in Lepidosiren there is, in just this region, 
a median perforation of the basis cranii (Bridge, 1898). 

At the postero-mesial corner of the ectethmoidal depression of 
Chimera, near the summit of the ridge that there bounds the 
depression, there is a groove which leads postero-mesially to a 
canal which traverses the cartilage of the basis cranii and trans- 
mits the efferent pseudobranchial, or so-called anterior carotid 
artery, this canal opening on the dorsal surface of the cartilage 


THE SKULL OF CHIMAERA. aa 


in the region of the line of attachment of the membranous inter- 
orbital wall, and directly ventro-lateral to the foramen by which 
the nervus opticus traverses that membranous wall. 

The foramen opticum lies directly dorsal to a well-marked and 
rounded transverse ridge on the floor of the cranial cavity, this 
ridge marking the anterior end of the pituitary fossa and hence 
being the presphenoid bolster of my descriptions of the Selachu. 
The cranial cavity is here abruptly constricted, latero-mesially, 
and this constriction, in its ventral portion, is produced by the 
pinching in of the mesial membranous walls of the orbits, the 
membrane of either orbit being pinched inward and downward 
so that the internal surface of its ventral portion lies closely 
upon the cartilage of the basis cranu. The membrane there 
usually, but not always, becomes adherent to the cartilage, and 
a second line of attachment of the membranous cranial wall, 
lying mesial to the primitive line, is acquired. A depression is 
thus formed in the ventral portion of the outer surface of the 
membranous wall of each orbit, and it is filled with tough fatty 
connective tissue. The original line of attachment of the mem- 
branous wall forms the ventro-lateral edge of this depression, 
and lies lateral to the foramen for the efferent pseudobranchial 
artery, while the secondary line of attachment lies mesial to that 
foramen, the membrane covering the foramen. Having issued 
from its foramen, the efferent pseudobranchial artery runs pos- 
teriorly beneath the covering membrane and issues in the cranial 
cavity at the base of the posterior wall of the depression, there 
lying posterior to the nervus opticus. It then immediately gives 
off a recurrent branch which perforates the posterior wall of the 
depression, traverses the tough fatty tissue that fills the depres- 
sion, and, joining the nervus opticus, accompanies that nerve to 
the eyeball. After giving off this branch, the pseudobranchial 
artery turns mesially and immediately separates into anterior 
and posterior cerebral arteries. 

When the secondary attachment of the membranous orbital 
wall to the dorsal surface of the basis cranii is not strong, the 
pseudobranchial foramen opens directly into the cranial cavity, 
but when the attachment is strong it opens external to that 
cavity but beneath the covering membrane. The nervus opticus, 
after issuing from its foramen, lies directly upon this adherent 
membrane, and the membrane may there become thinned to 
such an extent that, in dissections not carefully made, the 
pseudobranchial foramen appears as a perforation of the sub- 
orbital shelf lying wholly external to the interorbital wall; this 
apparently having been the condition in the specimen examined 
in connection with my earlier work on the arteries of this fish 
(Allis, 1912), where the foramen is said to open external to 
the cranial wall, The foramen however lies, morphologically, in 
the actual floor of the cranial cavity, as it is shown in Schauins- 
land’s figure of an embryo of Callorhynchus. In the Selachii 
this foramen always lies, in all the specimens that I have 


118 MR. E. PHELPS ALLIS, JUN., ON 


examined, dorsal to the trabecule, and where there is a sub- 
orbital shelf it lies dorsal to that shelf. This foramen of 
Chimera, which is quite unquestionably the homologue of the 
foramen in the Selachii, must then have become surrounded by 
and quite deeply enclosed in the dorso-lateral edge of the 
trabecula, the membranous lateral wall of this part of the neuro- 
cranium thus lying lateral to it and the foramen appearing to lie 
in the floor of the cranial cavity. The foramen is not shown 
in Dean's figure of an embryo of Chimera, doubtless because 
it is hidden, in dorsal view, by the supraorbital portion of the 
cranial wall. 

The branch of the pseudobranchial artery that accompanies 
the nervus opticus to the eyeball was called by me, in the earlier 
work just above referred to, the arteria ophthalmica magna, this 
identification of the artery being based on its apparent origin, in 
a dissection in which the tissues had evidently been torn, from 
the pseudobranchial artery before that artery enters the cranial 
cavity. This is, however, an error, the artery quite certainly 
being the arteria centralis retine, or optic artery, of my descrip- 
tions of the Selachii, and no arteria ophthalmica magna being 
found in this fish. 

At the hind end of the orbit five foramina, lying close together, 
perforate the cranial wall and transmit the nervi profundus, 
trigeminus, buccalis lateralis, facialis, and abducens. The fora- 
men for the facialis is always separated from the other foramina 
by cartilage, and the foramen for the abducens usually so 
separated, the other three nerves usually issuing through a single 
foramen but being separated from each other by membrane. 
Dorsal to these foramina there is a large foramen for the ramus 
ophthalmicus superficialis *, and two foramina that he near the 
edge of the membranous mesial wall of the orbit, one of them 
transmitting the nervus trochlearis and the other a venous vessel 
which 1s doubtless the anterior cerebral vein of my descriptions 
of Amia and the Teleostei. Slightly anterior to the five foramina 
that transmit the trigeminus, facialis, and abducens nerves there 
is the foramen for the nervus oculomotorius, which lies at the 
edge of the membranous mesial wall of the orbit, and ventro- 
lateral to that foramen there is a small foramen for the pituitary 
vein, this latter foramen opening into the pituitary fossa. The 
vein that traverses this latter foramen is small, and although it 
connects with ‘its fellow of the opposite side im the usual piscine 
manner, it is always in communication with a vein that issues 
from the cranial cavity through the foramen for the nervus 
vagus. There is in this fish no transverse pituitary canal such 
as is found in many of the Selachii. 

Lateral to the several foramina above described, there is a 
large perforation of the suborbital shelf which transmits the 
ramus byoideo-mandibularis facialis and the vena jugularis, and 
anterior to that foramen there is another perforation of the 


* A small foramen is, by error, shown in figure 3, Plate IT., lying slightly ventral 
to this large foramen. It does not ecist in the fish. 


THE SKULL OF CHIM-ERA. 119 


shelf which transmits the arteria carotis externa and the ramus 
palatinus facialis, these two foramina being persisting remnants 
of the palatoquadrate fissure of Dean’s (1906, p. 108) deser iptions 
of embryos. ‘That fissure of embryos Ge as shown in Dean’s 
figures, between the palatoquadrate, the trabecula, and the 
anterior edge of a shelf of cartilage that projects nee ally from 
the neurocranium beneath the hind end of the orbit and is 
fused, in its antero-lateral portion, with the palatoquadrate. 
The vena jugularis accordingly les definitely ventral to this 
laterally projecting shelf-like process of the neurocranium, and 
hence also morphologically ventral and internal to the dorso- 
posterior portion of the palatoquadrate. If then this shelf-like 
process is an outgrowth of the neurocranium, the dorso-posterior 
end of the palatoquadrate, as shown by Dean, must be the 
processus oticus of that cartilage, but it seems much more 
probable that the shelf itself is the processus oticus, or extra- 
branchial element of the arch. Dean (/. c. p. 129) considers this 
shelf to be the posterior portion of the pharyngeal elenient of 
the mandibular arch, but mm that case this element of the arch 
has fused with the neurocranium dorsal to the vena jugularis, 
which would be in marked exception to the conditions found in 
all other fishes that I know of (Allis, 1915). It accordingly 
seems to me, as above stated, that this shelf must represent 
the extrabranchial of the arch, or processus oticus quadrata, 
apparently found only partially fused with the palatoquadrate in 
Dean’s six-month embryo. The vena jugularis, ramus hyoideo- 
mandibularis facialis, and arteria carotis externa would then all 
lie ventral and internal to this element of the arch, as they 
normally should, and the foramina for these several structures 
would represent the trigemino-facialis chamber = the fish, as 
1 have already sug egested i in an earlier work (Allis, 1914). 

The orbit has postorbital and antorbital processes. The 
postorbital process is a pronounced and slightly curved ridge 
which is fused ventrally with the suborbital ‘shelf. The dorso- 
lateral antorbital process 1s a short but tall ridge which lies at 
the dorso-anterior edge of the orbit, and its base is perforated 
by a foramen which transmits the ramus ophthalmicus super- 
ficialis trigemini from the orbit to the ethmoidal canal. 
Immediately ventral to this process there is a foramen which 
transmits a vein and artery, and antero-ventral to this latter 
foramen there is another foramen which transmits the nervus 
ophthalmicus profundus; both these foramina leading into the 
median ethmoidal canal. A small foramen dorsal to these 
foramina also leads into the ethmoidal canal and gives exit to 
a small branch of the ophthalmicus superficialis. A small 
foramen on the dorso-mesial surface of the nasal capsule gives 
passage to a branch of the orbital branch of the external carotid, 
and anterior to 1t there is another small foramen for a delicate 
branch of the ophthalmicus nerve. The ventro-lateral antorbital 
process is simply a ridge which traverses the suborbital shelf 
and is continued onto the palatoquadrate, where it turns 


120 MR. E, PHELPS ALLIS, JUN., ON 


posteriorly and joins the ventral end of the postorbital process, 
this ridge forming the boundary between the horizontal portion 
of the suborbital shelf and its ventrally projecting eaves-like 
lateral edge. Between its dorso-lateral and ventro-lateral 
processes the remainder of the antorbital process is a wide and 
gently rounded surface which extends from the orbit to the 
nasal capsule, and is so completely fused ventrally with the 
dorsal edge of the palatoquadrate that even the line separating 
the two cartilages cannot be recognised. 

The suborbital shelf is a wide thin plate of cartilage which 
extends between the antorbital and postorbital processes, and 
has a narrow, ventrally projecting, eaves-like lateral edge, just 
above referred to. ‘The shelf is prolonged posteriorly beyond the 
postorbital process, this postorbital portion of the shelf projecting 
ventro-posteriorly at a slight angle to the orbital pertion, and 
apparently representing, as just above explained, the processus 
oticus quadrati. It gives origin, on its ventral surface, to the 
musculus protractor arcuum branchialium of Vetter’s (1878) 
descriptions. On the ventral surface of the continuous shelf 
these two portions are separated from each other by a groove 
which lies directly ventral to the ventral end of the postorbital 
process and is directed postero-mesially. That part of the 
orbital portion of the shelf that lies lateral to the foramen for 
the nervus hyoideo-mandibularis facialis belongs to the palato- 
quadrate, and is probably formed by the epal (quadrate) and 
pharyngeal portions of the mandibular branchial bar, the 
pharyngeal element not projecting as a processus basilaris. 

The cranial cavity has large anterior and posterior portions, 
separated by a narrow neck in the orbital region, this neck 
lying between the ventral edge of the azygous portion of the 
membranous interorbital wall and the summit of the rounded 
transverse presphenoid bolster. The posterior portion of the 
cavity is tall, with a concave anterior edge the dorsal end of 
which projects dorso-anteriorly above the orbit like the horn of 
a crescent. The postclinoid wall lies at about the middle of the 
length of this posterior portion of the cavity and_ projects 
anteriorly or antero-dorsally. Between it and the foramen 
magnum there is a depression in the cranial floor, and anterior 
to it, between it and the presphenoid bolster, is the large 
pituitary fossa. The postero-dorsal end of the hypophysial 
groove on the ventral surface of the basis cranii lies slightly 
posterior to the summit of the presphenoid bolster. <A slight 
transverse ridge at about the posterior quarter of the pituitary 
fossa marks off a small posterior and deeper portion of the fossa. 
The foramina for the nervi facialis, trigeminus, and abducens lie 
dorsal or postero-dorsal to the postelinoid wall, and the small 
foramen for the pituitary vein antero-ventral to it. 

The anterior portion of the cranial cavity increases gradually 
in height from its hind end up to the base of the median rostral 


THE SKULL OF CHIMERA, 121 


process, that point marking also the anterior end of the ethmoidal 
canal. This latter canal is large and is separated from the 
cranial cavity by cartilage slightly thinner than that of the 
basis cranii. Beyond the anterior end of the ethmoidal canal 
the cranial cavity contracts somewhat abruptly to its anterior 
end, that end being marked, on either side, by the foramen 
olfactorium. The cranial cavity of the adult accordingly shows, 
when compared with Dean’s figures of embryos of this fish and 
with Schauinsland’s figures of embryos of Callorhynchus, that 
the dorsal portion of the chondrocranium has undergone marked 
lateral compression in post-embryonic stages, the membranous 
mesial walls of the orbits being pressed together so that they 
meet and coalesce in the median plane in front of the midbrain 
and dorsal to the forebrain, a considerable portion of the cranial 
cavity of embryos thus being obliterated. Because of this, or 
correlated to it, the midbrain becomes, in the adult, widely 
separated from the hind end of the ethmoidal canal, and the 
forebrain is correspondingly elongated. 

The two foramina olfactoria are separated from each other by 
a stout column of cartilage which hes nearly at right angles to 
the cranial floor, and each foramen is closed by a membranous 
evibriform plate. Hach foramen opens directly into the mesial 
portion of the posterior end of the related nasal capsule, that 
capsule being directed antero-ventrally and bulging Jaterally so 
that it forms a pronounced swelling on the lateral surface of the 
chondrocranium. The ventral edge of the capsule reaches nearly, 
but not quite, to the ventral edge of this part of the chondro- 
cranium, and the capsules of opposite sides are in contact in the 
median line, a septum nasi separating them from each other. 
This septum nasi is continuous, posteriorly, with the column 
of cartilage that separates the foramina olfactoria, that column 
lying in large part posterior to the plane of the foramina and 
representing the rostral stalk of embryos of Callorhynchus and 
of the Selachii. The septum nasi thus lies morphologically ventral 
to the stalk, and is formed either by a subethmoidal ‘keel alone, or 
by that keel fused with the contiguous mesial sides of the nasal 
capsules, the rostral stalk forming the morphologically dorsal 
edge of the septum, as it also does in Mustelus (Gegenbaur, 
1872, fig. 1, pl. 5) and Scylliwm (Parker, 1876, fig. 4, pl. 37). 
The nasal capsules thus fuse with each other, in Chimera, ventral 
to the trabecule, as they do in the Selachu, this confirming the 
conclusion already arrived at that they have this position im 
Callorhynchus. 

With the contact and fusion of the nasal capsules in the 
mnedian line ventral to the trabecule, the ventro-lateral trabecular 
processes of opposite sides, if they existed in this fish, must also 
have been pressed together in the median plane, and their flaring 
ventral ends probably form some part of the beak-like prenasal 
process, that process being, as already stated, morphologically 


Proc. Zeon, Soc 197, No. LX. 9 


122 MR. E. PHELPS ALLIS, JUN., ON 


subnasal, instead of prenasal in position. ‘The palatine pro- 
cesses of the palatoquadrate must, however, also take some 
important part in the formation of this beak, for the anterior 
dental plates, which it supports, lying as they do posterior to the 
primary upper lips (Allis, 1917 6), must belong to the primary, 
or palatoquadrate dental arcade. These plates cannot accord- 
ingly be intermaxillary (premaxillary) teeth, as Schauinsland 
(1903, p. 14) suggests as possible in Callorhynchus, and they must 
be either vomerine or vomero-palatine teeth according as the 
term palatine is used to apply to teeth developed in relation to 
the palatine process of the palatoquadrate or in relation to some 
part of the palatoquadrate that lies posterior to that process. 
‘The posterior dental plate of either side would then be either 
a pterygoid or a’ palato-pterygoid plate, a pterygoid element 
quite certainly being included in it. Further facts in favour of 
considering the beak of Chimera to be formed in part by the 
palatine process of either side are :— 

1. That the external surface of the prenasal beak gives 
articulation to the anterior end of a cartilage that is quite 
unquestionably an anterior upper labial, as will later be explained, 
and this anterior end of this labial is frequently, in the Selachii, 
in contact with the dorsal surface of the palatoquadrate but 
never in such contact with the neurocranium, as is also later 
explained. 

2. That the beak of Chimcera is traversed, on either side, by a 
small canal which, in the specimen used for illustration, begins 
on its dorsal surface by a single foramen and opens on its ventral 
surface by three small foramina which he internal to the anterior 
dental plate. This canal is traversed by a branch of the nervus 
maxillaris trigemini which descends over the lateral edge of the 
nasal capsule and is evidently destined to innervate the anterior 
dental plate and the related tissues. ‘The corresponding nerve in 
Mustelus, Chamydoselachus, and Raia, and hence probably in all 
of the Plagiostomi, runs forward along the external surface of 
the palatoquadrate and then over its ventro-lateral edge, thus 
being separated from the trabecule by the full width of the 
palatoquadrate, and while it might become enclosed in the lateral 
edge of the palatoquadrate it seems impossible that it could 
become so enclosed in the lateral edge of any cartilage of trabe- 
cular origin. 

The mandibula presents no special features that seem to 
require consideration, but it may be mentioned that there are 
two surfaces for the articulation with the palatoquadrate, the 
antero-lateral one being an articular facet and the postero-mesial 
one an articular head. It is also to be noted that the line of the 
gape of the jaws, when the mouth is closed, is approximately 
parallel with the line of the trabecule, the plane of the epal and 
ceratal elements of the mandibular arch thus retaining its 
primitive perpendicular relations to the trabecule and hence 
being directed postero-ventrally instead of ventrally. 


THE SKULL OF CHIMAIRA. 123 


7 ae Nasal and Labial Cartilages. 


The so-called nasal and labial cartilages of current descriptions 
of the Holocephali are much more numerous than the similarly 
named cartilages of the Selachii, and the conditions in the latter 
fishes must fee st be considered. 

The nasal cartilages of the Selachit ave limited to the ala nasalis 
(Nasenfliigelknorpel), and this cartilage I have recently described 
in a coeiaan number of these fishes and compared 16 with the 
cartilage in Chimera (Allis, 19176). It is accordingly not 
necessary to here consider it in the former fishes, 

The labial cartilages of the Selachii are limited to one or two 
related to the upper jaw and one related to the lower jaw. 
Gegenbaur (1872) called the one or two former cartilages the 
auterior and posterior upper labials, and as he at that time 
considered the premaxillary and caiscillar y bones, respectively, of 

the Teleostei to be dev eloped superticial to, and in relation to, 
these cartilages, he also called them the. premaxillary and 
maxilary labials. These latter “terms I shall avoid because 
of the implied homologies, which I consider not yet established. 
The single labial related to the lower jaw ‘Gegenhaur called the 
lower, or premandibular labial, the latter term being used by him 
because he then considered this labial and the maxillary labial 
to together represent persisting remnants of a premandibular 
visceral arch. ‘This latter term I shall also nora using in its 
place the indifferent term mandibular labial. Gs it should be 
later established that this labial and one or both of the upper 
labials ave structures related to a premandibular arch or arches, 
they can then be given names that will show their relations to 
the arch or arches to which they belong. 

The hind end of the mandibular labial and the corresponding 
end of one or both of the upper labials articulate with each other, 
or are connected by ligament, immediately posterior to the outer 
end of the line of the apele of the g gaye, and these ends of the 
labials there lie at a certain distance from the cartilages of 
the upper and lower jaws, separated from them by the thickness 
of the anterior edge of the musculus adductor mandibule. This 
point of articulation of the labials les at a variable distance 
anterior to the quadrato- mandibular articulation, and itis shown 
in nearly all of Gegenbaui’s figures of these fishes lying external 
to the mandibula. 

In Chlamydoselachus there are two upper and one lower labials, 
and they have been described or figured by Goodey (1910), K. Fur- 
bringer (1903), and Luther (1909 a). The anterior labial gives 
attachment, along the dorsal edge of its anterior end, to a 
ligament which has its origin on the anterior wall of the orbit 
at about the middle of its height. Numerous short ligamentous 
strings run from the antero- Sohal edge of this ligament into 
the tissues of the upper lip, this seeming to indicate that. the 
ligament has been differentiated from an extensive dermal or 


G* 


124 MR. E, PHELPS ALLIS, JUN., ON 


subdermal fascia. The posterior uppér labial lies internal and 
aboral to the anterior one and is closely attached to the musculus 
levator labii superioris, the tendon of that muscle passing ventral 
to the ligament related to the anterior labial and. having its 
insertion on the ventro-lateral antorbital process. 

In a specimen that I have of Mustelus, probably vulgaris, there 
are two upper labials, and they are strikingly similar to those 
shown by Gegenbaur in a figure (1872, fig. 3, pl. 11) said, in the 
explanations of the plates, to be of Mustelus, but referred to in 
the text as of Galews. In this figure the jaws are furnished with 
sharp pointed teeth; and Marshall and Spencer (1881) and 
Ridewood (1895) have each independently called attention to 
the fact that, because of these teeth, the figure is probably 
of Galeus, as it is said by Gegenbaur to be in his text. But, 
curiously enough, in this figure with sharp teeth, and hence 
probably of Galews, there are two upper labials similar to those 
I find in my specimen of MZestelus, identified by the presence of 
pavement teeth, while in the figure said by Gegenbaur (J. ec. 
fig. 2, pl. 12) to be of Galeus, but furnished with pavement 
teeth and hence probably of Jfustelus, there is but one labial. 
Gegenbaur furthermore says (1. c. p. 214) that both Meckel and 
J. Miiller also found but one upper labial in JdZestelus; but 
K. Fiirbringer (1903) has since found two of them in d/ustelus 
equestris. If these labials do not vary in different specimens 
or species of A/ustelus, it is thus probable that Gegenbaur, in 
his figures above referred to, interchanged either the labials 
or the teeth, as well as the names, of the two fishes. 

But, whatever the error in these two figures of Gegenbaur’s 
may be, in my specimen, which has pavement teeth, there are 
two upper labial cartilages, and they both le in the maxillary 
portion of the labial fold. The anterior labial is about twice as 
long and much stouter than the posterior one, and lies external 
and oral to it, as does the anterior labial in Chlamydoselachus. 
Its hind end lies but slightly anterior to the hind end of the 
posterior upper labial and is attached to that labial, but not 
to the mandibular one, by ligamentous tissues. The anterior 
half, approximately, of the anterior labial lies internal to the 
ventral edge of the musculus levator labii superioris, in a pocket 
formed in the tough connective tissues that cover the external 
surface of the palatoquadrate. The labial is freely movable 
in this pocket, and its anterior end hes directly upon the palato- 
quadrate, in a scarcely perceptible depression in that cartilage, 
but is in no way attached either to it or to the chondrocranium. 
The walls of the pocket are differentiations of a dermal or 
subdermal fascia which is somewhat hgamentous in character, 
and which extends antero-mesially and is attached to the 
posterior surface of the nasal capsule; this fascia doubtless 
representing the one from which the ligament attached to this 
labial in Chlamydoselachus is developed. 

The posterior upper labial of J/ustelus is bound by ligamentous 


THE SKULL OF CHIMAIRA. £25 


tissues to the mandibular labial, and ean thus be said to articulate 
with it. From near its hind end a ligamentous strand runs 
antero-dorsally and joins a stouter ligament which arises from 
the median transverse SLi of the musculus adductor 
mandibule, at the inner end of the line of the angle of the gape. 
The ligament so formed is join, ed by a ligamentous slip from 
about the middle of the anterior labial, and then joins and fuses 
in part with the internal surface of the dorsal, maxillary portion 
of the adductor muscle and in part with the clos sely related 
musculus levator labii superioris. <A part of the ligament 
continues forward and is inserted on the peer surface of 
the nasal capsule at the lateral edge of the large surface of 
insertion, on that capsule, of a part of the Mvecuine levator 
labii superioris. There ave thus, in this fish, two Hgamentous 
structures connecting the upper labials with the nasal capsule. 
In Chlama Niners this connection is, as just above stated, 
with the antorbital wall. 

In Vriakis fasciatum there are two upper labials, and they 
closely resemble, in relative size and arrangement, those in 
Mustelus. The hind ends of the two labials lie close together, 
and both are bound by ligamentous tissues to the hind end 
of the mandibular labial, but the posterior upper labial alone 
articulates with that labial. ‘The anterior end of the anterior 
upper labial rests directly upon the external surface of the 
palatoquadrate, as in Mustelus, but it is not enclosed in a 
pocket of the connective tissues of the region. A broad stout 
ligamentous band arises frem the external surface of the palato- 
quadrate, at its oral edge and slightly anterior to the angle 
of the gape, and running antero-mesially across the external 
. surface of the anterior upper labial is inserted on the internal 
surface of the nasal latero-sensory canal, close to the postero- 
mesial edge of the nasal capsule. . 

In two small specimens of Scylliwm canicula I find, as 
Gegenbaur did, but one labial, and it quite certainly corre- 
sponds to the anterior upper labial of J/ustelus, as Gegenbaur 
concluded. It extends forward along the ventral edge of, or 
shightly internal to, the musculus levator labii superioris, and 
its anterior end lies, as does that of the anterior upper labial 
of Mustelus, directly upon the external surface of the palato- 
quadrate, in a slight depression in that cartilage, but 1t is not 
enclosed in a pocket of connective tissue. This end of the labial 
lies directly internal to the posterior portions of the nasal and 
rostral sections of the latero-sensory canals. In one of the two 
specimens, but not in the other, the mandibular labial hooked 
around the angle of the gape, as Luther (1909a@) shows it in 
his figure of Chiloscyllium, this suggesting the possible fusion 
of a much reduced posterlor upper ‘labial with the mandibular 
labial. 

The anterior end of the anterior upper labial is thus, in each 
of these few Selachii, either in contact with or closely related to 


126 MR. E. PHELPS ALLIS, JUN., ON 


the dorsal surface of the palatoquadrate, and it is also either 
attached to, or related to, ligamentous or fibrous tissues which 
are attached to the chondocranium in the nasal region; and 
these are probabiy universal conditions in these fishes. 

In Chimera Hubrecht (1877) describes five cartilages that 
are called by him nasal cartilages, and four that are called 
labials, the nasal cartilages being designated by the letters 
da 9, 1, m, and the labial cartilages by. the letters a, c, d, e. 
The letter 2 is used by Hubrecht “to designate a process of the 
cartilage &, this cartilage accordingly sometimes being referred 
to as the cartilage vn. 

The cartilage £, with its large process 7, is called by Hubrecht 
the Nasenmuschel, and as it certainly represents sume part of 
the ala nasalis of the Plagiostomi, it may be referred to as that 
cartilage. It encircles the antero-mesial or so-called ingress 
nasal aperture, and [ have recently fully described it in this 
fish (Allis, 1917 6). 

The cartilage /, shown in figure 4 (PI. IT.), is a small and irre- 
gular plate of cartilage which lies in the mucous tissues along 
the lateral edge of the process 7 of the ala nasalis, and it extends 
from that process to the inner surface of the cartilage Fg. Ns 
bounds the dorso-mesial edge of the postero-lateral nasal aper- 
ture, and is attached both to the process n and the cartilage fy 
_by connective or fibrous tissues, its point of attachment to the 
latter cartilage lying inmediately beyond the base of the nasal- 
fold process of that cartilage, to be described immediately below. 

The cartilage m is a thin, flat, curved, triangular cartilage 
which is attached by its pointed mesial end to the dorso-mesial 
surface of the ala nasalis (cartilage An) near its external edge, 
and from there extends ventro-latero-posteriorly in the nasal 
portion of the naso-labial fold. A small nodule of cartilage 
mmay be found attached to its latero-posterior edge. 

The cartilages fand g, said by Hubrecht to be found separate 
in Callor haeehae: are fused in Chimera, to form a single 
piece called by Hubrecht both a Lippenknorpelti viiger Sed au 
Nasenfliigelknorpel. The two parts f and g differ slightly in 
coloration and appearance. suggesting that they are of inde- 
pendent o1igin, the part g being a flat plate-like superficial 
cartilage, and the part f a stout rounded and curved rod, which 
arises from the ventro-posterior end of the plate-like portion 
of the cartilage and may be called the pedicel of the entire 
cartilage. The plate-like portion of the cartilage lies, in the 
posterior half of its length, in a nearly horizontal position 
along the outer edge of the nasal capsule, Its dorso-anterior 
half turns dorso- anteriorly across the outer edge of the nasal 
capsule, and there lies’ on the outer surface of that capsule, 
passing latero-posterior to the lateral rostral process and being 
strongly but flexibly attached to that process by ligamentous 
tissues. Approximately at the point where the process turns 
dorso anteriorly, there is a prcminent eminence on its internal 


THE SKULL OF CHIMERA. Le 


surface, and this eminence rests upon the little eminence on the 
dorsal surface of the ala nasalis (cartilage kn) that is described 
in my earlier work (Allis, 1917 6), the two cartilages there being 
strongly but flexibly attached to each other by connective or 
ligamentous tissues. The pedicel of the cartilage turns sharply 
mesially, in a rounded angle, and passing through a notch iu 
the postero-ventro-lateral edge of the nasal capsule runs along 
the postero-lateral edge of the postero-lateral nasal aperture and 
reaches a smail perforation of the chondrocranium that hes 
immediately ventral to the ventro-posterior edge of the fenestra 
nasalis. This perforation of the chondrocranium is filled with 
ligamentous tissues, and to these tissues the foot of the pedicel 
is strongly but flexibly attached, the position of this pedicel 
strongly resembling that of the anterior upper labial in L/etero- 
dontus (Allis, 19176). 

On the ventral edge of the cartilage fy, at about the posterior 
third of the length of its plate-like portion, there is a slender 
eurved process which projects ventrally into a ridge on the 
internal surface of the nasal portion of the naso-labial fold, 
this ridge quite certainly representing the tissues that enclose 
the process (3 of the ala nasalis of the Plagiostomi (Allis, 1917 6). 
Because of its position, this process of the cartilage /y may be 
called the nasal-fold process. It is thinner than the body of the 
cartilage fg, is flexible, and in many instances seems to be a 
primarily independent piece of cartilage that has secondarily 
and not completely fused with the cartilage fg. In several 
instances there was a foramen between the base of this process 
and the body of the cartilage, this foramen giving passage to 
a delicate nerve which was ‘apparently a branch of the nervus 
maxillaris trigemini. In other specimens this nerve passed 
between the process and a crescentic cartilage described imme- 
diately below. The cartilage J is attached to the inner surface 
of the cartilage fg immediately beyond the base of this nasal-fold 
process. 

In the hollow of the curve formed by the posterior edge of the 
nasal-fold process of the cartilage fg and the ventro-posterior 
portion of the body of that cartilage, lies the crescentic piece 
of cartilage just above referred to. This cartilage is of the 
same consistence as the nasgal-fold process, and les, as that 
process coes, in the ridge on the internal surface of the naso- 
labial fold, and it is apparently the caitilage e of Hubrecht’s 
descriptions of Chimera monstrosa. Both it and the nasal-fold 
process of the cartilage fg are shown in one of Luthev’s figures 
of the latter fish (1909 6, p. 37), and K. Fiirbringer (1903) also 
refers to both of them, the two together being considered by 
him to represent the cartilage e of Hubrecht’s descriptions. In 
Chimera collier the crescentic cartilage is connected with the 
cartilage d, which I consider to be the posterior upper labial, 
bya band of tough fibrous tissue which passes across the external 
surface of the cartila ge fy, a slip of the tissues being sent antero- 


128 MR. E. PHELPS ALLIS, JUN., ON 


dorsally along the external surface of the latter cartilage. This 
crescentic’ cartilage and the nasal-fold process of the cartilage fy 
thus seem to both be chondrifications of this fibrous tissue, one 
of them related to the cartilage d (posterior upper labial) and the 
other to the cartilage fg. 

The cartilage fy is thus a strongly curved cartilage which 
encircles the lateral half of the fenestra nasalis, lying against 
the external, and not the internal, surface of the edge of the 
fenestra, and it either sends a process into the ridge on the 
internal surface of the naso-labial fold, or becomes secondarily 
fused with a cartilage that is developed independently in that 
ridge. ‘The lining femlnon’ of the nasal capsule is not attached 
to this cartilage fy, as it is to the cartilage kn (ala nasalis), lying 
wholly internal to it. The cartilage is capable of a swinging, 
dorso-ventral motion around a line passing through its two 
points of attachment to the chondrocranium, the cartilage 
passing backward and forward across the outer, and not the 
inner, surface of the lateral edge of the fenestra nasalis. The 
posterior one of these two motions is impressed upon it by the 
musculus levator anguli oris anterior of Vetter’s (1878) de- 
scriptions, acting bouts through its own tendon and the long and 
slender hg: cient called by aouther (1909 6) the levator carti- 
laginis prelabialis, the contrary motion apparently being caused 
in part by the simple elasticity of the parts, and in part by the 
action of the musculus labialis posterior. The musculus labialis 
anterior, which is attached by one end to the dorso-anterior end 
of this cartilage and by the other to the cartilage d (posterior 
upper labial), would seem to exert its action on the latter rather 
than on the former cartilage. 

Hubrecht says that the process x of Gegenbaur’s descriptions 
of the Selachii, together with that part of the edge of the nasal 
capsule that, in ‘those fishes, bounds the mesial edge of the 
postero- -mesial nasal aperture, corresponds to the cartilage kn of 
Chimera, this latter cartilage thus betng considered by him to be 
cut off from the outer edge ‘of the nasal “capsule, and its process 7 
to correspond to the process # of the Selachii. The cartilage fy 
of Chimera 1s said to correspond to the process 3 of Gegenbaur’s 
descriptions of the Selachii, the cartilage e of Chimera to 
represent the anterior upper labial of the Selachii, and the 

cartilage 1 to be a remnant of that part of the ala nasalis that 
primar ‘ily connected the cartilages kn and fy. 

Both Fiirbringer (1903) and Luther (1909 6) apparently accept 
Hubrecht’s conclusions regarding the homologies of these several 
cartilages of Chimera; but Luther nevertheless, calls the car- 
tilage fg a premaxillary cartilage, which must mean that he 
considers it to be a labial and not a Nasenfliigelknorpel, and 
he neither mentions, nor shows in his figures, the cartilage e. 
Vetter (1878), who knew of Hubrecht’s work only by title, 
describes as a premaxillary cartilage a cartilage that must be 
the cartilage fy of Hubrecht’s descriptions, but he makes no 


THE SKULL OF CHIMARA. 129 


mention of the cartilage e. He shows, in one of his figures, 
a so-called Nasenfliigelknorpel, which is said to be longer than 
the premaxillary cartilage, to lie directly mesial to it, and to be 
derived from the nasal capsule. This cartilage, as shown in 
Vetter’s figure, is not found in Chimera colliei, and 1t 1s not 
shown in Hubrecht’s figures of Chimera monstrosa. Its dorsal 
end corresponds, in position, to the lateral rostral process of 
these fishes, the remainder of it apparently being the cartilage a 
of Hubrecht’s descriptions, or both that cartil ie and the 
cartilage kn. 

My conclusions regarding the homologies of these several. carti- 
lages differ somewhat prow those of these several authors, and 
they are based on my interpretation of the lips and nasal aper- 
tures of this fish as set forth in the work already several times 
referred to as now in press (Allis, 1917 6), and which should here 
be consulted. 

In the Plagiostomi it is always that part of the ala nasalis that 
encircles the antero lateral and ingress nasa] aperture that is the 
most developed, the part that encircles the postero-mesial and 
egress aperture always being less developed and in some cases 
wholly wanting. In Chimera, on the contrary, it is the part of 
this cartilage that encircles the antero-mesial and here so-called 
ingress aperture that is the most developed, that part of the 
cartilage that encircles the antero-lateral and originally ingress 
spertuce having undergone marked reduction. That part of the 
cartilage that encircles the antero-mesial aperture is represented 
in the cartilage An of Hubrecht’s descriptions, the cartilage /: 
representing that part of the ala nasalis of the Plagiostomi that 
lies mesial to and between the processes « and /3 of Gegenbaur’s 
(1872) descriptions of the latter fishes, and the process » of the 
cartilage & representing the process « of the Plagiostomi together 
with the process that Gegenbaur eal}s, 7n Mustelus, the process @’. 
The process (6 of the Plagiostomi is represented in Chimera in 
the little crescentic car tilage that lies in the ridge on the internal 
surface of the naso-labial fold, and a remnant of that part of 
the ala nasalis that originally lay between this process and the 
process & is represented in the cartilage ¢ of Chimera. 

The cartilages m and g of Chimera have no homologues in the 
Plagiostomi, but they, the nasal-fold process of the cartilage g, 
and the little adjacent crescentic cartilage are all evidently of 
fibrous origin and all quite certainly chondrifications of a sub- 
epidermal layer of fibrous tissue. Just what this layer of tissue 
is I have been wnable as yet to definitely determine, but it would 
seem to be the fibrous layer of the corium. The nasal-flap carti- 
lage of my descriptions of Raia (Allis, 1916) is certainly a 
Sy aanincon of this same layer of tissue, and as the process 2 
of the ala nasalis of that fish has exactly the same subepidermal 
position as the nasal-flap cartilage, that process must also be of 
fibrous origin. But, if this process & is of fibrous origin, the alar 
ring, of which it is a process, must also be of similar origin, all 


130 MR. E. PHELPS ALLIS, JUN., ON 


of these cartilages then being chondrifications of a single layer 
of subepidermal fibrous tissue. ‘This fibrous layer certainly 
passed, originally, beneath the epidermal tissues that were modi- 
fied to form the sensory epithelium of the nasal pit, and when 
this sensory tissue was invaginated to form that pit, the fibrous 
layer must have been invaginated with it. The nasal capsule 
might then itself also be a chondrification of this same layer of 
fibrous tissue, and hence not a part of the axial skeleton, as it is 
usually considered to be; and its development in the Plagiostom1 
is decidedly in favour of this assumption. 

The cartilage f of Hubrecht’s descriptions of Chimera, although 
it lies along the edge of the postero-lateral nasal aperture, has no 
relations whatever to the original antero-lateral nasal aperture, 
and hence is not a derivative of the ala nasalis, and its position 
strongly indicates that it is an anterior upper labial. It has 
approximately the position of that labial in /Heterodontus, and 
also that of the single labial of Ceratodus (Allis, 1917 6), and 
the fact that its anterior end is in contact with a part of the 
chondrocranium that is quite unquestionably derived from the 
palato-quadrate is in accord with the conditions that I have 
above described in several of the Selachii, and would be wholly 
exceptional for a cartilage derived from the ala nasalis. If, in 
one of the Selachii above referred to, the fibrous or ligamentous 
tissues that are related to the anterior upper labial, and that are 
attached to the chondrocranium in the nasal region, were to 
undergo chondrification, a cartilage would arise that would 
closely resemble the cartilage ih of Chimera, one part of this 
cartilage thus being of labial origin and the other of independent 
fisveuist origin. Furthermore, tie origin of the cartilage is in 
accord with its relations to the terminal branches of the nervus 
inaxillaris trigemini, for the more important terminal branches 
of that nerve pass internal to the cartilage, between it and the 
nasal capsule, which is not their relations either to any part of 
the nasal capsule of the Selachii or to the ala nasalis of those 
fishes, but is their relations to the anterior upper labial and its 
related ligaments in those fishes. What is apparently the carti- 
lage fin Schauinsland’s descriptions of embryos of Callorhynchus 
(1903, n?, fig. 127) has, however, decidedly the appearance of 
being a part of the ala nasalis, but the cartilages as there shown 
are so different from those in the adult Chimara that I am 
unable to make any comparison. Two of the cartilages de- 
scribed by Schaninsland, called by him US cartilages 0 “an i”, 
are said by him to lie in mucous folds “welche die Schnauze 
umgeben ” and to represent preoral visceral arches. Premandi- 
bular arches they may represent, as may also the labial carti- 
lages of the Selachu, but they certainly cannot represent preoral 
arches. 

The cartilages c and d of Hubrecht’s descriptions were con- 
sidered by him to together represent the posterior upper labial 
of the Selachii. They are first said by him to be separate and 


THE SKULL OF CHIMERA, Lou 


distinct cartilages, capable of a slight motion with each other, 
but they are later said to be so completely fused that there ts 
even but slight persisting indication of the line of separation 
between them. Vetter found them as separate cartilages and 
considered them to be, respectively, the maxillary and mandibular 
labials. K. Fiirbringer found them more independent of each 
other than they are said to be by Hubrecht, but less so than 
described and figured by Vetter, and he agrees with Hubrecht 
in considering them, together, to represent the posterior upper 
labial. Luther shows them as a single cartilage, and they to- 
gether form the maxillary, and hence posterior upper labial of 
his descriptions. I find them as separate and distinct cartilages 
lying immediately posterior to the angle of the supplementary 
secondary gape of the mouth and straddling the line prolonged 
of that angle, that angle lying near the inner end of the line of 
the angle of the gape, as fully described in my work now in 
press (Allis, 1917 6). One of these cartilages thus les dorsal and 
the other ventral to the line of the angle of the gape, in the 
positions respectively of the posterior upper and mandibular 
labials of the Selachii, and I accordingly consider them, as 
Vetter did, to represent, respectively, those labials, notwith- 
standing that they both, and particularly the posterior upper 
labial, seem to be chondr ication: of ae same fibrous layer that 
has given origin to the cartilages g, m, and e of Hubrecht’s 
descriptions. 

The posterior upper labial, thus identified, is a small and some- 
what rectangular cartilage that les immediately dorsal (morpho- 
logically posterior) to the ventro-posterior-lateral end of the 
plate-like portion of the cartilage fy. It is strongly but loosely 
attached, by its ventral (morphologically anterior) end, to the 
latter cartilage, and also strongly but quite rigidly attached, by 
the adjoining, posterior (morphologically ventral) edge, to ‘the 
dorsal end of the mandibular labial. This latter attachment 
allows of but little notion between the two pieces, such little 
motion as there is being latero-mesial, The labial gives inser- 
tion, on its dorsal (morphologically posterior) edge, to the 
musculus levator anguli oris anterior, and either on its external 
surface or its Mier (morphologically dorsal) edge, to the 
musculus labialis anterior, the other end of the latter muscle 
being inserted on the dorso-anterior end of the cartilage /g. 

The mandibular labial (cartilage c) is a stout bar of cartilage 
with a large triangular process on its anterior edge near its ventr: val 
end. The anterior edge of its dorsal end is str ongly attached to 
the posterior upper labial, as just above described. Its ventral 
end lies in the hind end of the labial part of the naso-labial fold. 
Its anterior process passes internal to the hind end of the supra- 
mandibular furrow, and les in supporting relations to the sup- 
plementary secondary lower lip, this process and the nasal-flap 
process of the cartilage g, together with the related crescentic 
cartilage above described, thus being opposed to each other in 


132 MR. E. PHELPS ALLIS, JUN., ON 


these supplementary secondary lower and upper lips. The ante- 
vior end of the anterior process of the mandibular labial gives 
insertion to a tendon of the musculus levator anguli oris pos- 
terior, that tendon passing downward across the internal surface 

the labial to reach its point of insertion, as shown in Luther's 
figure of Chimera monstrosa. Connected with this tendon, and 
as direct ventral continuations of it, there are two small liga- 
ments, one of which joins the ligament / of Luther’s descriptions, 
to be described immediately below, while the other runs ventro- 
mesially along the external surface of the mandibula, internal to 
the musculus labialis posterior, and has its insertion in tough 
fibrous tissues in the median line. The tendon of the musculus 
labialis posterior runs dorsally along the internal surface of the 
mandibular labial, internal (mesial) to the tendon of the musculus 
levator anguli oris posterior, and has its insertion on the internal 
surface of the mandibular labial near its dorsal end, this also being 
as shown in Luther’s figures of Chimera monstrosa. 

The cartilage a of Hubrecht’s descriptions was considered by 
him to be, together with a related ‘* Bandapparat,” the homologue 
of the mandibular labial of the Selachii. Vetter does not men- 
tion this cartilage. .K. Fiirbringer accepts Hubrecht’s statements: 
regarding it as correct. Luther considers this cartilage, alone, 
to be the homologue of the mandibular labial of the Selachii, 
and, following Gegenbaur’s nomenclature, he calls it the pre- 
mandibular labial. The Bandapparat he considers to be an 
independent and superficial structure, and he fully describes it 
in Chimera monstrosa. 

In Chimera colliei, the thick lower lip contains a mass of dense 
tough fibrous tissue which extends the full length of the lip and 
from its oral edge a certain distance posteriorly. This mass of 
tissue completely surrounds the musculus labialis posterior, that 
muscle traversing a canal-like perforation of the posterior portion 
of the tissue without being in any way attached to it excepting 
only at its origin near the symphysis of the mandibles. The 
anterior (oral) end of the cartilage a is strongly attached to this 
tough fibrous tissue, and it abuts against the posterior surface of 
the musculus labialis posterior, the cartilage being grooved to 
receive the muscle and the edges of the groove sometimes pro- 
jecting to such an extent that the cartilage is decidedly Y-shaped. 
External to the symphysial edge of the cartilage, and strongly 
attached to it, there is a small piece of strongly calcified cartilage 
which is grooved on its external surface to lodge a section of the 
mandibular latero-sensory canal. On its symphysial edge the car- 
tilage a gives origin to hgamentous tissues which cross the median 
line and are attached to the corresponding cartilage of the oppo- 
site side, this apparently being as shown in Luther's figure of 
Chimera monstrosa, but the musculus Jabialis inferior of that 
author’s descriptions of Chimera wonstrosa is not found in my 
specimens of Chimera collier. 

The ligament / of Luther’s descriptions (19096, fig. 28) is 


THE SKULL OF CHIMERA, : 133 


found in’ my specimens of Chimera collie arising from the 
tough fibrous tissues that cover the ventral surface: “of the sym- 
phys sis of the mandibles, and also from the external surface of the 
cartilage a. Running latero- -posteriorly in a curved line, this 
ligament is joined first by a ligament coming from the median 
line posterior to the cartilage a, and then by a ligament coming 
from the ventral edge of the mandibular labial and to which 
reference has just above been made. The ligament so formed 
runs posteriorly across the postero-ventral edge of the mandibula, 
in a slight groove in that edge, and then runs upward along the 
internal surface first of the mandibula and then of the palatc- 
quadrate, and is inserted on a little cartilage which seems’ to 
enone to the spiracular cartilage of Hubrecht’s descriptions 
of Chimera monstrosa, notwithstanding that it lies mueh farther 
from the hind edge of the mandibula. This little eartilage is 
probably a persisting remnant of a mandibular branchial ray 
such as is frequently found in the Selachii and there currently 
called a spiracular cartilage, but 16 cannot be the homologue of 
the spiracular cartilage of ae Batoidei, that cartilage being the 
extrabranchial, or suprapharyngobranchial, of the mandibular 
arch and being represented, in Chimera, in the processus oticus 
quadrati, as already stated. The ligament related to this little 
cartilage may then represent either certain persisting fibrous 
tissues of the mandibular arch, or be a ligament derived from 
certain fibres of the primitive constrictor of the arch such as are 
found in Astrape (Luther, 1909 a, p. 14), and to which I ie 
mde reference in a work now in press on the homologies of 
the muscles related to the visceral arches in the enathostome 
fishes (Allis, 1917 a). If this little cartilage be a persisting 
remnant of a-mandibular branchial ray, thon the cartilage a 
would also seem to be such a remnant. Luther (J. ¢. p. 46) 
considers the cartilage @ to be a chondrification of the membrane 
in which it lies. 


3. Rostral Processes. 


The three rostral processes of the adults of all of the Chime- 
roids are said by Garman (1904) to be attached to the chondyo- 
cranium by ligaments ‘in such a way as to admit of considerable 
movement of their distal extremities up and down,” the evident 
inference being that Garman did not find, in any of these fishes, 
the cartilage of these processes continuous Pick that of the hens 
drocranium. Hubrecht, however, shows all three of these 
processes directly continuous with the cartilage of the chondro- 
cranium, and he suggests that the median process may be the 
homologue of the emocbeall process of the Selachii, and that 
the lateral processes are probably represented, in the latter 
fishes, by ligaments. Schauinsland (1903) refers to these pro- 
cesses, 10 ono of Callor hynchus, as “init dem Schiadel fest 
verbundenen Knorpeln,” and in. his figures he shows all three of 
them as outgrowths of the septal cartilage of lis descriptions 


134 MR. E, PHELPS ALLIS, JUN., ON 


and as directly continuous with it, the processes thus being of 
trabecular origin. Schauinsland says that these processes support 
the rostrum. Garman (1904, p. 252) says there is no rostrum in 
these fishes, but the presence of the rostral processes suggests 
that a rostrum existed in ancestral forms and has become ob- 
solete. Gegenbaur (1898) says that the three rostral processes 
correspond to, but are not directly descended from, the three- 
limbed rostral basket of the Carchariide and Scylliide, this 
implying that they are directly continuous with the cartilage of 
the chondrocranium. Dean (1906) shows the median process 
directly continuous with the cranial cartilage in a six-month 
embryo of Chimera colliei, but says (J.c. p. 129) that: “I am 
inclined to interpret it as an element, 7. e., a fin support, trans- 
posed from a hinder position,” which would mean that it-was not 
primarily a part of the cranial cartilage. The lateral rostral 
processes he says are “later developed into long and separately 
jointed elements.” He considers the median bar of the rostral 
basket of the Carchariide and Scyllide to be probably represented 
in the septal plate of Schauinsland’s descriptions of Callorhynchus, 
and hence not in the median rostral process of that author's 
descriptions. The lateral bars of the rostral basket of the 
Selachi1 are said to be possibly the homologues of the little 
processes s of Schauinsland’s descriptions of Callorhynchus, to be 
later considered. 

In my specimens of Chimera colliei, I find the median rostral 
process articulating with and strongly bound by ligamentous 
tissue to, but not directly continuous with, a slight eminence 
of the chondrocranium that lies on its dorsal surface between 
the two anterior openings of the ethmoidal canal. The lateral 
process of either side arises from a thin and flexible portion of 
the anterior edge of the dorsal wall of the nasal capsule, and it is 
apparently directly continuous with the cartilage of that wall, 
the surfaces, when the process is removed, always appearing 
fractured. The median process runs at first dorso-anteriorly and 
then turns somewhat abruptly ventro-anteriorly, and extends ap- 
proximately to the level of the outer ends of the lateral processes. 
From about the middle of the length of the proximal portion of 
the median process a ligament arises and runs antero-ventrally in 
the median line until it reaches the level of the lateral processes. 
There it spreads dorsally, ventrally and laterally and is lost in 
a layer of tough fibrous tissue that lies internal to the latero- 
sensory canals on the ventro-anterior surface of the snout, and 
forms part of the fibrous layer of the corium, to be later described 
in connection with the ampullary tubules. A stout lgament, 
which extends mesially from the outer end of each of the lateral 
processes, also lies in this tough tissue, and is thus indirectly con- 
nected with the median ligament just above described as well 
as with its fellow of the opposite side. Gegenbaur (1898) found 
these three ligaments represented by cartilage in one specimen 
of Chimera, the ligaments of my specimen thus being capable of 


THE SKULL OF CUIMERA. pESi5) 


undergoing chondrification. The presence of these three liga- 
ments connecting the three rostral processes of Chimera, and 
their possible chondrification, thus suggest that they and the 
processes together represent a rostral basket similar to that 
found in the Carchariide and Scylliide, but everted. In 
certain specimens of JM/ustelus I find similar ligamentous or 
fibrous strands connecting the median and lateral limbs of the 
rostral basket, and if ear a basket were to be everted, by 
the lateral limbs passing antero-ventrally external to the median 
limb, the conditions found in Chimera would apparently arise ; 
the lateral processes of Chimera thus representing the basal 
portions of the lateral limbs of the basket of the Selachii, those 
portions of those limbs being attached to the nasal capsules as are 
the processes in my specimens of Chimera. 

Garman says that, in Chimera monstrosa, the median rostral] 
process “presents the appearance of having originally been 
attached near the nasal capsule, as in Callorhynchus, and otf 
having the basal portion, for a short distance, brought back 
against and fused with the frontal region of the skull.” This is 
certainly also the appearance presented by the process in Chimera 
colliei ; but it is probable that this apparent folding backward of 
this process is due to the marked antero-ventral growth of the 
anterior portion of the naso-ethmoidal cartilage, the rostral stalk 
being carried forward beyond the base of the median rostral 
process. Comparison of Schauinsland’s figure 125 of Callorhynchus, 
giving a median vertical sectional view of an embryo of Callo- 
rhynchus, with the accompanying figure 2 (PI. I.), giving a similar 
view of the adult Chimera colliei, will show that this must be so. 
Comparison with Sewertzoff’s figures of embryos of Pristiurus, 
and with Dean’s figures of embryos of Chimera colliet, then 
further shows that, becatiee of the antero-ventral growth of this 
part of the chondrocranium, the antero-ventral half of the 
fenestra precerebralis has been carried forward on either side 
of the median rostral process, that the edges of the fenestra 
anterior to the process have then fused with each other, and that 
posterior to the rostral process the fenestra has been closed by 
lateral compression of the cranium, the ethmoidal canals of oppo- 
site sides coalescing in the median line and their floors forming 
the roof of the cranial cavity and their roofs a secondary roof to 
the cranium. The fenestra precerebralis is thus completely closed 
in the adult Chimera, but in embryos of Callorhynchus a part of 
it still persists as the opening ¢ of Schauinsland’s figures. 

A stout ligament arises from the dorsal surface of the chondro- 
cranium at the posterior edge of the nasal capsule, there lying 
postero-lateral to the corresponding lateral rostral process. ‘This 
hgament lies at first at right angles to the chondrocranium, and 
as it is stout and stiff, it looks like a short ligamentous horn 
which extends to the internal surface of the fibrous layer of the 
corium above referred to as being related to the tubules of the 
ampullary organs and to be described immediately below. There 


136 MR. E, PHELPS ALLIS, JUN., ON 


the ligament breaks up into numerous branches which spread in 
every divection along the internal surface of the fibrous mem- 
brane. From the anterior surface of the columnar portion of 
the hgament a branch hgament is sent antero-ventrally beneath 
the fibrous membrane. The little cartilaginous eminence s, shown 
in Schauinsland’s figures of Callorhynchus on the dorsal surface 
of each nasal capsule, corresponds approximately, in position, to 
this hgament of Chimera. 


4. Ampulle. 


The ampulle were examined in two specimens. In one of 
these specimens there were five large occipital ampullary pores 
lying in line along the anterior edge of that portion of the 
supratemporal commissure (lateral canal, Cole, 1896) of the 
latero-sensory canals that lies between the posterior ends of the 
supraorbital and infraorbital canals. The tubules that start 
from these pores immediately penetrate a subepidermal mem- 
brane which extends over nearly the entire surface of the head, 
and, lying in that membrane, between external and internal 
layers of it, ran forward dorsal to the orbit and terminate in a 
group of five ampulle that le immediately dorso-lateral to the 
median rostral process, these ampulle also lying between the two 
layers of the subepidermal membrane. Dorso-anterior to the 
eyeball, between it and the supraorbital latero-sensory canal, 
there is a group of twelve supraorbital pores, the tubules from 
which also penetrate the subepidermal membrane and, running 
antero-ventrally, terminate in an equal number of ampulle that 
lie ventral to the ampullee of the occipital pores and, like them, 
between the two layers.of the subepidermal membrane. Anterior 
to these supraorbital pores, in the anterior end of the space 
enclosed between the supraorbital and suborbital latero-sensory 
canals, there are three groups of pores more or less contiguous 
one with the other. In the two dorsal groups together there 
were thirty pores, and in the ventral group twelve pores. The 
tubules from the two dorsal groups penetrate the subepidermal 
membrane, and running dorsally, dorso-posteriorly or posteriorly, 
terminate in ampulle that le between the two layers of the 
subepidermal membrane, close to the ampulle of the supra- 
orbital pores. The tubules of the ventral group perforate first 
the outer and then the inner layer of the subepidermal mem- 
brane and terminate in ampulle that he close against the 
internal surface of that membrane, in the region between the 
dorsal and lateral rostral processes. Ventral to this latter 
group of pores there are numerous pores, many of them minute, 
covering that portion of the ventro-lateral surface of the snout 
that Jies between the suborbital latero-sensory canal and the 
supramaxillary fold, and antero- mesial to the point where the 
supraorbital canal. joins the outer buccalis canal. The tubules 
leading from these pores all run dorsally between the two layers 


THE SKULL OF CHIMERA. 137 


of the subepidermal membrane, and apparently all perforate the 
inner layer of that membrane and terminate in ampulle that lie 
internai to it, in the region between the dorsal and lateral rostral 
processes, ‘The tissues were, however, here so dense and tough 
that the relations of all of these ap: alle to the membrane could 
not be definitely determined. 

The ampulle above described are all innervated by branches of 
the ramus ophthalmicus superficialis trigemini, these branches 
being all given off after that nerve issues from the ethmoidal 
canal through its anterior opening, the branches destined to the 
ampulle that lie between the two layers of the subepidermal 
membrane all perforating the inner layer of that membrane to 
reach the ampulle. ‘These ampulle correspond, in position, to 
the larger one of the two supraorbital groups of Cole’s (1896) 
descriptions of Chimera monstrosa, the smaller group of supra- 
orbital pores of that author's descriptions having approximately 
the position, in Chimera collier, of a group of ampulle that are 
innervated by the nervus buccalis. 

Cole says (1896, p. 655) that the ramus oticus facialis supplies 
“the most ventral of the ampulle opening on the surface by the 
large occipital pores.” No such branches of the ramus oticus 
could be found in my specimens of Chimera colliei, and as the 
ampulle related to these occipital pores all lie on the dorsal surface 
of the snout, at a great distance not only from the related pores 
but also from the branches of the oticus shown in Cole’s figure, 
this would seem to be an error. 

The buccalis ampullze are found in three groups, two inner- 
vated by the inner buccal nerve of Cole’s descriptions and one 
by the outer buccal. The dorsal group of inner buccalis ampuile 
lies on the dorsal surface of the snout, internal to, or slightly 
posterior to, the occipital and supraorbital groups of ophthal- 
micus ampulle, and, like those ampullew, between the two layers 
of the subepidermal membrane, these ampulle having approxi- 
mately the position of the posterior group of supraor bital : am pullee 
of Cole’s descriptions. The tubules of these ampulle run 
postero-ventrally, internal to the ophthalmicus ampulle and 
tubules and between the two layers of the subepidermal mem- 
brane, and with one exception they all open on the external 
surface in the space included between the latero-sensory canals 
innervated by the inner and outer buccalis nerves of Cole’s 
descriptions, the former canal being the one to which I have 
above referred as the suborbital canal. Nine of these pores form 
a sub-group which lies in the dorso-posterior corner of that space, 
while one of them opens somewhat ventro-anterior to that sub- 
group; and associated with the tubules of these pores there is 
the one exceptional tubule, which opens by an isolated pore that 
lies posterior to the posterior hyomandibular latero-sensory 
canal of Cole's descriptions. A second sub-group, of seven large 
pores, lies immediately ventral to the inner buccalis suborbital 
canal, in a large bend at about the middle of its length, and a 

Proc. Zoou. Soc.—1917, No. X. 10 


138 MR. E.,.PHELPS ALLIS, JUN., ON 


third sub-group, of ten pores, dorsal to the outer buecalis canal 
and postero-lateral to the point where that canal is joined by the 
supraorbital canal. 

The second, or ventral group of inner buccalis ampulle has the 
position of the group B?* of Cole’s descriptions, and it also lies 
between the two layers of the subepidermal membrane, about 
midway between the median and lateral rostral processes. From 
there the tubules run postero-ventrally, internal to the tubules 
of the dorsal buccalis ampulla, and open on the external surface 
in a long line which begins at the point where the outer buccalis 

canal falls into the anter ior one of the two hyomandibular canals 
of Cole’s deseriptions and extends along the dorso-anterior edge 
of the latter canal until it reaches the postero-ventral end of the 
supramaxillary fold. There the line of the ampullary pores 
turns dorso-anteriorly along the edge of the supramaxillary 
fold, and so continues nearly to the “aiaukens line. 

The outer group of buccalis ampulle is Cole’s group B*, and it 
lies ventral to the base of the lateral rostral process. The 
tubules of these ampulle all run ventrally and pass internal to 
the supramaxillary furrow. There two tubules separate from 
the others, and running mesial to the external nasal aperture 
open on the external surface immediately ventro- mesial to that 
aperture. The remaining tubules of the group pass postero- 
lateral to the external nasal aperture, and open along the ventral 
edge of the nasal portion of the naso-labial fold. These ampullee 
and their tubules all lie internal to the inner layer of the sub- 
epidermal membrane, but they are surrounded by delicate con- 
nective tissues that would seem to represent the corresponding 
layer of the corium. 

Closely elated to the ampullary pores on the ventral surface of 
the snout, there are several patches of small depressions which 
are the external openings of little glandular structures that are 
of epidermal origin and form little protuberances on the internal 
surface of the epidermis, suggesting undeveloped or degenerate 
ampullee. 

The subepidermal membrane in which, as above described, the 
larger part of the ampullary tubules lie is formed by the outer 
fibrous layer of the corium. The latero-sensory grooves, and the 
calcified cartilaginous rings that partly surround and support 
them, all lie external to this fibrous layer, as Wright (1884, 
p- 263) says that the latero-sensory canals of embryos of dmaurus 
also do to the corresponding layer in that fish. The membrane 
passes external to the three rostral processes, and there encloses 
a median, three-sided sprce which les between the three pro- 
cesses. ‘The membrane is here thick, because of the accumulation 
of the enclosed ampullary tubules and sacs, and the inner layer 
of the membrane is markedly reticulated, the meshes in the 
membrane being traversed by the tubules of certain of the 
ampulle and by the nerves that supply the others. The median 
portion of the three-sided space is filled with loose fatty con- 
nective tissue, and the nervus ophthalimicus superficialis of either 


THE SKULL OF CHIMERA, 159 


side, after issuing from the anterior opening of the ethmoidal 
canal, enters this space and there breaks up into numerous 
terminal branches. The two layers of the membrane extend 
into the supramaxillary fold, and apparently end in its ventro- 
anterior edge. The tubules of the outer buccal group of ampulle 
open on the external surface oral to the supramaxillary fold; and 
they and the related sacs he internal to the membrane that 
lodges. the other tubules, but, as already stated, fibrous sub- 
dermal tissues are found im the lips and the naso-labial folds 
that seem to correspond to this layer of the corium, but they do 
not form a definite membrane. 

In my work on Mustelus (Allis, 1901) no attention was given 
to the relations of the ampulle to this fibrous membrane, But i 
now find, on re-examining wy sections of this fish, that the con- 
ditions there are strictly similar to those in Chimera. In the 
work on Mustelus I came to the conclusion that each ampullary 
pore of the adult fish imdicated, approximately, the place of 
origin of the related ampullary organ, the long ampullary tubule 
of certain of these organs being formed by an exceedingly rapid 
growth of a primarily short tube, that tube being stretched out 
between the two relatively fixed points represented by the 
surface pore and the point where the sensory nerve enters the 
organ. This has been since confirmed by Coggi (1902), and 1s 
further confirmed by the conditions that I have sivee found in 
Chlamydoselachus, the ampulle of that fish all having short tubes, 
and the ampullary sacs all lying immediately beneath the related 
surface-pores. This marked difference in the positions of the 
ampullary sacs in this fish and those in J/ustelus and Chimera 
evidently needs explanation, and it would seem as if it must be 
in some way related to the amount of cranial flexure at the time 
the ampulle are developed. When the cranial flexure is at its 
greatest, those portions of the external surface of the head 
on which the ampullary pores are found in the adult must lie 
anterior or ventral to the curved anterior end of the central 
nervous system, and hence in the region of the future rostrum, 
If the ampullary sacs and the related nerves were well developed 
at this time, it would seem as if the tendency would be to 
hold the sacs there when the cranial flexure was later reduced 
and the brain drawn relatively backward. ‘The dermal tissues 
would, on the contrary, probably retain their relative relations 
to the underlying parts of the brain, and hence also be drawn 
backward ; and if the ampulle had already penetrated the fibrous 
layer of the corium and continued to he in it, their short tubes 
would be drawn out into long tubules lying in the fibrous layer 
of the corium, as is actually the case in MWustelus and Chimara, 
But if the ampullary sacs were not well developed when the 
cranial flexure was at its greatest, thei tubules could not be 
stretched, and it would be the related: nerve strands that would 
be lengthened, as in Chlamydoselachus. This would not, however, 
explain why these organs penetrate the fibrous layer of: the 
corium without wholly perforating 1t, nor why these organs alone 


10% 


140 MR. E. PHELPS ALLIS, JUN., ON 


enter this layer, the latero-sensory organs and terminal buds all. 
lying external to it, as do apparently also the corresponding 
tissues of the ear and eye. 


CONCLUSIONS. 


In the chondrocranium of the adult Chimera the trabecule 
arise from the ventral surface of the parachordals at a con- 
siderable distance posterior to their anterior ends, and they 
project antero-ventrally at a marked angle to the parachordals. 
This shows that there was not only marked cranial fiexure at the 
time the trabecule began to chondrify, but also that this flexure 
was not, as in the Plagiostomi, later greatly reduced ; for that 
there has here been reversion from the conditions found in the 
wlult Plagiostomi, the cranial flexure being first reduced and 
then later reacquired, seems wholly improbable. 'The Holocephali 
must accordingly be descended from some form in which con- 
ditions existed similar to those that are now found in embryos 
of the Plagiostomi, and as these conditions were probably not 
found in any adult form, the Holocephali must be descended 
directly from embryos of the Plagiostomi by conservation and 
modification of the conditions there found. 

The chondrocranium is generally considered to have been 
formed as a cast which lies between the brain, as a core, and the 
external epidermis, the form of the cast depending primarily 
upon the form of the brain. There must accordingly have been 
some potent influence determining, in these fishes, the retention, 
by the forebrain, of the position imposed upon it by the marked 
cranial flexure of embryos. The precocious development of the 
eyeballs, and their shifting forward and mesially dorsal to the 
forebrain, where they are actually found in the adult, would 
doubtless have furnished such an influence; but, unfortunately 
for this assumption, Schauinsland’s descriptions of Callorhynchus 
show that the eyeballs are still widely separated from each other 
in early embryonic stages, and that the midbrain hes between 
them. It would accordingly seem as if the determining influence, 
whatever it may have been, must have been related either to 
a precocious development of the olfactory organs, or to such a 
development of the functional mouth. 

In all embryos of the Plagiostomi the mouth is at first directed 
ventrally, and if it became functional before the cranial flexure 
was reduced, it would evidently tend to remain ventral, this 
possibly explaining why it is found in this position in the adults 
of most of these fishes. If in such an embryo the cranial flexure 
were relatively late in being reduced, or, what is the same thing, 
if the mouth became precociously functional, the perpetuation of 
the conditions then existing might give rise to the relations of 
trabeculae to parachordals actually found in the adult Holocephali. 
The eyeballs would then naturally tend to shift forward, and 
when the head was later compressed latero-mesially they would 
lie dorsal to the forebrain. The other peculiarities of the, 


THE SKULL OF CHIMASRA, 141 


chondrocranium would then arise as a result of the anterior ends 
of the trabecule turning dorsally anterior to the forebrain and 
between the nasal sacs, and then again forward in their terminal 
portions, represented in the median rostral process. 

The peculiar form of the chondrocranium of these fishes would 
not then be primarily due to developmental adjustments to 
feeding habits, but to the acquisition of feeding habits adapted 
to anatomical conditions which were wholly independent of those 
habits. 


literature cited. 


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muscles, and the Peripheral Distribution of certain of 
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——, 1912. “The Branchial, Pseudobranchial and Carotid 
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——. 1913. ‘The Homologies of the Ethmoidal Region of the 
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——. 1904. “ Beitrige zur Morphologie des Skelets der Dipnoer 
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142 MR. E. PHELPS ALLIS, JUN., ON 


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——. 1898. Vergleichende Anatomie der Wirbelthiere mit 
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——. 1909 b. “ Beitriige zur Kenntnis von Muskulatur und 
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EXPLANATION OF THE PLATES. 
The figures are all natural size. 


Pirate I. 


Fig. 1. Lateral view of the head of Chimera colliei, showing the latero-sensory 
canals and the ampnilary pores. 
2. Median view of the bisected skull. 


Puatez II, 


Fig. 3. Lateral view of the skull, with the labial and nasal cartilages in place. 
4. Lateral view of the anterior end of the skull, with the ala nasalis and the 
cartilage Z in place but the other labial and nasal cartilages removed, 
5. The same, with the ala nasalis and cartilage J removed, 
6. Ventro-anterior view of the end of the snout. 


Prafe III, 


Fig. 7. Dorsal view of the skull. 
8. Ventral view of the skull, 
9. Posterior view of the skull. 
Index Letters. 
a, cartilage a of Hubrecht’s descrip- | 9 md, mandibula. 
*  416NS.. - mdl, mandibular labial. 
ab, foramen for nervus abducens. mlf, maudibular labial fold. 
acy, foramen for arteria centralis n, process of cartilage k. 
retin. | ne, nasal capsule. 
acv, foramen for anterior cerebral vein. nlf, naso-labial fold. 
adp, anterior dental plate. o, foramen for nervus opticus, 
aec, anterior opening of ethmoidal obec, outer buccalis latero - sensory 
canal, | canal. 
ant, antorbital process. | oem, foramen for nervus oculomotorius, 
ecr, cavuin cranil. | opf, foramen for nervus ophthalmicus 
e, cartilage e of Hubrecht’s descrip- | profundus, 
tions. oph, foramen for nervus ophthalmicus 
ec, ethmoidal canal. superficialis. 
epsb, foramen for efferent pseudo- | pdp, posterior dental plate. 
branchial artery. pec, posterior opening of ethmoidal 
Jf, foramen for nerve and artery. canal. 
’", foramen for branch of nervus pf, foramen for nervus palatinus 
ophthalmicus. | facialis. 
fa, foramen for nervus facialis. | pfg, perforation for pedicel of carti- 
Jn, tenestra nasalis. | lage fg. . 
fe, foramen for branchofvenanasalis. | pnb, prenasal beak. 
fon, foramen for vena nasalis. pnd, prenasal depression. 
g, cartilage g of Hubrecht’s descrip- pq, palatoquadrate. 
tions. pre, premaxillary lip. 
hme, hyomandibular latero - sensory pst, postorbital process. 
canal. ptf, pituitary fossa, 
hmf, foramen for neryus hyoideo-man- pul, posterior upper labial. 
dibularis facialis. © pv, foramen for pituitary vein. 
hy, hypophysial groove. smaf, supramaxillary fold. 
k, cartilage & of Hubrecht’s descrip- soc, supraorbital latero-sensory canal, 
tions. sos, suborbital shelf. 
lrp, lateral rostral process, sube, suborbital latero-sensory canal. 
m, cartilage m of Hubrecht’s deserip- t, foramen for nervus trochlearis. 
tious, fr, foramen for neryus trigeminus, 


land i 
i” OMEE Rae a 


he 


P2Z51917, 3OULENGER St 


3b J. Green del.et lith. 
1,2. PHILOCHORTUS 'SPINALIS: 6,4) 2 Rais teres 


vee On led a 
. \ \ 


P2ZS.1917 BOULENGER Fi 


TOIMALLL LATS 


LTTE LS BN neal AAT 
LLL, L EKA a RRS 


eB) (@} 
i J.Green del.et lith. 


12,PHILOGHORTUS NEUMANN, 3,4. PD iNT 2 Eines 
5. P AAR DEMG GER. 


— 


ON LIZARDS OF THE GENUS PHILOCHORTUS, 145 


On the Lizards of the Genus Philochortus Matschie. 
By G. A. Boutenerr, F.R.S., F.Z.8.* 


'Received January 27, 1917: Read February 20, 1917. } 


(Plates I. & II.) 


INDEX. 
SYSTEMATIC : Page 
Philochortus Matschie, definition and description 
of the species mn ore ee ee ls 
Ph. intermedius, sp. ., Somaliland .................. 152 


PHILOCHORTUS. 


Latastia, part., Bouleng. Cat. Liz. 1. p. 54 (1887). 
Philochortus Matschie, Sitzb, Ges. nat. Fr. Berl. 1898, p. 30. 


Head-shields normal, save for the occasional absence of the 
interparietal. Nostril pierced between two nasals and bordered 
by the first upper labial or narrowly separated from that shield. 
Lower eyelid scaly, more or less transparent in the middle. 
Collar well marked. Back with two to six longitudinal series 
of large plate-like scales; ventral plates feebly imbricate, with 
Fue LS posterior Borden, smooth. Digits more or less com- 
pressed, with smooth or keeled lamellar scales inferiorly. 
Femoral pores. ‘Tail long, cylindrical. 

Southern Arabia, Eritrea, Abyssinia, Somaliland. 

This genus differs from Lacerta and Latastia in the longi- 
tudinal series of enlarged plate- like seales on the back, an 
approximation to the condition in Por omera, Lach ydr omus, and 
HTolaspis. 

In all the species the parietal foramen is absent, and a few 
small teeth are usually present on the pterygoids. 

The subdigital lamellee vary according to individuals; they are 
usually smooth or with two series of obtuse tubercles ; sometimes, 
however, the tubercles form obtuse keels, and in a female of 
P. neumanni there is a rather sharp keel along the middle. 
Steindachner has already observed that of the two types of 

hardeggeri one has the subdigital lamelle distinctly keeled, 
whilst in the other the keels are scarcely indicated. 

The species grouped under this genus thus afford another argu- 
ment against the unnatural division of the Lacertide into two 
main groups, Liedactyli and Pristidactyli, according to the absence 
or presence of keels on the lower surface of the digits, which the 
state of things in Psammodromus and Seaptira had already led 
me to abandon, 


* Published by permission of the Trustees of the British Museum, 


+ For explanation of the Plates see p. 157. 


146 MR. G. A. BOULENGER ON LIZARDS 


Synopsis of the Species. 


I. Dorsal plates smooth or very feebly keeled; interparietal usually absent or 
separated from the occipital by the parietals meeting in the middle; usually 
4 upper labials anterior to the subocular; 11 to 15 femoral pores on each side. 
30 to 42 dorsal plates and scales across the middle of the 
body ; hind limb not reaching the ear; foot 14 to 12 
times length of head. ee tcee LP. Spinalis Peters. 
28 or 30 dorsal plates and scales across the middle of the 
body; hind limb reaching the car; foot 12 times 
length OF ead. 60... Se5 ovens sinslen'nsshstcenn ems sou! bantlacuha lege aa alle BypaeD UU Iie canemnea heat 


II. Dorsal plates more or less strongly, rarely feebly keeled ; interparieta! in contact 
with the occipital or separated from it by a small shield; usually 5 upper 
labials anterior to the subocular. 


38 to 42 dorsal plates and scales across the middle of the 

body; 12 to 14 large keeled scales in a transverse 

series between the hind limbs; supraoculars 1n con- 

tact with the frontal; 29 to 34 gular scales in a 

longitudinal series; 14 to 16 femoral pores on each 

side; 33 to 35 lamellar scales under the fourth toc.. P. neumanni Matsch. 
33 to 40 dorsal plates and scales across the middle of the 

body; 12 to 18 large keeled scales in a transverse 

series between the hind limbs; supraoculars in con- 

tact with the frontal; 21 to 28 gular scales in a 

longitudinal series; 12 to 18 femoral pores on each 

side; 24 to 30 lamellar scales under the fourth toe.. P. 7itexmedius Bley. 
24 to 30 dorsal plates and scales across the middle of the 

body; 8 keeled plates in a transverse series between 

the hind limbs; supraoculars usually separated from 

the frontal by a series of granules; 25 to 28 gular 

scales in a longitudinal series; 11 to 13 femoral 

pores on each side; 25 to 31 lamellar scales under 

the fourth oe... 0.0.0 4:. cgaccnsenese vans cntssuvssaebucs scutes - ce UOMO eInymrties 


1. PuiLocHortus sprnanis. (PI. I. figs. 1,.2.) 


Lacerta spinatis Peters, Mon. Berl. Ac. 1874, p. 369, pl. —- 
fig 2. 
Latastia spinalis Bouleng. Cat. Liz. ill. p. 57 (1887); Stejneg. 
Proc. U.S. Nat. Mus. xvi. 1894, p. 717; Bouleng. Ann. Mus. 
Genova, (2) xvi. 1896, p. 551; Tornier, Zool. Jahrb., Syst. xxii. 
1905, p. 375; O. Neumann, t.c. p. 395. 

ilead and body feebly depressed. Head 14 to 12 times as long 
as broad, its depth equal to the distance between the anterior 
corner of the eye and the tympanum, its length 4 to 43 times in 
length to vent; snout obtusely pointed, as long as postocular part 
of head, with obtuse canthus. Pileus twice as long as broad. 
Neck as broad as the head or a little narrower. Hind lhmb 
reaching the shoulder or between the shoulder and the ear in 
males, the axil or the shoulder in females; foot 1} to 12 times 
as long as head; toes slender, feebly compressed. Tail 2 to 
24 times as long as head and body. 

Nostril pierced between 3 aves nasals forming a suture 
behind the rostral, the suture 4 to 3 the length of the fronto- 

nasal, which is men broader than long and ee: than the 
far ecnactal space ; prefrontals forming a short median suture; 
frontal as long as its distance from the end of the snout, 13 to 


OF THE GENUS PHILOCHORTUS, 147 


12 times as long as broad, angular in front, much narrower 
behind, not or but very feebly grooved in front; parietals 1} to 
14 times as long as broad, meeting on the median line, the inter- 
parietal being small and separated from the occipital, or alto- 
gether absent *; occipital small}, triangular. Two large subequal 
supraoculars, preceded and followed by a small one (first and 
fourth), often broken up into two or more small shields or 
eranules, the first in contact with the frontal; 6 or 7 super- 
ciliaries, separated from the supraoculars by a series of granules. 
Lower eyelid opaque or somewhat translucid, with feebly enlarged 
scales in the middie, some of which are deeper than the others. 
Rostral not entering the nostril; a single postnasal; anterior 
loreal. shorter than the second}; 4, rarely 5, upper labials 
anterior to the subocular, which is usually narrower beneath 
than above. <A long narrow upper temporal, usuaily in contact 
with the fourth supraocular, followed by one or two small shields ; 
upper temporal scales small and granwar or hexagonal, lower 
larger ; a more or less distinct tympanie shield. 

4 pairs of chin-shields, first two or three in contact in the 
middle; 21 to 29 gular scales between the symphysis of the chin- 
shields and the median collar-plate, granular in front, gradually 
or abruptly enlarged and imbricate behind, those of the last row 
often as large as the plates of the collar; no gular fold. Collar 
with strongly serrated edge, composed of 7 to 10 plates. 

Scales granular on the nape; 2, 4, or, rarely, 6, rows of 
hexagonal plates along the back, usually smooth, sometimes very 
feebly keeled; these plates rather small and usually merging 
gradually into the granular, flat, smooth, or obtusely keeled 
scales on the sides; 30 to 42 plates and scales across the middle 
of the body. Ventral plates in 6 longitudinal and 27 to 32 trans- 
verse series, the border of the transverse series notched between 
the plates, the median pair of which 1s narrower than the others. 
Preanal plates small and irregular, or two median enlarged. 

11 to 15 femoral pores on each side. 27:-to 30 lamellar scales 
under the fourth toe. 

Caudal scales forming Beer longer and shorter whorls, 
upper oblique, rather strongly and diagonally keeled, rounded or 
obtusely pointed behind, lower feebly keeled. basals aioothi: 24 
to 28 scales in the fourth or fifth whorl behind the postanal 
granules. 

Young dark brown or black on the body, ad 6 yellowish- 
white longitudinal streaks, the median pair bifurcating on the 
nape, the outer branch extending to the superciliary edge, and 
uniting on the base of the tail; the upper lateral streak from the 
eye to the tail, passing above the tympanum, the lower from 


* Absent in the type-specimen and in most of the 59 specimens from Ghinta 
examined by me. J was wrong in thinking the absence of this shield in the type- 
specimen to be an individual anomaly. 

+ Divided into two in the type-specimen, 

ty Absent in the type-specimen. 


148 MR. G, A. BOULENGER ON LIZARDS 


the upper lip to the thigh, passing through the tympanum and 
above the fore limb, reappearing on the back of the thigh; upper 
surface of head pale brown, wath or without darker mottling; 
upper surface of fore limb pale brown, the shields dark-edged, of 
hind limb brown with round white spots ; lower parts white; tail 
coral-red. ‘These markings may entirely disappear in the adult, 
or the dark spaces between the light streaks may be replaced by 
longitudinal series of dark brown or black spots on a yellowish- 
brown ground, 


Measurements, in millimetres. 
1. 2. 3. 


From end of snout to vent ......... 55. 49 52 

‘ PE fore limb ... 19 17 18 
Headend base ct elec ones teem, 12 12 
Width of herd Pee rn een a ih 
Depth of head’ (ie.cccersts oe OO 6 6 
Kore limb: o:2.0.sece ccc eee ee 17 17 
Fanaclimil: aos... acest eee ho ee 31 31 
Ph OOGE Ma ne ee Oe eee Las 16 16 
alts bo eee ete SIR Oe es a 122 123 


1. &, Rugdeia Sogheira. 2. ¢, Ghinda. 3. 9, Ghinda. 


Particulars of specimens examined. 


I =2: 8h 24. a6; 6. 7. 


6, Rugdeia Sogheira, Adal... 55 34 29 8 26 13-12 29 
ee Gruhn akc at ee pS 34 29 9 21 1415 30 
a Pm eae tar. repr asses VY) 33 27 9 24. 12 27 
as BaD ofS Se ae nat eA 37 29 ef 22 14-15 28 
3 se dcaiishsosah anaes Stan ee OO 37 32 8 29 15 29 
n Ton -cdesiaduaite: Soceiy eee UDO OGRE coo mene eno 12 28 
Ree lu cn eh oe ya igs Babette ae ee aes an 


1. Length to vent (in millimetres). 2. Plates and scales across middle of body. 
3. Transverse series of ventral plates. 4. Plates in collar. 5. Gular scales in a 
straight line between symphysis of chin-shields and median collar-plate. 6. Femoral 
pores (right and left). 7. Lamellar scales under fourth toe-—Same tabulation for 
the following species. 


Habitat. Eritrea. The type-specimen is from Bogos. 

I have examined the type-specimen and one from Adal in 
the Berlin Museum, two from Rugdeia Sogheira, and 59 from 
Ghinda, in the Genoa Museum. 


2. PHILOCHORTUS PHILLIPSI. (PI. I. figs. 3, 4.) 


Latastia phillipsii Bouleng. Ann. & Mag. N. H. (7) i. 1898, 
Di Lol: 


Head and body rather depressed. Head about 13 times as long 
as broad, its depth equal to the distance between the centre of 
the eye and the tympanum, its length a little more than 4 times 
in length to vent; snout obtusely pointed, as long as postocular 
part of head, with obtuse canthus. Pileus twice as long as broad. 


OF THE GENUS FHILOCHORTUS, 149 


Neck a little narrower than the head. Hind linb reaching the 
ear-opening ; foot 14 to 13 times as long as head ; toes slender, 
compressed, ‘T'ail 24 to 23 times as long as head and body. 

Nostril pierced between 3 shields; nasals forming a suture 
behind the rostral, the suture } to 3 the length of the fronto- 
nasal, which is much broader than long and broader than the 
ferearial space; prefrontals forming a short median suture ; 
frontal as long as its distance from the end of the snout, nearly 
twice as long as broad, angular in front, much narrower behind, 
feebly 2 erooved ; parietals 13 times as long as broad, meeting on 
the median line between the narrow or very small interparietal 
and the small triangular occipital. 4 supraoculars, first small, 
divided into two and in contact with the frontal, second and 
third large and subequal, fourth smali and sometimes divided 
into two; 5 or 6 superciliaries, separated from the supraoculars 
by a series of granules. Lower eyelid transparent in the middle, 
with feebly enlarged scales, some of which are deeper than the 
others. Rostral not entering the nostril; a single postnasal ; 
anterior loreal shorter than second; 4 upper labials anterior to 
the subocular, which is a little narrower beneath than above. 
A long narrow upper temporal, in contact with the fourth 
supraocular, followed by two small shields; upper temporal 
scales small and granular, lower large; a distinct tympanic 
shield. 

4 pairs of chin-shields, first three in contact in the middle; 
21 gular scales in a straight median line, granular in front, 
gradually enlarged and imbricate towards the collar, which is 
formed of 9 plates. 

Scales flat, granular on the nape, subimbricate and smooth or 
faintly keeled on the body, with the two series on the spine much 
enlarged and plate-like, hexagonal, and twice as broad as long ; 
28 or 30 plates and scales across the middle of the body. Ventral 
plates in 6 longitudinal and 26 transverse series, the border of the 
transverse series notched between the plates, the median pair of 
which is narrower than the others. ‘Two enlarged preanal plates, 
one in front of the other, with smaller plates anteriorly and 
laterally. 

13 femoral pores on each side. 27 to 30 lamellar scales under 
the fourth toe. 

Caudal scales forming alternately somewhat longer and shorter 
whorls, upper oblique, rather strongly keeled, obtusely pointed 
behind; 20 scales in the fourth or fifth whorl. 

Young dark brown above, with 5 yellowish longitudinal streaks, 
the median bifurcating on the nape, each branch extending to 
the middle of the posterior border of the parietal shield; the 
lateral streaks occupying the same position as in P. spinalis. 
Adult grey above, with 3 lemon-yellow longitudinal streaks, the 
median forked on the nape, and with crowded black dots forming 
a band from the temple to above the hind limb; limbs speckled 
with black. Lower parts white. Tail red in the young. 


150 MR. G. A. BOULENGER ON LIZARDS 


Measurements, in millimetres. 


On 

From .endiof sioub torventi 5; 4%. nee oe 
53 fore lhnb:<..5 see ee, 

Haka, waciditteena ds 2 batts dhaln de uanecdi eR MRCP ee ee ae 
Widthofihead: 2 da Geseaieiioipe ad Rat Soy HCE oe es EE 
Depthrorihend: cc2 4h eeate staat, ata te tae mn 
Fore Si pase ace os tos als ae eee ee el 
Bind Tiina. % sos cosas wetack po ence ne oon Oe ee ee 
1 U0 (0) ra eee RM eye ee dee ei Ta mcd TR 


This species is known from two specimens obtained at Berbera, 
Somaliland, by Mr. E. Lort Phillips. Distinguished from the 
preceding by the longer foot and by the light streaks being in 
odd number on the body. 


3. PHILOCHORTUS NEUMANNI. (PI. IT. fig. 1.) 

Philochortus newman Matschie, Sitzb. Ges. nat. Fr. Berl. 
1393, p..30: 

Latastia neumannt Anders. Proc. Zool. Soc. 1895, p. 643, 
pl. xxxvii. fig. 1; Steind. Denkschr, Ak. Wien, Ixix. 1901, aa 330, 

Latastia hardeggeri, part., Anders. Proc. Zool. Soc. 1901, i 
p. 145. 

Head and body feebly depressed. Head about 12 times as long 
as broad, its depth equal to the distance between the centre of 
the eye and the tympanum, its length 4 to 43 times in length to 
vent ; snout obtusely pointed, as long as poctoeal uw part of head, 
with ‘bbuse eanthus. Pileus twice as long as broad. Neck as 
broad as the head or a little narrower. Hind limb reaching the 
collar or between the collar and the ear; foot 12 to 14 times as 
long as head; toes slender, compressed. ‘Tail 2 2 to near ly 3 times 
as long as ‘head and body. 

Nostril separated from the upper labial and the postnasal by a 
narrow rim *; nasals forming a suture behind the rostral, the 
suture 7 to 4 the length oF the frontonasal, which is much 
broader than lone and ioreaves than the iternen ial space; pre- 
frontals forming a short median suture; frontal as long as its 
distance from the end of the snout, 14 to 2 times as long as 
broad, angular in front, narrower behind, not grooved ; parietals 
14 to 1$ times as long as broad; interparietal small, usually 
separated from the occipital by a el shield. 4 supraoculars, 
first small and usually in contact with the frontal, second and 

third large and subequal, fourth small and sometimes broken up 

into two or three; 6 or 7 superciliaries, separated from the supra- 
ocular's by a series of granules. Lower eyelid somewhat trans- 
parent, with feebly enlarged scales in the middle, some of which 
ave deeper than the others. Rostral not entering the nostril ; 
2 single postnasal; anterior loreal shorter than the second; 5 


* Exactly as in Lacerta perspicillata. 


‘OF THE GENUS PHILOCHORTUS. ED1 


rarely 4*, upper labials anterior to the subocular, which is a little 
narrower beneath than above. A long narrow upper temporal, 
usually in contact with the fourth supraocular, followed by one 
or two small shields; temporal scales mostly hexagonal, very 
small above, larger beneath ; a narrow curved tympanic shield, 

5 pairs of chin-shields, first three in contact in the middle; 29 
to 34 gular scales ina straight ne in the middle, granular in 
front, gradually or abruptly enlarged and imbricate towards the 
collar; no gular fold. Collar with strongly serrated edge, com- 
posed of 7 to 10 plates. 

Scales granular and smooth on the nape, rhombic, juxtaposed 
or subimbricate, and obtusely keeled on the body ; 4 or 6 series 
-of hexagonal plates along the back, more or less strongly keeled ; 
38 to 42 plates and scales across the middle of the body f; 12 or 
14 large keeled scales in a transverse series between the hind 
limbs. Ventral plates in 6 longitudinal and 28 to 31 transverse 
series, the border of the fetiovet se series feebly notched between 
the plates, the median pair of which is narrower than the others. 
Preanal plates small and irregular, or two large one in front of 
the other, or three large forming a triangle. 

14 to 16 femoral pores on each side. 33 to 35 lamellar scales 
under the fourth toe. 

Caudal scales in alternating somewhat longer and_ shorter 
whorls, upper oblique, str ongly and diagonally “keeled, rounded 
or obtusely pointed behind, lower keeled ; 26 to 30 scales in the 
fourth or fifth whorl behind the postanal granules. 

Brown to blackish above, with six yellowish-white streaks, two 
along the back, diverging on the nape, and two on each side, the 
upper from behind the eye to the base of the tail, passing above 
the tympanum, the lower from the upper lp, through the tym- 
panum and above the fore limb, to the base of the thigh ; the 
dark band between the two light lateral streaks sometimes 
spotted with whitish; upper surface of head and limbs pale 
brown ; lower parts white; tail pale brown above, yellowish 
beneath, orange-red distally i in the young, 


Measurements, in millimetres. 
i. 2. 3. 


From end of snout to vent .......... 73 82 80 

5 fore limb .... 29 29 28 
ere) TER AUC re Aes Ae Ee a Fs 19 18 
Width an nends Re ace Me eetetais Toe 12 dat 
Depth or head. co. tcce seni  8'S 10 9 
orem iclesces sear ater ccvecs “2O 30 26 
Hand Gina) 2.2.3. eo eee 48 53 47 
Foot.. ST dha eek sce le A 28 25 
As Ss ae Eg be ee 0S 190 200 


k. 3.3 Mt. MeniES 2 9 Me Manif. 3. ¢, Lahej. 


* Tn the female from Mt. Manif. 
+ Anderson’s count, 47, is taken higher up the bedy. 


152 MR. G. A. BOULENGER ON LIZARDS 


Particulars of specimens examined. 


mE Oe SOs aos 6. 7. 


G., Mt. Manif7.... 73 38 30 df 33 «15-16 34 
ts x Ae agin p Pats 38 29 ff 29 14 34 
Hore tc ) spires 50 39 88 38 a4 hay eee 
Or avahe), cniaceeeO 42 31 8 31 16 33 
Het.,,,0 sc. 68) 40 (30 10 “932 — aoa 


The type-specimen, preserved in the Berlin Museum, has been 
compared with the specimens from Lahe}j. 

Habitat. This species is only known from South-Western 
Arabia inland of Aden, specimens having been obtained at Lahe}j 
and in the Wadis below Mt. Manif, near Lahej. 


4, PHILOCHORTUS INTHERMEDIUS, sp. n. (PI. II. figs. 2, 3.) 


Latastia hardeggert (non Steind.), Bouleng. Ann. & Mag. N. H. 
(7) 11. 1898, p.-130. 

Latastia hardeggert, part., Anders. Proc. Zool. Soc. 1901, ii. 
p. 145. 

Head and body rather depressed. Head 12 to 12 times as long 
as broad, its depth equal to the distance between the anterior 
corner or the centre of the eye and the tympanum, its length 
4 to 44 times in length to vent in males, 44 to 5 times in 
females ; snout obtusely pointed, as long as postocular part of 
head, with obtuse canthus. Pileus 2 to 2; times as long as 
broad. Neck as broad as the head or a little narrower. Hind 
limb reaching the elbow or the axil in females, the axil, the 
shoulder, or the collar in males; foot 14 to 14 times as long as 
head; toes slender, compressed. ‘Tail 2 to 22 times as long 
as head and body. 

Nostril separated from the upper labial and the postnasal by a 
narrow rim; nasals forming a suture, behind the rostral, the 
suture } to 2 the length otf the frontonasal, which is broader 
than long and broader than the internarial space; prefrontals 
forming a short median suture; frontal as long as its distance 
from the end of the snout or a little shorter, 13 to 2 times as 
long as broad, angular or rounded in front, narrower behind, not 
or but feebly grooved ; parietals 14 to 12 times as long as broad ; 
interparietal 2 to 3 times as long as broad, in contact with the 
occipital, which is usually broader and about half as long. 
4 supraoculars, first small, sometimes broken up into 2 or 3, and 
as often as not in contact with the frontal, second and third 
large and equal or second a little longer, fourth small and usually 
broken up into 2 or 3; 6 or 7 superciliaries, separated from the 
supraoculars by a complete series of granules, or first in contact 
with the first and second supraoculars. Lower eyelid somewhat 
transparent, with feebly enlarged scales in the middle, some ot 
which may be deeper than the others. Rostral not entering the 


OF THE GENUS PHILOCHORTUS. Las 


nostril; a single postnasal; anterior loreal shorter than the 
second; 5, rarely 4*, upper labials anterior to the subocular, 
which is narrower beneath than above. <A long narrow upper 
temporal, usually in contact with the fourth supraocular, usually 
followed by one or two small shields; temporal scales very small 
and granular above, much larger beneath; a curved tympanic 
shield. 

4 pairs of chin-shields, sometimes followed by a small fifth, 
first three in contact in the middle; 21 to 28 gular scales in a 
straight line in the middle, granular in front, gradually or 
abruptly enlarged and imbricate towards the collar; no gular 
fold. Collar with strongly serrated edge, composed of 7 to 10 
(exceptionally 4) plates. 

Scales granular and smooth on the nape, rhombic and keeled 
on the body ; ; 2 or 4 series of hexagonal plates along the back, 
more o1 less strongly keeled, avely faintly keeled; 33 to 40 
plates and scales across the middle of the body; 12 to 18 large 
keeled scales in a transverse series between the hind limbs. 
Ventral plates in 6 longitudinal and 27 to 32 transverse series 
(27 to 30 in males, 30 to 32 in females), the border of the trans- 
verse series feebly notched between the plates, the median pair of 
which is narrower than the others. Preanal plates small and 
irregular, or two or three enlarged ones in the middle, in a series 
or three forming a triangle. 

12 to 18 femoral pores on each side. 24 to 30 lamellar scales 
under the fourth toe. 

Caudal scales in alternately somewhat longer and shorter 
whorls, upper oblique, strongly and diagonally keeled, rounded 
or obtusely pointed behind, lower keeled ; 24 to 28 scales in the 
fourth or fifth whorl behind the postanal granules. 

Young black on the upper sur face of the body, with 6 yellow 
streaks, the median pai diverging towards the occiput, to em- 
brace an additional pair of short eee or each streak bifurcating 
just before reaching the occiput; the upper lateral streak ex- 
tends from behind the eye to the base of the tail, passing above 
the tympanum, the lower from the upper lip, through the tym- 
panum and above the fore limb, to the thigh; head and upper 
surface of fore limbs brown, hind limbs and tail coral-red, throat 
and belly white. 

These markings may persist more or less distinctly in the adult, 
which vary much in colour: the blackish bands between the light 
streaks may be much spotted with whitish, or broken up into 
spots, or the upper parts may be grey, brown, or reddish with 
7 blackish longitudinal streaks (9 behind the occiput) or merely 
with two series of blackish spots on each side, Tail often reddish 
towards the end. 


* 4 in two specimens only. 


Proc. Zoo. Soc.—1917, No. XT. 11 


154 


MR. G. A. BOULENGER ON LIZARDS 


Measurements, in millimetres. 


Li 2. 3. 4. 
From end of snout to vent ......... ae 68 56 85 
~ fore limb... 27 25 20 ae 
Head eh 18 1074 13 17 
Width of ead Jil 11 8 12 
Depth of head 9 8 67519 
Fore limb Och ROR eae RL 24. 18 26 
aimed him 52 oe pew cee 42, 42 29 45 
Foot.. 22 23 16 25 
Tail .. Sy SE “175 180 145 200 
di . anne 213% as Berbera. 4,5. 2 (gravid), Berbera. 
Particulars of specimens examined. 
1. 2. 3. 4. D. 6. 
Os Wiaeeay nti tO 40 30 9 28 16-14 
se os 71 34 2.202 8 %O7  NSSIy 
E B2- 36 "428 © 7 bad ial 
on 3 SEE Are ro LO 38 30 7 26 =14-16 
64 Berberavec® ois oe ae, 43 34 27 9 22 12 
» Inland of Berbera ... 68 40 28 9 27 16 
‘ a 68-87. 99. 99 | Ob. Neate 
. P 56. ©3629 11 eon eed 
2 fy 55 39° 29. Bh beat 
- 55 6386S 28 10 oak 14 
e 5 53 36 29 4. 23 14 
53 * 53 34 28 8 23 14-13 
Aa 55 ol 40 27 9 26 13 
oy . 95) 7) 36. 80, 107 aut 
Ke . 85 8%. 82 Bo eee GEIS 
. . 80. 868 «80. 30m 24s aroead 
. ' 15 B74 30 ae 0 eae Gols 
55 a 71 39 31 10 28 17-16 
i i 67 40, 30° “ge eae aE 
is . 58 34° (30° teow Nenu i a4 
. 56 | (80. 31> “Men a oats 
: S 63:// 38... 808p7 10. 2b A588 
a a 511986 +80 By ~) Bardens 


Habitat. Northern Somaliland at and near Berbera. 


5, PHILOCHORTUS HARDEGGERI. 
Latastia hardeggert Steind. Ann. 


Derails (ple: xaL3, 


Zeal Jaheb., Syst. xxu. 1905, 


(Pl. LL. figs. 4,:5.) 
Hofmus. Wien, 


vil, 13a, 


Bouleng. Zool. Rec. 1893, Rept. P- 23; ‘Tornier, 


p. 375. 


Eremias heterolepis, Boettg. ae Anz. 1898, ve 115y 193: 


Latastia degent Bouleng. 


p. 55. 


Ann. & Mag. N. FLY (7) 


Head and body rather depressed. 
long as broad, its depth equal to the distance between the 
anterior corner or the centre of the eve and the tympanum, 


Head about 12 


x1 eta. 


times as 


OF THE GENUS PHILOCHORTUS, 155 


its length 44 to 5 times in length to vent; snout pointed, as 
long as postocular part of head, with rather sharp canthus. 
Pileus 1? to 2 times as long as broad. Neck as broad as the 
head or a little narrower. Hind limb reaching the shoulder, 
the collar, or between the collar and the ear; foot 12 to 2 times 
as long as head ; toes slender, compressed, Tail 2? to 37 times as 
long as head and body. 

Nostril between three shields, or separated from the upper 
labial and the postnasal by a narrow rim; nasals forming a 
suture behind the rostral, the suture ? to 2 the length of the 
frontonasal, which is much broader than long and broader than 
the internarial space; prefrontals forming a short median suture ; 
frontal as long as its distance from the end of the snout or a little 
shorter, 14 to 13 times as.long as broad, angular or rounded in 
front, narrower behind, distinctly grooved; parietals 1} to 14 
times as long as broad, in contact with the upper postocular ; 
interparietal extremely narrow, in contact with or narrowly 
separated from the very small occipital *. Two large supra- 
oculars, subequal or anterior the shorter, entirely surrounded 
by a series of granules t, of which larger ones represent the 
first and, fourth supraoculars; 5 or 6 superciliaries. Lower 
eyelid somewhat transparent, with feebly enlarged scales in the 
middle, some of which may be deeper than the others. Rostral 
not entering the nostril; a single postnasal; anterior loreal 
shorter than second; 5 upper labials anterior to the subocular, 
which is narrower beneath than above. <A long narrow upper 
temporal, followed by a shorter shield; temporal scales very 
sinall and granular above, much larger beneath ; a small curved 
tympanic shield usually present. 

A pairs of chin-shields, first three in contact in the middle; 25 
to 28 gular scales in a straight line in the middle, granular in 
front, gradually or abruptly enlarged, and imbricate towards 
the collar; no gular fold. Collar with strongly serrated edge, 
composed of 7 or 8 plates. 

Scales granular and smooth behind the occiput, rhombic, sub- 
imbricate, and keeled on the body; 4 or 6 series of hexagonal 
strongly keeled plates along the back, 8 between the hind limbs. 
24 to 284 plates and scales across the middle of the body. 
Ventral piates in 6 longitudinal and 27 to 30 transverse series, 
the border of the transverse series feebly notched between the 
plates, the median pair of which is narrower than the others. 
Preeanal plates small and irregular, or one enlarged. 

11 to 18 femoral pores on each. side. 25 to 31 lamellar scales 
under the fourth toe. 

Caudal scales in nearly equal whorls, upper oblique, strongly 


* A small sbield between the interparietal and the occipital in one young 
specimen. 

+ Inone of the two type-specimens described by Steindachner, the circle of granules 
is incomplete, the supraoculars being in contact with the frontal. 

+ 30 in the type of Hremias heterolepis, according to Boettger. 


11 


156 MR. G. A. BOULENGER ON LIZARDS 


and diagonally keeled, pointed and more or less distinctly 
mucronate behind, lower keeled, except the basals; 22 to 28 
scales in the fourth or fifth whorl behind the postanal granules. 

Young black above with five yellowish-white longitudinal 
streaks, the median bifurcating on the nape, the two laterals 
as in the preceding species; head and limbs brown above, hind 
limbs with round white spots; throat and belly white, lower 
surface of hind limbs and tail coral-red. Adult brown above, 
with a more or less distinct yellow vertebral streak, bifurcating 
on the nape and extending some way down the tail; each of the 
dorsal plates with a small blackish spot; sides of body pale 
greyish brown, with a series of large blackish spots or bars; 
lower parts white. 


Measurements, in millimetres. 


1 2. 3 

From end of snout to vent ......... 70 70 56 
3 53 fore limb ... 23 24, 19 
Feats) A 9th. tay Moos teehee ote: 14 13 
Width-oftherd re eee AO 9 8 
Depth rofrhend 1457... a3 08 sent ip 8 6 
Hore limbs gy.ssnet et cee eee 22 18 
Pindilimnibs.26 2) eee ee eee 38 34 
GOOG. fates cd ee eae me 21 18 
Pails pean Sareea oaceh ch) 170 105y5) 


1. $, Berbera. 2. 9, Berbera. 3. 9, Mandah, type of L. degeni 


Particulars of specimens examined. 
1B Oo 35 4. 5. 6. Vie 


6), Berbera... cen... 70 24, 27 7 25 11 26 
OO ed te eee 70 26 30 Us 28 fel 25 
ty 0h ORE eet PGS. SEM NES trem OR eal sans 
40 Mandal. .2ht0..7 96 24 30 7 26 i 31 


Habitat. Northern Somaliland and Abyssinia. The type- 
specimens, preserved in the Vienna Museum, are from between 
Hensa and Artec, near Harrar. 

This species is distinguished from the two preceding by the 
lower number of plates and scales across the body, and by 
the nearly constant presence of a series of granules between 
the supraoculars and the frontal; also by the light vertebral 
streak. 


List of the Specimens in the British Museum. 


Philochortus spinalis Peters. 
Tae Rugdeia Sogheira, Adal, Eritrea. Marquis O. Antinori. 
2-9. 62 & yg. Ghinda, Eritrea. Signor Ragazzi. 
Philochortus phillipsii Blgr. 
1-2. g &her., types. Berbera, Somaliland. E. Lort Phillips, Esq. 


1-10. 2 & yg,, 
types. 

ll. ¢, type. 

12-30. 62 & yzg,, 
types. 


1. g, type of La- 
tastia degent. 
2-6. 62 & ve. 


Fig.1. Philochortus spinalis, p. 146, 9. nat. size. 
Upper view of head, X 2. 


ig. 


OV G9 LO LY LO bo po 
a 8 


OF THE GENUS PHILOCHORTUS. 


Philochortus newmanni Matschie. 


Lahej, near Aden. 
Wadis below Mt. Manif, N. of Lahej. 


Philochortus intermedius Blgy. 


Wagga, Goolis Mts., 3000-4000 ft., 
Somaliland. 

Berbera, Somaliland. 

Inland of Berbera, up to 400 ft. 


Philochortus hardeggeri Stdr, 
Mandah, Somaliland. 


Inland of Berbera, up to 400 ft. 


157 


Col. Yerbury. 
A. B. Percival, Esq. 


G. W. Bury, Esq. 


E. Lort Phillips, Esq. 
G. W. Bury, Esq. 


Mr. E. Degen. 
G. W. Bury, Esq. 


EXPLANATION OF THE PLATES, 


Pirate I. 


29 2) § 


Side - ,, 


99 ce} 


99 29 
Young, nat. size. 


99 99 
. = Side 8 
bb) 39: 


3 3 Young, nat. size. 


Prats II. 


Philochortus neumanni, p. 150, &, nat. size, 
intermedius, p. 152, 2, nat. size. 
Upper view of head, X 2. 


” 
bP) 22. 
Side 


2) CB) 
39 22) 


phillipsii, p. 148, ¢, nat. size, 
Upper view of head, X 2. 


Scales in middle of back, x 3. 


Scales in middle of back, x 4. 


Scales in middle of back, <s. 
Young, nat. size. 


5 hardeggeri, p. 154, 2, nat. size. 


39 29 


Young, nat. size. 


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ON THE MIGRATION OF WOODCOCK. 159 


9. An Experimental Investigation of the Migration of 
Woodcock breeding in the West of Ireland. By S. R. 
Doveras, M.R.C.S8., L.R.C.P. Lond., Captain I.M.S. 
(retired), F.Z.8., Ist Assistant Bacteriological Depart- 
ment, Medical Research Committee, National Insurance 


Act. 
[Received February 15, 1917: Read March 6, 1917. ] 


To ascertain if the woodeock breeding in the British Islands 
are migratory in their habits has from time to time been the 
subject of investigations. These investigations have usually taken 
the form of marking a number of nestling birds, generally by 
means of metal rings placed round the legs, and collecting the 
data furnished by the recovery of these birds in various 
localities. 

The best known of such experiments which have been 
published up to the present time are :— 


(i.) That made on the Duke of Northumberland’s estate at 
Alnwick and reported by Lord William Percy in ‘ Country 
Life,’ 1909, Feb. 27th. 

(ii.) That made at Baron’s Court, Co. Tyrone, and reported by 
Hamilton in ‘The Field,’ 1908, Oct. 17th, p. 717, and 
Oct. 24th, p. 745. : 

(iii.) Other references dealing with smaller experiments are to be 
found in ‘ British Birds,’ vol. iv. p. 280, and vol. v. p. 186; 
and also in the report of the Aberdeen University Bird 
Migration Inquiry. 


The experiment, the details of which are here given, has been 
carried out at the instigation of Col. W. W. Ashley, M.P., of 
Broadiands, Romsey, Hampshire, and the Palmerston Estates, 
Co. Sligo, who has most kindly given me permission to publish 
the results obtained up to the present time. 

Before giving the actual details of the numbers of birds. 
marked year by year and the numbers recovered, etc., it is 
necessary to describe accurately the situation where the expe- 
riment was carried out, and to give a description of the breeding- 
grounds, together with a few notes on the nesting-habits of 
woodcock 1n this locality. 

Col. Ashley’s property consists of a strip of country lying 
between the Ben Bulben range and the southern shore of the 
Bay of Donegal in the County of Sligo, the position being almost 
exactly 8° 30’ west and 54° 30’ north. 

It is composed largely of small holdings which have been 
vested in the tenants under the various Land Acts, but it also: 
contains a considerable extent of bog-land and some fine coverts 
made up of both hard and soft wood trees. These woods during 
the winter months harbour a considerable number of woodcock,, 


160 CAPT. S. R. DOUGLAS ON 


affording excellent sport: in exceptional years large bags have 
been made, for instance, in January 1892, 92 and 98 cock were 
shot on consecutive days; the more usual bags, however, are 
about 120 cock for two guns in a week’s shooting. 

Woodcock have only comparatively recently nested regularly in 
these parts, for, from information received from a former keeper 
of Col. Ashley’s, Mr. R. Bracken, who had been brought up in 
this part of the country and who has several relatives keepers on 
neighbouring estates, it appears quite certain that a woodcock’s 
nest was considered the greatest rarity before the year 1875. 
Since that date they have become more and more numerous. On 
Col. Ashley’s property, and especially in respect of the principal 
breeding-ground described below, woodcocks’ nests were practically 
unknown before the year 1900. 

The principal breeding-ground is a wood about 150 acres in 
extent, situated about the centre of the property. This wood, 
which was planted about 1830, consists almost entirely of Scotch 
and Maritime fir, and lies within half a mile of the sea. The soil 
is very sandy, and the ground between the wood and the sea- 
shore ts occupied by sand dunes covered with bent grass. 

That part of the wood where most of the woodcock nest is 
occupied by well-grown pine-trees, and is practically destitute of 
any undergrowth, the ground between the tree-trunks being 
thickly carpeted with moss and fallen pine-needles, with here and 
there heaps of fallen sticks. 

The nests are most frequently placed at the foot of a well-grown 
tree ; they consist of a saucer-shaped depression in the moss with 
no more lining than a few pine-needles, and are often partly 
shielded from view by some fallen dead wood. 

The usual number of eggs laid is four, but nests containing 
five have been found. One egg in the clutch frequently fails to 
hatch out. The average of 50 nests is now found every breeding- 
season in this wood. 

The nesting-season commences in March, a nest with eggs 
having been found as early as the fifth of this month, and 
continues throughout the summer until the end of July or the 
beginning of August; however, most of the young are hatched 
in the months of April, May, and the early part of June. 

The young birds grow very rapidly after being hatched out, 
and within two or three days leave the locality of the nest, so 
that the marking of the young birds has to be carried out very 
soon after they are hatched. 

No definite proof of a woodcock raising two broods in the year 
has been obtained, but all the keepers are of the opinion that this 
at any rate occasionally happens. 

The young birds remain near the place of their birth until 
about the middle of September. About this time, however, they 
practically all disappear until the middle of October, when a 
number of birds are again seen, and these appear to constitute the 
regular winter inhabitants of the different coverts. 


THE MIGRATION OF WOODCOCK. 161 


‘The experiment was commenced in the year 1910, and has 
been continued every year since then. The number of marked 
birds that have been recovered varies considerably in different 
years, this being mainly due to there being practically no shooting 
during certain seasons. 

For the first three years the birds were marked by having a 
single ring placed on one leg; this ring was made of aluminium 
and stamped with the lettering. “ A.C. 10” in the year 1910, 
while in 1911 and 1912 the lettering was ‘ W.A A. Sligo, ual ” 
and “ W.A. Sligo, 12” respectively. 

After the year 1912 two rings were used, one ring being placed 
on each leg. The reason for this was that it was ascertained 
that the postal authorities would not deliver letters which were 
addressed with initials only, a fact that may account for the 
searcity of information from outside sources received in the 
earlier years of the experiment. 

The lettering on these rings was ‘“W. Ashley” on the one ring, 
and on the other “ Sligo 13”-14”—15 ”-16,” according as the year 
was 1913, 1914, 1915, or 1916. 

The following table gives the number of birds marked and the 
lettering on the rings for each year :— 


TABLE I, 
Date. Paes Lettering on rings. 
1910. 33 One ring, “ A.C. 10.” 
1911. 48 One ring, “ W.A. Sligo, 1911.” 
1912. 50 One ring, “ W.A. Sligo, 12.” 
1918. 50 Two rings, * W. Ashley” & “Sligo 13.” 
1914, 50 | Two rings, “ W. Ashley” & “Sligo 14.” 
1915. 50 Two rings, ‘‘ W. Ashley” & “Sligo 15.” 
1916. 50 Two rings, ‘‘ W. Ashley” & “Sligo 16.” 


This gives a total of 331 bills marked in seven years. All 
these birds with the exception of seven were hatched in the large 
pine-wood described above. The seven exceptions, consisting of 
two in 1910, two in 1911, and three in 1912, were young birds 
taken from nests in some of the outlying coverts. 

The total number of marked birds that have been recovered 
is 55, that is 16°6 per cent. 

The following table gives (i.) the number of birds recovered 
year by year; (i1.) the locality in which they were found ; 
(ii1.) the date the bird was marked ; (iv.) the date the bird was 
recovered ; (v.) the method of recovery. 

In respect to the various names of places on the estate, 
Classiebawn includes the large pine wood and _ its immediate 


162 


CAPT. 


. R. DOUGLAS ON 


surroundings, Cloonkeen, Carnduff, Bunduff, and Castlegal being 
the local names of various portions of Col. Ashley’s estate, and all 
of them lie within 3 miles of the principal breeding-ground, To 
simplify the reading of the table the letter (.), signifying estate, 
is placed after these place-names. 

For convenience the years are reckoned in shooting-seasons. 

In the case of birds recovered outside the estate the appnommmaye 
distance and bearing are given. 


TaBueE LI. 


heey ie No. of mtg aes and ta ee Date and method of 
July 3st. ag marked. hs ales 8 
1910-1911. | 1, Hazlewood, an estate 10 miles 1910. Nov. 1910. Shot. 
to the south. : 
I *Cloonkeen (WH: exec 5.'4 sacs ees 1910. Feb. 1911. Shot. 
ADIT =VO Il, Castlecali(li.)) (gr esavece. esc 1911. .| Nov. 1931. Shot: 
i Carnduil (Hoa et ete seas nee: 1911. | Nov. 1911. Shot. 
4, Classiebawn (1.).....60.. 0080 crs TOM: Nov. 1911. Shot. 
1; Castleral (Ea 75. 1911. Dec. 1911. Shot: 
be @lassvepbawMebiain.tenease eae meee 1911. | Jan. 1912: Shot. 
TC@loomkeen (Ch ice.cacin sue ete 1911. Jan. 1912. Shot. 
le\Castlepalls(W)y cam. it -cccosssoan ee eee 1911. | Jan. 1912. Shot. 
i Cloonkeen (6 )s. pesos cee sens 1910. | Jan. 1912. Shot. 
1912-19138. | 4, Classiebawn (E.).......0........0000 1912. Dec. 1912. Shot. 
3. Cloonkeen (Moje cere ticsss car teres 1912. Dec. 1912. Shot. 
| 1, Cloonkeen (E.)..00.0.....nc00.0.| 911.) Jan. 19182 > Shot: 
| 1, Classiebawn (KE. ) DORR ee Race 1911. June 1913. Found dead. 
lee Classiebawany CH) eter nee ees 1910. | June 1913. Found dead. 
1913-1914. | 1, Classiebawn (E.) .............0.c00-+ 1911. | Jan. 1914. Shot. 
:8, Classtebawm (H.) 320% tees. cssens 1913. Jan. 1914. Shot, 
1, Bunduff (E.) 1912. | Jan. 1914. “Shot. 
| re Buna wilds) ees ee tere 1913. Jan. 1914. Shot. 
| 1, Hazlewood, an estate 10 miles 1913. Jan. 1914. Shot. 
| to the south. 
1914-1915. | 1, Morga, Biscay, Spain, about 1914. | Nov. 1914. Shot. 
| 800 miles due south. 
1, Castlegore, Co. Mayo, about 1914. | Dec. 1914. Shot. 
| 40 miles to south-west. 
| Lo Glencarss@o. sligOm sa.:.0 cee. cae 1914. Feb. 1915. Shot. 
14, Classiebawmt (Ha) ee.1 ie fone: 1914. April 1916. Found dead, killed 
| by a hawk. 
1915-1916. | 8, Classiebawn (H.) ...........0......-5- 1914. May 1916. Found dead, killed 
| by a hawk. 
1916-1917. 1, Bridgetown, Co. Donegal, 60 miles} 1918. | Nov. 1916. Shot. 
to the north-east. 
1, Mullins, Co. Donegal, 15 miles to} 1915. | Nov. 1916. Caught in rabbit- 
the north-east. trap. 
i @lassiebawim (Mo) strc aceeens 1910. | Jan. 1917. Shot. 
lM Classiebawn (ls) 2h0.c8esres oe 1913. | Jan. 1917. Shot. 
i Classiebawany (Hi) 02.14. vedas eee heres 1915. | Jan. 1917. Shot. 
6;:Classiebawmi (HN)... casccenten ee 1916. | Jan. 1917. Shot. 
EP Carnduiii(bs) i o..ccccsccaccoee se 1914. | Jan. 1917. Shot. 
1, Carndutff (E.) 1915. “| Jan. 19172-" Shot 


THE MIGRATION OF WOODCOCK, 163 


Besides these, two other ringed birds have been reported, but 
were not actually obtained by any responsible person. 

One with one ring only was caught by a cat in 1915, and 
therefore was a bird marked in 1910, 1911 or 1912; the other 
with rings of the year 1916 is said to have been shot by a 
poacher in 1916. | 

On looking over this table the most striking fact is the large 
number of marked birds which have been recovered on the estate, 
the actual number being 48 out of the total of 55 birds recovered, 
and of these no fewer than 33 were obtained either in the large 
wood forming the.main breeding-ground or its immediate vicinity. 

Of the seven birds which have been recorded as obtained in 
other situations three were shot within a radius of ten miles in a 
southerly direction, two at Hazlewood. and one at Glencar, about 
6, 7, and 9 months after being marked. 

One was shot at Castlegore, Co. Mayo, which is about 40 miles 
to the south-west, about six months after being marked. One 
was noticed in the market-place of Bilbao, having been shot at 
Morga, in the province of Biscay, Spain, within six months 
of being marked, the distance in a direct line being about 
800 miles. 

It is therefore to be noticed that all the birds which have been _ 
recovered south of the breeding-place have been obtained within 
9 months of their being marked. 

Two birds were recovered north of the breeding-place, one 
about 15 miles to the north-east in the townlands of Mullins, 
Co. Donegal, 14 years after being marked, and the other near 
Bridgetown, Co. Donegal, 60 miles to the north-east, 33 years 
after being marked. 

The small number of birds recovered during the years 1914— 
1915 and 1915-1916 is accounted for by the fact that owing to 
the war there was very little shooting in these years. 

Table II. (p. 164) gives (i.) the year, (ii.) the number of birds 
marked, (iii.) the number of birds recovered, (iv.) the date the 
birds were recovered. _ 

This table is imserted so as to bring out some of the data 
already given in a more convenient form; one point to which 
attention may be drawn is that a bird marked in 1910 was shot 
in Jan. 1917, so that it must have been at least 64 years old. 

On examining this bird no evidence of old age could be 
detected, and its weight, which was 13 oz., was rather above 
the average. 

With regard to the weight of woodcock, on two occasions when 
a considerable number of ringed birds have been obtained, the 
weights of these birds have been compared with the weights 
of the unmarked birds obtained on the same day. Thus, on 
Jan. 2nd, 1914, nine ringed birds and eight unringed birds were 
shot. 

The weights of the nine ringed birds were :—11, 11:5, 11°5, 
11°75, 11°75, 12°25, 12°25, 12°75, and 13 ounces respectively, 


giving an average of 11°97 oz. 


164 CAPT. 8S. R. DOUGLAS ON 


TABLE ITI. 


No. of birds: | No. of birds Dates the birds were 
marked. recovered. | recovered. 


1910. | 33 


Or 


1, Nov. 1910. 
1, Feb. 1911. 
1, Jan. 1912. 
1, June 1918. 
1, Jan. 1917. 


1911. 48 13 | 6, Nov. 1911. 
1; Dee. 1972, 
3, can. 1912, 
1, Jan., 1913: 
1, June 1913. 
1, Jan... 1934; 


1912. | 50 8 | 7, Dec. 1912. 
| |} 1, Jan. 1914. 


1913. 50 12 110, Jan. 1914. 
1, Nov. 1916. 
1, Jan. 1917. 


1914, 50 8 | 1, Nov. 1914, 
, 1, Dec. 1914. 
1, Feb. 1915. 
1, April 1915. 
3, May 1916. 
| L Jans aon 


1915. 50 3 1, Nov. 1916. 
2, Jan. 1917. 


1916. 50 6 6, Jan. 1917. 


The weights of the eight unringed birds were :—10, 11, 11, 
11°5, 11°5, 11:5, 11°75, and 12°25 ounces respectively, giving an 
average of 11:3 oz. 

Again on Jan. 15th, 1917, eight ringed and sixteen unringed 
birds were shot. The weights of the ringed birds were :—11°25, 
12, 12°25, 12°25, 13, 18, 13°5, and 14°25 ounces respectively, 
giving an average of 12°7 oz. 

‘Those of the sixteen unringed birds were :—10°5, 11°5, 11°5, 
11°75, 11°75, 12,-12, 12, 12°25, 12°25, 12:5, E303, sao, 
and 14 ounces respectively, giving an average weight of 12°3 oz. 

The following day 31 unringed birds gave an average weight 
of 12°1 oz. 

Although these numbers are not great, it 1s worthy of note 
that on both occasions the average weight of the ringed birds 
was greater than that of the unringed birds. 

These being the data obtained up to the present time, is it 
justifiable to draw any conclusions from them ? 

The first point, and this may be taken as conclusively proved, 


THE MIGRATION OF WOODCOCK. 165 


is that many of the woodcock which are hatched out in this 
locality remain there throughout the following winter months. 

The second point is that at any rate a proportion of the wood- 
cock hatched out in this locality remain in the neighbourhood for 
several years, and, even if they at some period or other migrate 
to other places, they return to nest: this is indicated not only by 
the recovery of several marked birds during the nesting-season, 
but also by the fact that on more than one occasion a nesting- 
bird flushed from the nest has been noticed to be a ringed bird. 

The third point is that five woodcock have been recovered in 
localities situated at various distances south of the breeding- 
centre, all within a few months of their being hatched out of the 
egg, and, although the numbers are few, 1t indicates that some of 
the birds tend to migrate south in the autumn months. 

A fourth point worthy of notice is that the woodcock appears 
to show a decided preference for a large-sized pine-wood free from 
undergrowth, as a nesting-ground, £0: any other type of covert. 

The fifth point is that on two occasions a number of ringed 
birds have given a larger average weight than a number of 
unringed birds, indicating that the “resident birds” are rather 
heavier than the winter migrants. 

In conclusion, the data already collected point to their being 
three classes of woodcock in this part of the west of Lreland, 
namely : 


(i.) Woodcock that are hatched out and remain in this locality, 
sometimes for years, that is “resident birds.” 


(ii.) Woodcock that are hatched out in this locality and then 
migrate in a southerly direction. 


(iii.) Woodcock that arrive from the north during the winter 
months. 


These classes are comparable to those into which Eagle Clarke 
divided the starlings of Shetland. 

Finally, let me point out most emphatically that any success 
which this experiment may have achieved in elucidating the 
habits of woodcock is largely due to the care and zeal with which 
Col. Ashley’s keepers, Messrs. Jules, Arthur and Walter Bracken 
have carried out their work. Their knowledge and powers of 
observation have furnished many of the points of interest 
recorded in this paper. 


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ON PRE-JURASSIC TETRAPODS, 167 


10. A Sketch Classification of the Pre-Jurassic Tetrapod 
Vertebrates. By D. M. 8. Warson, M.Sc, F.Z.S., 
Lecturer in Vertebrate Paleontology in University 
College, London. 


[Received February 20, 1917: Read March 20, 1917. ] 


(Text-figures 1 & 2.) 


INDEX. 
SysTEMATIC :— Page 
PANNA aeat acer ine mecca ae wear eersicaeeten LOO 
Ue Ul etrenter send wrnyrate os Te eee cae eae onnaare tenets Lk 


The enormous expansion of our knowledge of early Tetrapods 
during the last twenty years, which we owe especially to the 
work of Broili, Broom, Case, v. Huene, Moodie, and Williston, 
has led to the general realisation of the inadequacy of our existing 
scheme of classification. 

It is the purpose of this paper to produce a classification of 
these animals which, whilst including all existing information 
and paying attention to the taxonomic views of other students, 
shall be so far as possible a consistent whole expressing my own 
view of the relationships of the forms which fall within its 
scope. 

The difficulties of classification of early Tetrapods are identical 
with those which lead to divergence between those classifications 
of the early Hocene placental mammals characteristic of the 
American and European schools. 

Most American authors, for example, follow Osborn in dividing 
the Lower Eocene Perissodactyls, which are all very much alike 
in structure, between the families of Horses, Tapirs, Rhino- 
ceroses, Lophiodonts, Calicotheres, and Titanotheres, whilst 
European authorities include them all in the one family Lophio- 
dontide, ancestral to all other families of Perissodactyls. 

Both methods are quite legitimate, expressing as they do 
different aspects of the subject. 

Prof. Osborn’s method has the great merit of forcing atten- 
tion to the consideration of the small details which persist 
throughout families, and of bringing out clearly our knowledge 
of actual lines of descent. Its drawbacks are that, without a 
very considerable knowledge not only of one animal, but of its 
descendants, it is impossible to be certain: of its position in the 
system, and that the families are with difficulty, if at all, 
definable. 

The other method, of having large primitive familics ancestral 
to all later lines of an order, has the advantage of emphasising 
the great resemblances between all members of an order in its 
early youth and of giving readily definable families nto which 
any relatively well-known type can be securely placed. It 


168 MR. D. M. S. WATSON ON 


suffers from the disadvantage that whilst emphasising resem- 
blances it is liable to obliterate remembrance or recognition of 
differences. 

As in my opinion the study of early Tetrapods is at present 
suffering from an insufficient appreciation of the differences that 
do exist, the following scheme will follow Prof. Osborn’s method, 
although I fully realise that this course leads to a multiplication 
of Orders and to the placing of many types as incerta sedis. 


BATRACHIA. Macartney, 1802 


Tetrapods which, either in a larval stage or persistently, 
breathe by gills. 


Super-Order LABYRINTHODONTIA (Owen). 


Amphibia with a roofed skull, a lower jaw consisting of at 
least eight bones on each side, and vertebre consisting ote neural 
arches aid intracentra in all fone with pleurocentra in addition 


in most. 


Grade EMBOLOMERI Cope. 


Labyrinthodonts with large well-ossified basioccipital and basi- 
sphenoid. Occipital condyle single or triple. Pterygoids with 
a large palatal part, articulating by movable facets with definite 
basipterygoid processes of the basisphenoid. Interpterygoid 
vacuities very small. Tabulars and dermo-supraoccipitals with- 
out occipital extensions. 


Family ANTHRACOSAURIDA Cope, 1875. 


Embolomeri with a single occipital condyle. No specialised 
sacral vertebre. Clavicles flat plates with parallel anterior and 
posterior margins. A ventral armour of scutes. Primitively 
aquatic. | : 

Anthracosaurus Hux. Lower Carboniferous, Scotland. 

Pteroplax Hancock & Atthey. L.? Coal Measures, North- 
umberland. 

Pholiderpeton Hux. M. Coal Measures, Yorkshire. 

? Hrpetosuchus * Moodie. U. Coal Measures, Kansas. 


Family LoxommMip&, nov. 

Embolomeri with triple occipital condyles and enlarged orbits. 
Post-cranial skeleton net known. ¢ No ventral armour. 

Loxomma Hux. IL. Carboniferous, Scotland. 

‘“ Loxomma allmant.” Lower and Middle Coal Measures, 
Lanarkshire, Fifeshire, Northumberland, Staffordshire. 

Baphetes Owen. Coal Measures, Nova Scotia. 

** Loxomma bohemicum.” L. Permian, Ny¥ran, Bohemia. 


* Name preoccupied. E. T. Newton, 1893, 


PRE-JURASSIC TE!RAPODS. 169 


Family PHOLIDOGASTERIDA, nov. 


Embolomeri showing a passage to Rachitomi? With clavicles 
expanded on the ventral surface. Ventral armour of scutes. 
(Secondarily aquatic ?) 

Pholidogaster Huxley. Lower Carboniferous, Scotland. 


Family Cricotipa Cope. 


Embolomeri with elongated skulls. Clavicles expanded on the 
ventral surface. Ventral armour of small scutes, (Secondarily 
aquatic ?) 

Cricotus Cope. Artinskian, Texas, and ? U. Coal Measures, 
Tilinois. 

Embolomeri incerta sedis. 

Diplovertebron Fritsch. L. Permian, Bohemia. 

Nummulosaurus  ,; - - 


Macromeriwm 55 * a 
Spondylerpeton Moodie. U. Carboniferous, Illinois. 


Grade RACHITOMI Cope. 


Labyrinthodonts with ossified basioccipital and basisphenoid. 
Occipital condyle triple or double. Pterygoids usually with a 
medium-sized palatal part, interpterygoid vacuities of medium 
to large size. Pterygoids articulating with both parasphenoid 
and basisphenoid. Tabulars and dermo-supraoccipitals with 
occipital flanges. Paroccipital always visible from behind. 

Vertebre rachitomous, 7. e., with small paired pleurocentra 
and half-moon shaped intercentra. 


Grade STEREOSPONDYLI Cope. 


Labyrinthodonts with reduced basioccipital and basisphenoid. 
Occipital condyle double. Pterygoids with a reduced palatal 
ramus, interpterygoid vacuities large to very large. Pterygoids 
supported by the parasphenoid. Exoccipital meeting the occi- 
pital flange of the tabular so as to hide the paroccipital in an 
occipital view. 

Vertebre stereospondylous, 7. e., with very reduced or absent 
pleurocentra and enlarged intercentra. 

I have discussed the classification of the grades Rachitomi and 
Stereospondyli in very great detail in a paper which will, I hope, 
soon be published ; it is therefore unnecessary to reproduce the 
division into families on the present occasion. 


Super-Order PH YLLOSPONDYLIA Credner. 


Small, very highly specialised Batrachia with a roofed skull, 
palate with very widely open interpterygoid vacuities, palatines 
and lower jaw very reduced. Coracoid and pubis not ossified. 
Four-fingered hand. Ventral armour of small round scutes. 


Proc. Zoot. Soc.—1917, No, XII. 12 


170 MR. D. M. S. WATSON ON 


Family BRANCHIOSAURID Fritsch. 


With the characters of the super-order. 
Lranchiosaurus Fritsch. Upper Coal Measures and Lower 
Permian, Bohemia, Germany, and France. 


Micrerpeton Moodie. Coal Measures, Llinois. 
Melanerpeton Fritsch. L. Permian, Bohemia and Saxony. 
Pelosaurus Credner. % Saxony and France. 


? Dawsonia Fritsch. - Bohemia. 


Super-Order LEPOSPONDYLIA Zittel. 


Small Batrachia with a roofed skull aa lepospondylous 
vertebre. 

Very few members of the super-order are at all completely 
known, and these differ in many respects. In the structure of 
the skull and lower jaw Batrachiderpeton and Diplocaulus show 
clear resemblances to the Labyrinthodontia, perhaps only owing 
to a common descent from Crossopterygian fish. 


Family Necrripra Miall. 


Lepospondyli with the posterior corners of the skull produeed.. 
Two occipital condyles. Palate with a small parasphenoid and 
small vacuities. ‘Tail with long neural and hemal spines, ex- 
panded and fluted at the ends. Accessory articulating facettes 
between the vertebre. 

Keraterpeton Hux. Coal Measures, Ireland and England. 

Urocordylus Hux. 

Batrachiderpeton Hancock & Atthey. Coal Measur es, En gland. 

? Scincosaurus Fritsch. L. Permian, Bohemia. 

? Oestocephalus Cope. Coal Measures, Ohio: 

? Ptyonius Cope. a 3 

? Sawravus Thevenin. U. Coal Measures, France. 

? Diceratosaurus Jaekel. Coal Measures, Ohio. 

? Crossotelos Case. L. Permian, Oklahoma. 


Family DreLocauLip& 

Lepospondyli with enormously produced corners of the skull. 
Palate with a large parasphenoid and moderate-sized vacuities. 
Tail long, with well interlocked vertebre. 

Diplocaulus Cope. lL. Permian, Texas; U. Carboniferous, 
Illinois. 

Family AIsToPpoDID. 

Legless Lepospondyls. 

Dolichosoma Hux. U. Carboniferous, Ireland; and other 
forms not necessarily closely connected. 


Batrachia incerta sedis. 


Lysorophus (primitive Urodele 2) ; Cardiocephalus, 
Gymnarthrus, etc. 


PRE-JURASSIC TETRAPODS. Lél 


Class REPTILIA. 


It is now impossible to give any definition of the class Reptilia 
which, whilst including all members of the group, will exclude 
all other Tetrapods. The essential feature of a reptile is that it 
can carry out the whole of its life-history on dry land, not pro- 
ducing a gill-breathing larva, and that it is not a mammal or a 
bird. Reptiles lay a shelled egg except in viviparous forms, in 
which the egg is hatched before it is laid. 


Super-Order COTY LOSAU RIA (Cope). 


Reptiles with a roofed skull and _ plate-like pelvis. The 
members of this super-order are merely held together by these 
primitive characters, the typical forms also by many other 
common primitive reptilian characters lost by the advanced 
members of this group. 


Order Seymouriamorpha, nov. 


Cotylosaurs with a skull resembling in nearly all known 
details that of the Anthracosauride. Otic notches small, quad- 
vate inclined backward. Tabulars and dermo-supraoccipitals 
on the skull roof, but with occipital flanges. Pro-otie reaching 
the skull roof. Inner ear widely open to the cranial cavity in 
the lateral wall of the cranium. Vertebre with very heavy and 
expanded arches and very large intercentra. 

Limbs very primitive, like those of the Rhachitomous amphi- 
bian Hryops in many features. 

Seymouria Broili, Artinskian, Texas. 


Order Diadectomorpha, nov. 


Cotylosaurs with exaggerated laterally placed otic notches and 
a vertically placed quadrate. 


Super-Family DiApEctip& Cope. 


Diadectomorphs with a long low brain-cavity. Tabulars and 
interparietal turned down onto the occipital surface, closed post- 
temporal vacuities. Inner ear widely open to cranial cavity. 
Vertebree with heavy neural arches. Limbs primitive. 


Diadectes Cope. Artinskian, Texas. 


Diadectoides Case. ‘, ‘ 

Nothodon Marsh. ? U. Coal Measures, New Mexico. 

Animasaurus Williston. _ ,, 59 a 

? Desmatodon Case. B Pennsylvania, 
Diasparactus Case. ; a New Mexico. 


? 
Chibonyx Cope. Artinskian, Texas. 
? Stephanospondylus Geinitz & Deich. M. Rothliegende, 
Saxony. 
Peg 


2 MR. D. M. 8. WATSON ON 


Super-Family Partasaurip#& Seeley. 


Diadectomorphs with a long low brain-case. Tabulars and 
dermo-supraoccipitals on the dorsal surface. Large post-temporal 
vacuities. Otic notch secondarily obliterated. Inner ear sepa- 
rated from cranial cavity by bone. 

Vertebie with heavy neural arches. Pectoral limb advanced 
in the nearly vertical position of the fore arm in life. Pelvic 
girdle advanced in the expanded backwardly sloping ilium. 


‘Pariasaurus Owen. U. Permian, S. Africa. 
Propappus Seeley. 
Anthodon Owen. - bs ue 
Lradysaurus Watson.  M. Permian, 
Embrithosaurus Watson - 4 
Higinia K. T. Newton. U. Permian, Scotland. 

Pariasuchus Broom. < S. Africa. 

? Sclerosaurus H. v. Meyer. L. Trias, Switzerland. 


ae 29 


99 


Super-Family PRocoLopHonip& Seeley. 


Diadectomorphs with a short high Sphenoden-like brain-case. 
Tabulars partly on the oceipital surface. 

Vertebree with heavy neural arches. 

Fore limb specialised in the loss of the screw-shaped glenoid 
cavity. 

Procolophon Owen. LL. Trias, 8. Africa. 

Telerpeton Mantell. M. Trias, Scotland. 

Koiloskiosaurus v. Huene. L. Trias, Germany. 


Order Capitorhinomorpha, nov. 


Cotylosaurs with an obliterated otic notch and vertically 
placed quadrates. 


Super- Family CAPproRHINIDA, 


Captorhinomorphs with short high brain-cavity, widened 
supraoccipital (inner ear placed low down ?%). Dermo-supra- 
occipitals on the occipital surface. Stapes perforate, with a very 
large foot-plate articulating with pro-otic, paroccipital, basi- 
sphenoid, and basioccipital, and distally reaching the quadrate. 

Limbs primitive. 


Fam. CAPTORHININ#. 
Captorhinus Cope. Artinskian, ‘Texas. 
Labidosaurus ,, es a 


¢ Fam. PaRioricHID&. 
Parvotichus Cope. Artinskian, Texas. 
Lsodectes ie - Re 
Puercosaurus Williston. Artinskian, New Mexico. 


PRE-JURASSIC 'TETRAPODS. 173 


Super-Family LIMNOSCELID#, nov. 


Captorhinomorphs. Brain-case unknown. With primitive 
limbs, carpus and tarsus only partially ossified. 
Limnoscelis Williston. U. Coal Measures, New Mexico. 


Super-Family PAnryLip&, nov. 


Brain-ecase unknown. 
With light neural arches and limbs. 


Pantylus Cope. Artinskian, Texas. 


Super-Order ANOMODONTIA Owen. 


Reptiles with a single temporal vacuity. A short high brain- 
case with the inner ear placed low down. Supraoccipital very 
broad. ‘“Stapes” articulating with the quadrate. Tabulars and 
dermo-supraoccipital on occipital surface. In typical forms the 
lower jaw laterally compressed and with a notched angular. 

Derived from a common ancestor with Captorhinide. 


Order Caseasauria, nov. 


Doubtfully belonging to super-order. 

Skull short, with a single laterally placed temporal fossa, sur- 
rounded by the postorbital, squamosal, ? quadratojugal and jugal. 

Brain-case not well known but apparently considerably different 
from the normal type of the super-order. Stapes extending 
nearly to quadrate. Lower jaw showing a projection of the 
articular on the inner side and perhaps a lateral compression of 
the angular which may lead on to the typical superordinal 
arrangement. 

Casea Williston. Artinskian, Texas. 

¢ Trispondylus Williston. Artinskian, Texas, 


Order Pelycosaurias 


Anomodonts with the postorbital and squamosal meeting over 
the temporal fossa, with a screw-shaped glenoid cavity carried by 
scapula, coracoid, and precoracoid, and primitive limbs. 


Super-Family PoLrosauRID& Case. 


Pelycosaurs with a straight tooth-row and undifferentiated 
dentition. A small supratemporal; stapes articulating with the 
pro-otic, paroccipital, basioccipital, and basisphenoid round the 
fenestra ovale only. Cervical neural arches usually heavy and 
with nearly horizontal zygapophysial facets. 

Varanosaurus Broili. Artinskian, Texas. 


Varanoops Williston. i 5 
? Poliosaurus Case. » 
Pecilospondylus Case. - ” 


¢ Mycterosaurus Williston. ,, 56 


174 MR. D. M. Se WATSON ON 


-Super-Family OPpHIACODONTIDA. 


Pelycosaurs with a curved tooth-row and slightly differentiated 
dentition. A small supratemporal. Stapes articulating with 
the edges of the fenestra ovale and by a special head with the 
under surface of the paroccipital process. Cervical neural arches 
not heavy. 

Deiopeus Cope. Artinskian, ‘Texas. 

Theropleura Cope. e i 

Ophiacodon Marsh. U. Coal Measures, New Mexico. 

? Stereorachis Gaudry. L. Permian, France. 


Super-Family SPHENACODONTIDA. 


Pelycosaurs with a curved tooth-row and well-differentiated 
dentition. 

Stapes of two parts, a small element articulating with the 
fenestra ovale and distally with a large element which articulates 
with the paroccipital process, the pterygoid and quadrate. A 
typieal compressed and notched angular. 


- Family CLEPSYDROPID Cope. 


Sphenacodontide with a carnivorous dentition and a_pro- 
nounced step between the dentigerous edges of the premaxilla 
and maxilla. 

Clepsydrops Cope. Artinskian, Texas. 

Upper Coal Measures, Illinois. 

Dimetrodon Cope. Artinskian, Texas. 

Sphenacodon Marsh. ‘ New Mexico. 

Tetraceratops Matthew. ,, Texas. 

?“ Geosaurus cynodus” Gervais. L. Permian, France. 


Family EDAPHOSAURID& Cope. 


Sphenacodontide with a powerful palatal dentition of small 
teeth. 
Edaphosaurus Cope. U. Coal Measures and L. Permian, Texas, 
New Mexico, Bohemia, Saxony, Urals. 
Naosaurus Cope. L. Permian, Texas. | 


Super-Family Botosauripé. 


Systematic position very doubtful, but as the occiput seems to 
be of Anomodont type and the lower jaw is undoubtedly com- 
pressed and the angular apparently notched, they may be placed 
here. 

Skull with large orbits and very short pre-orbital and temporal 
region, upper surface passing smoothly into the occiput. Tem- 
poral vacuity entirely on the side of skull and placed low down. 


PRE-JURASSIC TETRAPOULS, lye 


Bolosaurus Cope. Artinskian, Texas. 
?Glaucosaurus Williston. Artinskian, ‘Texas. 
? Paleohatteria Credner. L.. Permian, Saxony. 


Pelycosaurs of undetermined position. 


Arribasaurus Williston. U. Coal Measures, New Mexico. 
Bathygnathus Leidy. Permian, Canada. (?% Clepsydropid.) 


Order Deinocephalia Seeley. 


Anomodont reptiles, with unreduced quadrates. No supra- 
temporal element. Postorbital and squamosal meeting above 
temporal fossa. ‘The basicranium forming a deep wall below the 
condyle. Glenoid cavity supported solely by the scapula and 
coracoid in typical forms. Limbs modernised. 


Super-Family TAPINOCEPHALID. 


Deinocephalia with a short skull and shoit mouth. 
Tapinocephalus Owen. M. Permian, 8. Africa. 


Mormosaurus Watson. 3 55 
Pnrigalion Watson. - a 
Lamiasaurus Watson. * > 
Struthiocephalus Haughton. _,, = 
Moschosaurus . " 
Moschops Broom. 3 35 
Moschognathus Broom. re 7 
Taurops m 8 3 
Eceasaurus - es ¥ 
Delphinognathus Seeley. i . 
Deuterosaurus Eichwald. Permian, Russia. 
etc. 


Super-Family TrranosucHID&. 


Deinocephalia with an elongated mouth. 


Titanosuchus Owen. M. Permian, 8S. Africa. 
? Rhopalodon Fischer. . i Russia. 

? Cliorhizodon Twelvetrees. __,, ‘ - 

? Dinosaurus Fischer. a " - 


Order Dromasauria. 


Anomodonts with very long slender limbs. Large orbits, 
short pre-orbital and temporal region, rounded dorsal contour of 
skull, and a T-shaped squamosal. 

¢ descended from Bolosauride. 

Galechirus Broom. M. Permian, S. Africa. 

Galepus 9? 99 9 

Galeops ” ” 20 

¢ Paleohatteria Credner. L. Permian, Germany. 


176 MR. D. M. S. WATSON ON 


Order Dicynodontia. 


Anomodonts with a reduced quadrate ‘and quadratojugal. 
T-shaped squamosal without otic groove. Fenestra ovale carried 
down by along process composed of basioccipital, basisphenoid, 
prootic, and paroccipital. 

Scapula with acromion. 

Glenoid cavity borne only on aca and coracoid. Pelvis 
with a pubo-ischiac vacuity. 


Dicynodon Owen. M. and U. Permian, 8S. Africa and Russia. 


Tropidostoma Seeley. 10 

Endothiodon Owen. M. Permian, S. Mees, 
Cryptocynodon Seeley. U. Permian, = 
Prodicynodon Broom. is ri 
Pristerodon Huxley. ne a 
Gordonia Newton. Be Scotland. 
Geikia of x * 
Kisticephalus Owen. a S. Africa. 


Dielurodon Broom. 
Taognathus  ,, 
Kannemeyeria Seeley. M. Trias, 
Lystrosaurus Cope. L. Trias, 
Eubrachiosaurus Williston. M. Trias, Wyoming. 
Placerias Lucas. M. Trias, Arizona. 


Order Theriodontia Owen. 


Anomodonts with carnivorous specialisation and reduced 
quadrate and quadrato-jugal. Glenoid cavity on scapula and 
coracoid. 


Sub-Order GorGonopsra Lydd. 


Theriodonts with the parietal excluded from the temporal 
fossa. Palate with no suborbital vacuities. 


Gorgonops Owen. M. Permian, 8. Africa. 
Scymnognathus Broom. U. as cs 
Arctops Watson. M. ; 3 
Galesuchus Haughton. M. ¥ Sipe 
Scymnosaurus Broom. M. a a 

? Cynodraco Owen. jy cone <f 

? Tigrisuchus Owen. linea de 
Atlurosaurus Owen. M. ‘ 3, 
Arctognathus Broom. Us ss ‘ 
Inostransevia Amalitzki. U. b Russia. 


Sub-Order CYNODONTIA Owen. 


Theriodonts with a narrow intertemporal bar formed by the 
parietals. Secondary palate with no suborbital vacuities. 


PRE-JURASSIC TETRAPODS, Lid 


Fam. CYNOGNATHID&. 
Cynognathus Seeley. M. Trias, 8. Africa. 


Diademodon ,, se ” 
Trirachodon _ ,, 35 9 
Cynochampsa Owen. 9 


Fam. NyTHOSAURID. 
Galesaurus Owen. M. Trias, S. Africa. 
Nythosaurus ,, 3 A 


Sub-Order Z'vzROCEPHALIA Broom. 

Theriodonts with a narrow intertemporal bar formed mainly 
by the parietals. No secondary palate. Large suborbital 
vacuities. F 

Scylacosaurus Broom. M. Permian, 8. Africa. 


Lycosaurus es 99 ” 

Alopecodon - _ ” 

Hycenosuchus ne 5 9 

Pardosuchus 3 ” ” 
etc. 


Sub-Order BAURIAMORPHA. 


Theriodonts with short temporal vacuities separated by the 
parietals. A secondary palate and large suborbital vacuities. 


Bauria Broom. M. Trias, S. Africa. 
Microgomphodon Seeley. M. ,, = 
Sesamodon Broom. an Oars . 
Melinoden Broom. M4; " 


Super-Order CHELONTIA. 


Reptiles with a roofed skull and the middle eight of the ten 
dorsal vertebre provided with widened ribs, supporting a dermal 
armour. 


Order Eunotosauria. 


?if rightly referred. 
Chelonia with costal plates not fused with the ribs and the 
pectoral and pelvic girdles not within the ribs. 


ELunotosaurus Seeley. M. Permian, 8. Africa. 


Order Testudinata. 


Chelonia with the pectoral and pelvic girdles within the dorsal 
ribs. | 


Family PLEURODEIRA. 


‘Testudinates which withdraw the head sideways. 


178 MR. D. M. S. WATSON. ON 


Family AMPHICHELYDID&. 


Proganochelys Baur. U. Trias, Wiirttemberg. 
Proterocherys K. Fraas. 


39 99 


Super-Order SAU ROPTERYGIA, 


Aquatic or semi-aquatic reptiles with a single temporal vacuity. 
Neck long, the cervical ribs articulating solely with the centrum, 
dorsal ribs single-headed. 

A single coracoidal element. 

Pelvis with an obturator foramen. 


Sub-Order VOTHOSAURLA. 


Seml-aquatic Sauropterygia. Paroccipital expanded and reach- 
ing the squamosal, pterygoid, and quadrate so as to close the 
middle ear behind. 

Radius and ulna, tibia and fibula long bones. 

Nothosaurus Minster. lL. & M. Trias, Germany. 

Cymatosaurus Fritsch. M. Trias, Germany. 

etc. 


Sub-Order PLACODO NTA’ 


Shantou ygia with paroccipital not eeiculaneae with ptery- 
goid and quadrate. Massive crushing teeth on the palatines. 
(Body with a heavy armour of bony scutes.) Limb-bones long. 

Placodus Agassiz. M. Trias, Germany. 

Cyamodus  ,, .. 

Placochelys Jaekel. ns Hungary. 


Order Thalattosauria Merriam. 


Aquatic reptiles with a single lateral temporal vacuity. A 
supratemporal present. A single coracoidal element. Occipital , 
region of skull unknown. Humerus twisted with expanded ends. 


ae ; : : 
Thalattosaurus Merriam. M. Trias, California and Nevada. 
Nectosaurus 


Order Ichthyosauria. 


Aquatic reptiles with a single temporal vacuity, very large 
orbits, and much elongated premaxille. 

Basisphenoid without definite basipterygoid processes, ptery- 
goids underlying much of its lower surface and that of the basi- 
occipital. Neck short. 


Family M1ixosauRID&. 


Ichthyosauria with relatively small orbits and short rostrum. 
Upper end of the scapula expanded. Pubis and ischium broad. 


PRE-JURASSIC TETRAPODS. 179 


Kpipodials long and separated. Caudal region not much 
deflected. 


Fam. Mrxosaurin&. With five digits. 


Mixosaurus. M. Trias, Italy, Germany, Spitzbergen, 
W.N. America. | 


Fam. SHASTASAURIN®, Narrow paddles with three or four digits. 


Cymbospondylus. > 
Toretocnemus. 
Merriamia. M. Trias, Nevada. 


Shastasaurus. 
Delphinosaurus. 


Super-Order ARCHOSAURIA. 

Reptiles with a skull with two temporal openings. Brain-case 
much narrowed between the ears, usually surrounded by bone in 
advance of the prootic. Paroccipital process antero-posteriorly 
compressed and long, with the fenestra ovale opening on its 
lower edge. 

A single coracoidal element. 


Order Thecodontia. 


Primitive Archosaurians with clavicles and an interclavicle. 
Pelvis “ plate-like.” 


Family ‘“‘ KosucHip.” 


Thecodonts with intercentra throughout the column. 


* Kosuchus” Watson. U. Permian, 8. Africa. 
Youngina Broom. 5 5 


Family ERyrtHrosucHID&, nov. 


Very large semiaquatic Thecodonts with a twisted humerus’ 
with much expanded ends. Anterior margin of pubis suddenly 
deflected. Feeble dorsal armour. 


Lrythrosuchus Broom, M. Trias, 8. Africa. 


Family Puyrosaurip# McGregor. 


Large Thecodonts, with an elongated rostrum formed mainly 
by the premaxilla. Dorsal and ventral scutes. 

Mesorhinus Jaekel. L. Trias, Germany. 

Palworhinus Williston. M.? Trias, Wyoming. 

Phytosaurus Jaeger. U. Trias, Wiirttemberg and U.S.A. 

Mystriosuchus Fraas. 3 . 

Rhytidodon Emmons. Toe secre 

¢*Stagonolepis Agassiz. M. Trias, Scotland. 


180 MR. D. M. S. WATSON ON 


Family OrnrirHosucHip# v. Huene. 


Small, slightly built Thecodonts. Pointed skull without pro- 
duced rostrum. Pubis and ischium much produced. 

Hinder limbs longer than the fore. 

? Directly ancestral to Theropodous Deinosaurs. 

Oriithosuchus EK. 'T. Newton. M. Trias, Scotland. 

Huparkeria Broom. . S. Africa. 

Scleromochlus A.S. Woodward. _,, Scotland. 

Sphenosuchus Haughton. U. Trias, 8. Africa. 


Family AErosaurRID& Baur. 


Small Thecodonts with a pointed skull without rostrum, pubis 
and ischium short. Hind limbs not greatly larger than the fore. 
A very heavy dorsal and ventral armour. 

Aetosaurus Fraas. U. Trias, Wiirttemberg. 

? Dyoplax 3 _ cs 

¢Stegomus Marsh. * WAS .As 

¢ Notochampsa Broom. ,, S. Africa. 


Family Howest1p&, nov. 


?Theecodonts with several rows of teeth in the maxilla. 
Howesia Broom. M. Trias, S. Africa. 

? Mesosuchus Watson. a sf 

?¢ Proterosuchus Broom. e ss 


Family ERPETOSUCHIDA, nov. 


Small Thecodonts with a pear-shaped skull, not very elongated 
rostrum. Palate ineipiently secondary, with the posterior nares 
in a deep groove. 

¢ Ancestral to the Crocodilia. 


Erpetosuchus E.T, Newton. M. Trias, Scotland. 


Order Saurischia Seeley. 


Archosauria with the astragalus very closely and immovably 
articulated with the tibia. Pelvis with the pubis and ischia long 
and projecting downward, usually with an ischio-pubic vacuity. 

Thecodontosaurus Riley & Stutchbury. U. Trias, Somerset- 

shire, Swabia, 8. Africa, Queensland ; 
and very many forms representing numerous families, defined 
especially by v. Huene. 


Order Rhynchocephalia Giinth, 


Reptiles with a two-arched skull, a short, high brain-case not 
ossified in advance of the prootic, and large fenestra ovale. Teeth 
on the dentary biting into a groove between the maxillary and 
palatine teeth. 


PRE-JURASSIC TETRAPODS. 181 


Family RHYNCHOSAURIDA. 
With plate-like pelvis. 
Rhynchosaurus Owen. U. Trias, England. 
Hyperodapedon Hux. M.& U. Trias, England, Scotland, India. 


Stenometopon Boulenger. M. Trias, Scotland. 
? Palacrodon Broom. M. Trias, S. Africa. 


Order Proganosauria Baur. 


Aquatic reptiles with long tails. Skulls elongated. Vertebrze 
with small notochordal centra and very heavy arches and no 
intercentra. A single coracoidal element, five distal carpals and 
tarsals. Pelvis plate-like. 

Mesosaurus Gervais. L.? Permian, 8. Africa, Brazil. 

Noteosaurus Broom. S. Africa. 


79. 
Stereosternum Cope. »  . Brazil. 


Order Protorosauria Seeley. 


Lightly built reptiles with pointed skulls and a single tem- 
poral vacuity. A single coracoidal element and a_ plate-like 
pubis. 

There ig not the slightest evidence that the series of small 
Paleozoic reptiles listed below are related to one another. Prof. 
Williston believes Arwoscelis to be related to the lizard ancestry. 
Broomia may also have affinities with the Squamata. <Adelosaurus 
may be related to the Rhynchocephalha. 


¢ Protorosaurus Meyer.  U. Permian, Germany and England. 
2 ( Arceoscelis Williston. Artinskian, Texas. 
° 1 Kadaliosaurus Credner, L. Permian, Saxony. 
Broomia Watson. M. Permian, S. Africa. 
24 Heleosaurus Broom. 
Heliophilus a; Pe <3 
¢Adelosaurus Watson.  U. Permian, England. 
? Aphelosaurus Gervais. L, Permian, Autun. 


39 wy) 


Noves ON THE AMPHIBIAN CLASSIFICATION. 


The superordinal a ore of the Labyrinthodontia, Phyllo- 
spondylia, and Lepospondylia is based on the fact that the early 
members of these groups differ from one another as much as do 
their ultimate descendants, despite the fact that in a number of 
characters such as the gradual development of large interptery- 
goid vacuities, the replacement of basipterygoid processes with 
definite articulating surfaces by expansions of the posterior end 
of the parasphenoid with which the pterygoids are suturally 
united, and the loss of a finger in the hand, the course of 
evolutionary change is the same in the first and last group. The 
structure ot the skull and lower jaw is known to be similar in 


182 


TRIAS 


DEVONIAN /L.CARBONIFEROUS|U.CARBONIFEROUS| BASAL PERMIAN] M.PERMIAN | U.PERMIAN 


SILURIAN 


' MR. D. M. S. WATSON ON 


Text-figure 1. 


--> 


oe 
SO TEREOSPONDYLI 


Capitosaurus 


Microphalidce 


Lydek'kerinide 


1 
i] 
! 
' 
‘ t 
U 
4 
U 


RACH TOMI, 


Eve 
Loxommiide ) 


t 
) 


ELES? 


URO 


w--- Oe ee ee ee ee 


Phylogenetic tree illustrating the relations of the early Amphibia, each group 


being placed in its correct time. 


183 


PRE-JURASSIC TETRAPODS. 


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184 MR. J. M. S. WATSON ON 


the Labyrinthodontia and Lepospondylia, but the resemblance 
may easily be due to inheritance from common ancestors amongst 
the Crossopterygian fish. 

The ordinal division of the Labyrinthodontia marks three 
stages in the evolution of that group. It is founded primarily 
on the skull, which becomes more and more depressed, the basi- 
occipital and basisphenoid being gradually reduced from the quite 
large bones they are in Pteroplax to the small slightly ossified 
tracts lying on the parasphenoid which represent them in 
Stereospondyls. The development of the vertebral column is 
used as a check on this classification. 


NovreEs ON THE REPTILIAN CLASSIFICATION. 


The super-order Cotylosauria is retained simply because of its 
use as a dumping-ground for those primitive reptiles which retain 
a roofed skull. The orders are also probably somewhat unnatural 
groups, and it would perhaps have been more satisfactory to raise 
the super-families to ordinal rank. 

The real classification, 7. e. that into families, is founded as far 
as possible on the characters of the brain-case, which have been 
discussed by me in a series of papers. 

The separation of the super-order Anomodontia is founded on 
its very characteristic brain-case, and for the typical forms on the 
equally characteristic lower jaw. 

The remarkable animal Casea is obviously ordinally different 
from all other well-known forms, its position in the super-order 
Anomodontia is very doubtful, but it is not improbable that it 
represents a very early offshoot from this stock. 

The order Pelycosauria is referred to the super-order Anomo- 
dontia with certainty ; in Dimetrodon the brain-case and lower 
jaw are absolutely typical, and the other forms included are 
obviously allied to this type by the whole of their structure. 

The super-family division is founded on differences in the 
stapedial articulation with the brain-case, which in the three 
suborders seem to form a morphological series. 

The order seems to be self-contained, culminating in Dime- 
trodon and Edaphosaurus, towards which the other types lead. 
At the same time the trend of evolutionary change in the brain- 
case is the same as that which, continued in later times by the 
South-African forms, leads up to the mammals. | 

The super-family Bolosauride is founded for very badly known 
reptiles, which it might perhaps have been wiser to leave as 
Anomodontia incerta sedis. The position of the super-family in 
the super-order depends mainly on the badly preserved lower 

AW. 
The order Deinocephalia is certainly a member of the super- 
order, having the typical brain-case and lower jaw. It is distin- 
guished from the Pelycosaurs by its advanced limbs, shown most 


PRE-JURASSIC. TETRAPODS. 185 


clearly in the loss of the screw-shaped articular surface of the 
head of the humerus, and the corresponding restriction of the 
glenoid cavity to the scapula and posterior coracoidal element 
alone. 

It is separated from other Anomodonts with advanced limbs 
by the retention of the large quadrate. The mode of articulation 
of the stapes shows that it cannot have been derived from any 
Pelycosaur more advanced than a Poliosaurid. 

The order Dromosauria is referred to the super-order by its 
lower jaw and occiput. It is distinguished from all other South- 
African types by the short face, very large orbits, slender limbs, 
and long tail. 

Paleohatteria is only placed here provisionally ; it is certainly 
an Anomodont, and in the structure of its temporal region very 
strongly recalls Galepus. The absence of the posterior coracoidal 
element may only mean that, as in Varanoops, it is unossified, 
although present as a cartilage. 

It will possibly be found, when fuller knowledge of Bolosaurus 
becomes available, that that type is really allied to the 
Dromosaurs, and that the two orders should be combined. 

The order Dicynodontia includes the first known and typical 
Anomodonts. The order is a very compact group, thoroughly 
distinct from all others and of uncertain derivation, perhaps from 
the Deinocephalian stock. 

The order Theriodontia includes many very diverse animals, 
still known almost wholly from skulls. Its members are held 
together by the presence of a reduced quadrate in all of them and 
by a well differentiated carnivorous dentition. 

The suborder Gorgonopsida appears to be ancestral to the Cyno- 
dontia, the Therocephalia to the Bauriamorpha, and it is probable 
that other lines will be distinguished. Each suborder certainly 
contains many families, but in the absence of detailed knowledge 
of the skull and of all knowledge of the post-cranial skeleton in 
most forms, it seems at present useless to found families on the 
dentition. 

I have recently discussed at length the relations of Hunoto- 
saurus to the Chelonia; if rightly referred, it is so much more 
primitive than any other Chelonian that ordinal separation seems 
justified. 

The various animals included in the Sauropterygia differ so 
widely amongst themselves, and the whole group is so distinct 
from all others, that its ordinal rank 1s unquestionable. 

Merriam’s quite distinct order Thalattosauria is still rather 
incompletely known, but its members certainly have only a 
single arch and the temporal region of the skull is reminiscent of 
the early Pelycosaurs. 

The Ichthyosaurs form a very compact group, whose origin is 
quite unknown. 

The inclusion of the two arened reptiles in one super-order is 
now commonly accepted. I have excluded the Rhynchocephalia 


Proc. Zoot. Soc.—1917, No. XIII. 13 


186 ON PRE-JURASSIC TETRAPODS. 


from the super-order because of certain differences in the brain- 
case, Which may, however, be due to the small size of Sphenodon, 
in which alone is the region known. The exclusion is really for 
the purpose of drawing attention to the extreme smallness of our 
knowledge of early two-arched reptiles. 

The small aquatic J/esosaurus and Stereosternum are still un- 
represented by well-preserved skulls. They seem to be definitely 
ordinally distinct, but their affinities are quite obscure. 

Finally, the order Protorosauria is retained merely because it 
is already in existence. Included in it is a series of small 
Permian reptiles which resemble one another in having slender 
limbs and a single coracoidal element. There is no evidence that 
these animals are in reality in any way related. - 

In this classification I have refrained from throwing weight on 
the peculiarly modified 5th metatarsal which occurs in Chelonia, 
Rhynchocephalia, Thecodonts, Crocodiles, Deinosaurs, and Squa- 
mates, because it is difficult to believe that all these forms can 
have beew derived from the same advanced Cotylosaurian ancestor. 
It is perhaps an arboreal adaptation, which may have originated 
separately. Possibly the broadened ribs of Hunotosaurus, which 
recall those of sloths, are also to some extent an arboreal 
adaptation. 

This classification is on the whole consciously conservative, 
but contains many new features, chiefly in the definitions and the 
super-ordinal grouping. An attempt has been made to make 
the structural differences separating orders approximate, having 
regard to the total variation, to those used in separating the 
orders of mammals, but in many cases suborders should perhaps 
be raised to ordinal rank. Super-orders are used to group together 
orders which seem to have had a common origin, 


ON THE VISCERA OF AN OKAPI. 187 


11. Notes on some of the Viscera of an Okapi (Okapiu 


.johnstont Selater). By R. H. Burng, M.A, F.ZS. 
[Received March 9, 1917: Read April 3, 1917.] 
(Text-figures 1-23.) 


INDEX. 
Page 
Structure or Morphology ..scci.0...5sccasavasseoseeeese LST 


During the winter of 1915-1916 certain parts of the viscera of 
an Okapi were sent by Dr. Cuthbert Christy to the Royal College 
of Surgeons for investigation. 

They included the tongue and the soft parts of the anterior end 
of the throat, the contents of the thorax, the stomach, and the 
colic coil of the intestine. 

A superficial examination revealed, as was to be expected, a 
general similarity to the corresponding viscera of the Domestic 
Ruminants, but a more detailed comparison with other genera * 
made it clear that underlying this general Ruminant type of 
structure were many small and individually unobtrusive pecu- 
liarities that pointed definitely to a very close relationship 
between the Okapi and the Giraffe. 


The Tongue (text-figs. 1 & 2). 

The tongue of the Okapi, though of the Ruminant type, 
resembles most nearly that of the Giraffe, both by reason of its 
great length and slenderness (it measures some 14 in, in length 
by from 14 to 2 in. in breadth) and its sharp pointed extremity 
and smooth base. It is thus distinguished from the tongues of 
Antelopes and Deer, the tip of which is as a rule either blunt oi: 
broadly spatulate, and in which the surface of tte elevated base 
or intermolar fixed portion 1s covered with large fleshy papuille. 

It is manifestly dificult to state with accuracy the relative 
measurements of a tongue or the proportions that its different 
parts bear to one another, as they depend on its degree of exten- 
sion; but after comparison with numerous tongues of Antelopes 
and other Ruminants, both fresh and preserved, it seems fairly 
clear that the free part of the Okapi’s tongue is considerably 
more developed relative to the base than is that of other 
Ruminants, with the exception of the Giraffe. 

The form and disposition of the various papille upon the sur- 
face of the tongue differ little from those in the Giraffe, and are 


* Tam indebted to the Zoological Society for material for use in the comparative 
study of the throat and tongue,—Giraffe, Crowned Duiker, Indian Antelope, Black 
Buck, Scemmering’s Gazelle, and Gazella rufifrons. 

Use was also made of preparations in the Royal College of Surgeons Collection of 
the tongue and larynx of many Ruminants, of the heart and lungs of Ox, Sheep, and 
Goat, and of the stomach of a young Giraffe and of several Antelopes and the 
Domestic Ruminants. 

Ji 


MR. R. H. BURNE ON THE 


188 


‘QA1OU [BSSOTSodAFT “TTX ‘oAdou pvosutreyd-ossopy "XJ  “oAtou jeusury “A 
‘gonp Ss, Wowel AA “PA § [Isuoy “2 Soposnut plorkgy-ouI94ys “w-y7'7s + aPOSMA 
jvoSufreyd-op{ys “wyd'gs Sayosnut prokyorsys Sw hy'7s Soposnut prosy-ou.to4s “wrys Spwkyorsys “y'7s foposnut snssopsopAys 
“wm bys ‘ pueps Aleypixeurqus Jo syoup Surqoo[foo “pews * purps Areppixvugus “op eu's §pueps pensuryqns 747s £pueys [eusuty 
-olqjad “76774 § anSu0} Jo vaay poyuwusid Jo qty “d £ eposnut snapropA4s-091d1990 “2U"78"0 faposnur prosyop dua See'ue £ SULOA [RNOUTT 
Gary $spuvps-ydurky "7 $ Srojre peusul| “py Faposnut snssopsody “eps 'y + aposwut ploxqoiuas “wyh faposnul sussopsorues 
uO Suda perry “af Sapided [ensuly [v10yv] JO asury cdf ¢ Saaqae perorg Cos * oposnut plorkyyoorts Sw yzda § L1oyAB prIyoreo “yo 


-wuogsuyol vrdvyoO jo xufieyd pus onsuo} oy} JO MELA [B.10}UT 


‘] oansy-} 297, 


VISCERA OF AN OKAPI. 189 


characteristically Ruminant. ‘The dorsal surface and sides are 
clothed with a close layer of sharp-pointed filiform papillee which 
are more or less cornified, particularly so upon a central area of 
the dorsal surface close behind the tip, where they form a patch 
of small horny recurved spines. 

Fungiform papille are scattered sparsely all over those parts of 
the free extremity and base that are coated with filiform papille 
except, as one frequently sees among Ruminants, along the 
central line of the dorsum. 

Circumvallate papille, to the number of 15 ox so on either side, 
are arranged in two indistinct irregular rows far back upon each 
side of the base of the tongue. This is the normal Ruminant 
position and arrangement, but the indistinctness of the individual 
papille and of the limits of the group asa whole recal] more 
particularly the condition in the Giraffe. 

There are no foliate papille, and, as previously mentioned, the 
base of the tongue is smooth, as in the Giraffe. 


Text-figure 2. 


Lingual papille of Okapia johnstoni. 


In a large number of Ruminants (nearly all Antelopes, the 
Giraffe, some Sheep and Oxen, but not in Deer) parts of the 
free extremity of the tongue are deeply pigmented. This also is 
the case in the Okapi, where the pigmented area includes the 
extreme tip of the tongue and extends some way backwards on 
either lateral surface, gradually fading away (text-fig. 1, p.). 

In the valley between the base of the tongue and the gums, 
there is in all Ruminants a fold of mucous membrane fringed by 
a number of sharp-pointed papille similar in character and 
probably also in function to the conical papille that clothe the 
inner surface of the cheeks. 

A similar fringe of fleshy papilla les beside the base of the 
tongue of the Okapi; but whereas in all the other Ruminants 
examined the papille are simple conical processes flattened from 
side to side and usually recurved like the thorns of a rose, in the 
Okapi they have quite a peculiar shape, being swollen and blunt 
with two three or more short, sharp horny spines projecting 
from their rounded free extremities (text-fig. 2). 


190 MR. R. H. BURNE ON THE 


The Muscles of the Tongue and Pharyna. 


The muscles of the tongue conform in general to the common 
Luminant type, but in the details of their arrangement and 
individual development disclose more points of agreement with 
the Giraffe than with any of the other Ruminants examined. 

A comparison of the accompanying figures (figs. 1, 3, 4,5, 6) 
shows more readily than a verbal description how close the 
similarity between the Okapi and Giraffe in respect of these 
muscles is. Attention may, however, be drawn to the following 
particulars. 


Digastric, Mylohyoid, Geniohyoid, and Genioglossus Muscles. 


Blending of the various muscles in the floor of the mouth to a 
varying degree is typical of Ruminants in general. In the 
majority, 1f not in all genera, the anterior belly of the digastric 
muscle and the mylohyoid are so closely united that where they 


Text-figure 3. 


Outline of the tongue and pharynx of a Giraffe, for comparison with text-fig. 1. 


Lettering as in text-fig. 1. 


blend in the superficial sheet of the mylohyoid it is impossible to 
distinguish the one from the other. The deeper muscles (genio- 
hyoid and genioglossus) are, as a rule, more independent. 
Generally the former has a weak fibrous attachment along the 
central line of the mylohyoid, and very occasionally (e. g. Antilope 
cervicapra) it unites with the genioglossus close up to the sym- 
physis menti, but as a rule it is a well-developed paired muscle 
independent of its neighbours. 

In the Okapi and Giraffe, not only is the association of the 
digastric and mylohyoid peculiarly intimate, but there is also a very 
close fibrous union between the deep layer of the mylohyoid and 
the geniohyoid and a blending of the geniohyoid and genioglossus, 

The geniohyoid is also very much reduced in both these 


VISCERA OF AN OKAPI, 191 


genera, and can only with difficulty be separated into its two 
component halves. 

Tne condition is shown in text-figs. land 3. It will be noticed 
that the belly of the geniohyoid extends only about halfway 
from the-hyoid to the symphysis menti, and is continued upon 
the ventral surface of the genioglossus as a flat tendon from 
which a large part of the latter muscle arises. 


Text-figure 4. 


SRPHM HLM 
be 


\ 
‘ M4 

HY PHM 

THe S.LK THY 

TRAY 


GV, 


ORTH PL 


ST.TH.M 
Dissection of the pharynx of Okapia johnstoni. 


er.c., cricoid cartilage; c7r.ph.m., crico-pharyngeal muscle; h.l.m., hyoideus latus 
muscle; hy.a., hyoidean branch of lingual artery; hy.ph.m., hyopharyngeal 
muscle; /.g., cervical lymph-gland ; s./.a., superior laryngeal artery; st.g.m.?, 
lesser styloglossus muscle; st¢.ph.m., slips of stylo-pharyngeal muscle ; 
th.c., thyroid cartilage; th.b., thyroid body; th.hy.m., thyro-hyoid muscle 
th.ph.m., thyro-pharyngeal muscle. 

X!_ Pharyngeal branches of vagus. X2. Superior laryngeal branch of vagus. 


Other lettering as in text-fig. 1. 


Stylohyoid Muscle. 


The round tendon of origin of this muscle differs a good deal 
in length in different genera of Ruminants. It is relatively short. 
in Antelopes, considerably longer in the Okapi and Sheep, and 
longer again, though not by much, in the Giraffe. 


e 


192 


cr.ar.m., crico-arytenoid muscle ; 


MR. R. H. BURNE ON THE 


Text-figure 5. 


nyetim STPAM 


ST.TH.M 
Outline of the pharynx of a Giraffe, for comparison with text-fig. 4. 


h.tr.m., hyoideus transversus muscle. Other lettering as in text-fig. 4. 


Text-figure 6. 


OT PMP 


CRAR LT: 


CR TY py 
Superficial dissection of the larynx of Okapia johnstoni. 
cr.ph.m., crico-pharyngeal muscle; ¢7.th.m., crico- 


thyroid muscle; hy.ep.m., h yo-epiglottidean muscle; hy.ph.m., ‘hyo- pharyngeal 
muscle; s.d.a., superior laryngeal artery ; st¢.ph.m., stylo-pharyngeal muscle ; 


t., tonsil; th.c., thyroid cartilage; th.ph.m., thyro-pharyngeal muscle. 
IX. Glosso- pharyngeal nerve. X}. Phar yngeal branches of vagus. X?. Superior 


laryngeal nerve. . 


VISCERA OF AN OKAPI. 193 


Occipito-styloideus (text-figs. 7, 8, 9, 0.st.m.). 

This muscle is peculiar to Ungulates, and extends from the 
styloid process of the occipital to the upper portion of the styloid 
bone. 

In the Giraffe and Okapi it was far larger than in any of the 
other Ruminants examined (Antelopes and Sheep), and offers in 
these two genera a striking contrast to the same muscle in 
Antelopes, particularly as regards the area of the styloid bone 
covered by its insertion. | 

In connection with the insertion of this muscle it should be 
noted that the styloid bone of the Giraffe and Okapi differs 
markedly from that of other Ruminants in the form of its upper 
extremity (text-fig. 10). : 


Text-figure 7. 


O.oT.™M 


The upper end of the stylohyal of Okapia johnstoni, inner surface, showing 
the insertion of the occipito-styloideus muscle. 
o.st.m., occipito-styloideus muscle ; s¢.i.m., tendon of stylohyoid muscle ; 
st.ph.m., stylo-pharyngeal muscle. 


In Ruminants in general, the posterior margin of this bone, 
close to its upper end, is produced to form a prominent angular 
process, to the tip of which is attached the tendon of the stylo- 
hyoid muscle. This process stands out more or less transversely 
to the length of the bone and shows a tendency to a downward 


194 . MR. R. H. BURNE ON THE 


curve, forming in some Antelopes (text-fig. 10, B) a veritable 
hook, and in all cases offers a very broad surface for the insertion 
of the occipito-styloideus muscle. 

In the Giraffe and Okapi (text-fig. 10, A) and in the Camels 
and the Perissodactyl Ungulates the process is relatively small, 
and is directed upwards, forming a simple swelling or tuberosity 
of the upper end of the styloid bone. 


Text-figure 8. Text-figure 9. 


0.ST.™M 


BSUS 
Ur oxS 2 


3 
U 


- aie Yyy cy 
[ZB 


-32 


Text-fig. 8—The stylohyal of a Giraffe, showing the occipito-styloideus muscle. 


Text-fig. 9.—The stylohyal of Scemmering’s Gazelle, showing the occipito- 
sty loideus. 
Lettering as in text-fig. 7. 


Salivary Glands. 

The submaxillary gland (text-figs. 1, 3, smaxgl.) is similar in 
position, form, and relative size in the Giraffe and Okapi, and 
is closely comparable also to the same gland in Antelopes and 
Domestic Ruminants. 

In the Giraffe and Okapi the-.gland gives off two main 
tributary ducts, one from its posterior and one from its anterior 
end. The parts drained by these two collecting ducts can be 
.readily and completely separated from one another in the Giraffe, 
but this is not the case in the Okapi. 


VISCERA OF AN OKAPI. 195 


The two tributary ducts unite in the Giraffe ne junction was 
destroyed in the Okapi) close in front of the gland to form a 
single Wharton’s duct (text-fig. 3, Wd.), and there is no reason to 
think that the separate lobule drained by the anterior tributary 
is anything but a part of the submaxillary gland, for the retro- 
lingual gland, which at first sight it might be supposed to be, is 
represented in Ruminants, including the Okapi and Giraffe, by 
a mass of gland-tissue surrounding the anterior extremity of 
Wharton’s duct close behind the symphysis of the jaw, between 
the origin of the genioglossus muscle and the sublingual mucous 
membrane (text-figs. 1, 3, rl.gl.). 

The sublingual sland calls for no comment. The parotid gland 
had been removed with the skin. 


Text-figure 10. 


A B 


Upper end of stylohyal of (A) Okapia and (B) Scmmering’s Gazelle. 


The Blood-vessels of the Tongue (text-figs. 11, 12, 13). 


In his description of the anatomy of the Giraffe, Owen * 
mentions a peculiar plexiform arrangement of the veins close in 
front of the body of the hyoid bone. In this position there is 
generally among Ruminants a large venous anastomosis uniting 
the lingual veins of either side. This anastomosis was strongly 
developed in the Giraffe’s tongue (text-fig. 11, v.a.) dissected for 
comparison with that of the Okapi, and in front of it, upon the 
left side (text-fig. 11, v.pl.), there was some slight branching and 
reunion of the veins, suggesting the plexus mentioned by Owen. 
There was no indication of a plexus i in the Okapi (text-fig. 12). 

Posteriorly, the lingual vein passes through the substance of 
the submaxillary gland, and where it emerges again to the sur- 
face receives the facial vein (text-fig. 1, fv.). At this spot lies a 
very definite lymph-gland, which in the Giraffe lies embedded in 
the substance of the salivary gland, and by its dark colour forms 
a very striking object when the surface of the gland is exposed. 


* Owen, Trans. Zool. Soc. vol. ii. 1841, p. 223. 


196 MR. R. H. BURNE ON THE 


In the Okapi, Antelopes, and Sheep this lymph-gland is not 
so deeply embedded as in the Giraffe, and in the Okapi (text- 
fig. 1, 2.g.) consists of two globular masses separated from the 
submaxillary gland by a capsule of fibrous tissue. 


Text-figure 11. 


SLAYV 66L.M 


» CLM G.HAY.M 


Blood-vessels at the root of the tongue of a Giratte. 


g.gi.m., genioglossus muscle; g.hy.m., geniohyoid muscle; h.gl.m., hyoglossus 
muscle; hy.a., hyoidean branch of lingual artery ; /.a., lingual artery ; La, 
lingual vein; s/.a. & v., sublingual artery and vein; st.gl.m., styloglossus 
muscle; v.a., venous anastomosis; v.pl., venous plexus. 

XII. Hypoglossal nerve. 


The lingual arteries of the Okapi and Giraffe agree in many 
particulars and show several, though small, differences from those 
of Antelopes. 

In both, the lingual artery arises from the carotid independently 
of the facial (text-figs. 1, 3, f.a.), as it is stated to do normally 


VISCERA OF AN OKAPI. 197 


in the Sheep* and Goat, and occasionally in the Ox. In 
Antelopes, on the other hand, a common trunk (the arteria 
maxillaris externa) intervenes between the carotid and the lingual 
and facial arteries, dividing to form these two arteries about the 
middle of the submaxillary gland. 


Text-figure 12. 


G.HY.M 


MM 


Blood-vessels at the root of the tongue of Okapia johnstoni. 
m.m., mylohyoid muscle turned back. 


Other lettering as in text-fig. 11. 


In passing the hyoid region of the tongue, the main trunk of 
the lingual artery may lie either deep or superficial to the hyo- 
glossus muscle. In the Antelopes (text-fig. 13, /.a.) and Sheep 
it lies deep to this muscle, and in this position forms in ‘the 
Antelopes a strong anastomosis (text-fig. 13, aa) with its fellow 
of the opposite side, from which are given off the continuation of 
the main lingual artery, the sublingual artery, and branches to 
the muscles of the hyoid region. 


* In the Sheep dissected it vas given off from an external maxillary trunk as in 
the Antelopes. 


198 MR. R. H. BURNE ON THE 


In the Okapi (text-fig. 12) and on one side in the Giraffe (text- 
fig. 11), the lingual arteries, after giving off a large branch to the 
hyoid region, passed superficially to the hyoglossus muscle, and 
in neither case was there any anastomosis between the vessels in 
this region. Nor was I able to trace in either of these genera 
the large connection between the lingual arteries at the com- 
mencement of the free portion of the tongue described by Owen 
in the Giraffe. 

On the other hand, I can corroborate the fact mentioned by 
Owen that in the Giraffe the left lhngual artery greatly exceeds 
the right in size (text-fig. 11), and carries practically the whole 
blood-supply to the free part of the tongue. 


Text-figure Lay 


Blood-vessels at the root of the tongue of an Indian Antelope. 
a.a., anastomosis of the lingual arteries. 


Other lettering as in text-fig. 11. 


This is not, however, the case in the Olapi or among Antelopes 
and Sheep. In them the two lingual arteries are of equal size, 
but it should be noted that in the Okapi the left artery divides at 
the commencement of the free part of the tongue into a large 
ventral and a small dorsal branch, and thus provides the chief 
source of supply to the ventral parts of the extremity of the 
tongue, as it does in the Giraffe, the dorsal parts being supplied 
by the right artery and the smaller branch of the left. 


The Laryn« (text-figs. 14, 15, 16). 

The larynx of the Okapi presents a remarkably close resem- 
blance to that of the Giratte, and by its structure suggests that 
the animal, like the Giraffe, 1s practically mute. In general it 


VISCERA OF AN CKAPI, 199 


Text-figure 14. 


CAP. 


FS Reoeeers 


; io CRAM | THAN HY.EM, 
ERT 14 


Mascles of the larynx of Okapia johnstoni. 


a.m., arytenoideus muscle; c.a.dm., crico-arytanoideus lateralis muscle; c.a.p.m., 
crico-arytenoideus posticus muscle; e.ph.m., crico-pharyngeal muscle ; 
c.th.m., crico-thyroideus muscle ; e¢.g/., epiglottis; hy.e.m., hyo-epiglottidean 
muscle; th.a.m., thyro-arytenoideus muscles; s.c., cartilage of Santorini. 


Text-figure 15. 


Longitudinal section of the pharynx and iarynx of Okapia johnstoni. 


a.c., arytenoid cartilage; a.p.ph., arcus palato-pharyngeus (posterior pillar of fauces) ; 
er., cricoid cartilage; ws., wsophagus; p., soft palate; ¢f., tonsillar fossa ; 
th.c., thyroid cartilage; v.c., vocal cord. 


Other lettering as in text-fig. 14. 


200 MR. R. H. BURNE ON THE 


has the simple form common to Ruminants. The eminence sup- 
ported by the arytenoid and Santorini’s cartilages is peculiarly 
large and prominent (text-fig. 14, s.c.). The general cavity (text- 
fig. 15) is quite simple, without recesses or pouches of any kind, 
and, as in the Giraffe, the vocal cords (text-fig. 15, v.c.) are 


Text-figure 16, 


Longitudinal section of the larynx of Okapia johnston, showing the cartilages. 


Lettering as in text-figs. 14 & 15. 


extremely rudimentary, represented only by minute folds con- 
necting the ventral extremities of the enormous arytenoids to the 
mid-line of the thyroid cartilage. The cartilages (text-fig. 16) 
closely resemble those of the Giraffe. The muscles (text-fig. 14) 
are similar to those of other Ruminants, and call for no special 
comment. 


VISCERA OF AN OKAPI. 201 


The Tonsils (text-figs. 1, 4, 6). 


The tonsillar fossa of the Okapi lies in the usual position (text- 
fig.15, ¢.f.), between the anterior and posterior pillars of the fauces 
somewhat in advance of the hinder free margin of the palate. It 
is a simple oval pit, with its fundus subdivided into two or more 
subsidiary recesses. The fossa and the area surrounding it are 
covered externally by a nodulated mass of tonsillar tissue, which 
is encapsulated by a thin longitudinal sheet of muscle derived 
from the muscles of the palate. 

The tonsils are thus of quite a usual Ruminant type, and very 
similar to those of the Giraffe. It is interesting, however, to 
note that in many Ruminants the single fossa is replaced by a 
row of separate incisions, which no doubt represent the subsidiary 
recesses in the fundus of the common fossa of the Okapi. 


The Palate. 


In 1915 the Royal College of Surgeons purchased the roughly 
prepared skeleton of a young male Okapi collected by Dr. Christy, 
in which the soft tissues covering the hard palate had not been 
removed. They are shown in the accompanying photograph 
(text-fig. 17). 

In front of the molar region the palate is extremely narrow, 
and throughout this region is traversed by a series of about 15 
major palatal ridges. The anterior three or four are weak and 
do not extend across the entire breadth of the palate; they 
are succeeded by some twelve or so more complete ridges bowed 
forward in the centre and with their free papillate edges directed 
backwards. In the middle line these ridges are interrupted, the 
right half of each being situated in the majority of cases slightly 
posterior to the left. Behind the level of the anterior molar 
teeth, the palatal ridges become less and less pronounced and 
more nearly transverse in direction and gradually fade away, 
leaving the greater part of the intermolar area of the palate 
quite smooth. 

Between the major palatal ridges are series of papille arranged 
transversely, forming minor ridges. 

As in the Giraffe, there is no papilla incisiva, but at the 
extreme anterior end of the palate, in front of the palatal ridges, 
are two little longitudinal slits—the openings of the naso-palatine 
canals. 

The soft palate terminates posteriorly in a simple curved 
border, without any indication of a uvula. Its lateral parts are 
prolonged, as the posterior pillars of the fauces, and become con- 
fluent upon the dorsal wall of the pharynx, forming, as in the 
majority of quadrupeds, a complete circular lip (arcus palato- 


Proc. Zoou. Soc.—1917, No. XTV. 14 


202 MR. R. H. BURNE ON THE ~ 


pharyngeus) (text-fig. 15, a.p.ph.) through which the nasal passage 
communicates with the pharynx, 


Text-figure 17. 


J 


Palate of Okapia johnstoni. 
The Thyroid Body. 

The thyroid body, as in many other Ruminants (Sheep, Ante- 
lopes, Giraffe), consists of the lateral lobes only, without a 
connecting portion or isthmus between them. 

Each lobe is an oval body measuring 47 mm. in length x 
16 mm. in breadth, and occupies much the same position as in 
Domestic Ruminants in the angle between the cesophagus the 
hinder part of the larynx and the trachea. 


The Lungs (text-fig. 18). 


The chief subdivision of the lungs follows the plan common to 
Ruminants, there being two lobes to the left lung and three and 
an azygos lobe to the right. The upper lobe of the right lung 
is partially divided into two minor lobes, the upper of which 
(text-fig. 18, 7.a./.") is the larger of the two, and is bent in the 


VISCERA OF AN OKAPI. 203 


characteristic Ruminant way transversely across the front of the 
heart. The lower (text-fig. 18, 7.a./.*) is leaf-like and resembles 
the same lobule in the Sheep or Goat, but is relatively larger. The 
middle and lower lobes and the azygos lobe resemble very closely 
those of the Goat. 


§ 


Text-figure 18. 


RAL 


B.R.AJ 


RPL 


Rib 


The lungs of Okapia johustoni, ventral aspect. 


b.l.a.l., bronchus of left apical lobe; 6.r.a.l., bronchus of right apical lobe; 
l.a., azygos lobe; J.a.l.1, left apical lobe, longitudinal process; J.a.l.?, 
left apical lobe, transverse or cardiac process; J.i.1., left inferior lobe; 
Li.l.a., cardiac appendage of left inferior lobe; 7.a.0.!, right apical lobe, 
transverse process ; r.a.l,*, right apical lobe, cardiae process; 7.7.2., right 
inferior lobe; 7.m.U., right median or cardiac lobe; ¢7., trachea. 


The upper lobe of the left lung is very similar to that of the 
‘Goat or Sheep, except for the greater length of its anterior 
14* 


204 MR. R. H. BURNE ON THE 
° 


(text-fig. 18, /.a.0.‘) (longitudinal) lobule. The lower lobe (text- 
fig. 18, 2.2.2.) is remarkably prolonged at its upper and outer angle, 
forming a long finger-like cardiac process (text-fig. 18, J.i.l.a.) that 
lies alongside the lower (transverse) lobule of the upper lobe 
(text-fig. 18, /.a./.*), bound to it by a fold of pleural membrane. 
In the natural position these two elongated lobules embrace the” 
left side of the heart. 

The trachea, chief bronchial tubes, and pulmonary veskele call 
for no particular notice, as they resemble closely those of the 
Sheep. 


The Heart. 


The heart has a long narrow form, and measures 140 mm. in 
total length from base to apex, and 95 mm. in breadth at the 
upper part of the ventricles. The length of the ventricular 
portion is 120 mm. 

It is of the same type as that of other Ruminants. In 
dissecting it, the musculi papillares of both atrio-ventricular 
valves gave one the impression of being more than ordinarily 
developed, particularly as regards the muscular swelling that 
forms the foundation of those of the anterior cusp of the mitral 
valve. The wall of the left ventricle seemed also to be thick for 
a heart of the size. 

The moderator band in the right ventricle is noticeably small. 
The great arteries spring from the arch of the aorta by a single 
large innominate trunk, as in other narrow-chested Ungulates. 
Between the aorta and pulmonary artery is a large vestige of the 
ductus arteriosus. 

There is at the root of the aorta in Ruminants a very cha- 
racteristic splinter of bone (os cordis) supporting the attach- 
ment of one of the cusps of the aortic valve. In the Okapi 
this bone is represented by a plate of cartilage, but whether its 
unossified condition is a permanent character or is due to youth 
it is difficult to say definitely, but it should be noted that in the 
calf ossification * sets in almost immediately after birth, and that 
in Antilocaprat the os cordis is permanently cartilaginous. 


The Thymus Body. 


The thymus is a relatively small triangular mass of tissue 
attached by a fold of membrane to the antero-ventral surface of 
the pericardium. It measures 64 mm. in length x 55 mm. in 
breadth at its posterior end. 


The Stomach < (text-figs. 19, 20, 21, 22, 23). 


In its general forin (text-fig. 19) the stomach of the Okapi 
resembles that of the Giraffe and many Antelopes more nearly 


* Vaerst, ‘“ Herzknochen bei Wiederkauern,” Deutsch. Zeitschr. f. Thiermedicin, 
Bd. 30, 1888, p. 53. 

+ Maurie, Proc. Zool. Soc. 1870, p. 348. 

t Garrod, Proc. Zool. Soc. 1877, p. 5. 


VISCERA OF AN OKAPI. 205 


than that of the various other Ruminants figured by Cordier *. 
All four chambers (rumen, reticulum, psalterlum, abomasum) 
are fully developed. 

The paunch (rumen) is subdivided by folds into secondary 
chambers in a manner similar to that of Domestic Ruminants, 
and is lined throughout by coarse papiile (text-fig. 20), mostly of 
large sizeand flattened lanceolate form, like those in the rumen of 
the Giraffe. Upon the free edges of the dividing partitions the 
papilla are reduced in height and resemble flat scale-like plates 
rather than papille. 


Text-figure 19. 


SPL oe eT 


gy 
é : ) 4 es ; pet ene 
ISTE e 
ae pe ee LER AS os 
cB Seas ar ag, 5 o, 


SSS eT ep se 


Re aecaek 


Outline of the stomach of Okapia, with the cavities opened. 


ab., abomasum; duo., duodenal dilatation; cs., cesophagus; cs.g., cesophageal 
gutter; psal., psalterium; pyl., pylorus; 7., longitudinal ruge in aboma- 
sum ; ret., reticulum seen through opening from the rumen ; ret.’, outline of 
the reticulum ; spl., spleen. 


The reticulum is a roomy, oval chamber, and is remarkable, like 
that of the Giraffe and Antilocapra ty, for the extreme shallowness 
of its cells (text-fig. 21). In this particular it is approached by 
the reticulum of many Deer, and offers a striking contrast to that 
of Domestic Ruminants and the majority of Antelopes. 

The psalterium is large, of about the same size as the reti- 
culum, and contains some 14 chief lamine. The spaces between 


* Cordier, Ann. des Sci. Nat. t. xv. 1898, pl. 6. 
+ Murie, Proc. Zool. Soc. 1870, p. 349. 


206 MR. R. H. BURNE ON THE 


the chief lamine are occupied by others of less height, consti- 
tuting according to their size and position secondary, tertiary, 
and quaternary series. 

The position and height of the different series are shown to scale 
in the accompanying diagram (text-fig. 23). It is to be noticed 
that of the quaternary series those only interposed between the 
secondary and tertiary series are represented by definite lamine. 
Those between the tertiary and primary, when they are repre- 
sented at all (and they apparently are not in every case), are 
indicated only by a row of minute papille (text- fig. 23, p.), as 
the whole ee ee series is in the Giraffe and certain other 
Ruminants* 


Text-figure 20. Text-figure 21. 


Text-fig. 20.—Papille of rumen, x 2. 


Text-fig. 21.—Cell of reticulum, nat. size. 


Text-figure 22. 


Specialised papille of cesophageal gutter, nat. size. 


1., lamina of psalterum ; cs.g., roof of esophageal gutter. 


Towards the esophagus the lamine gradually fade away, and 
are succeeded by rows of enlarged papillee similar in character but 
not in size to those that cover the surface of all the lamine. Some 
of these papille are enormously developed (text-fig. 22) and form, 
when the cesophageal gutter 1s closed, a coarse filter to prevent 
unsuitable food from passing into the psalterium. 

Papille similar to these, though fewer in number and smaller 
in size, are present in the stomach of the adult Ox, at the entry 
of the cesophageal gutter into the psalterium. 


* Garrod, Proc. Zool. Soc. 1877, pe 5. 


VISCERA OF AN OKAPI. 207 


In the Okapi each of these special papille of the cesophageal 
gutter consists, like those on the free margin of the leaves of the 
psalterium, of a fleshy base and a horny apex. The separation 
of the base from the apex is quite abrupt and clear, giving to the 
papilla the appearance of a long horny rod set upon the summit 
of a rounded fleshy boss. 

The digestive stomach (abomasum) calls for no particular com- 
ment. Its walls become gradually more and more muscular 
towards the pylorus, béyond which the commencement of the 
duodenum is marked, as in other Ruminants, by a sudden 
dilatation. The lining membrane, towards the cardiac end, is 
longitudinally folded. 


Text: figure 23. 


uu 


Diagram, to scale, of laminze of psalterium. 


1, 2, 3, 4, laminee of primary, secondary, tertiary, and quaternary series. 


p., lamina of quaternary series represented by row of papille. 


In the small portion of cesophagus preserved there was found 
amass of partly masticated food. This was composed of small 
fragments of the leaves of trees. Though the fragments were too 
small to offer much likelihood of detailed identification, Mr. L. A. 
Boodlé, of the Jodrell Laboratory, Kew Gardens, most kindly 
undertook to examine them. He reports that they “ consist, 
chiefly at any rate, of fragments of Dicotyledonous plants, grasses 
being apparently absent. A few specimens show considerable 
resemblanee to leaflets of certain Leguminose, but these and the 
fragments of other plants present have not been identified. No 
definite opinion has been arrived at as to the stature of ulate plants 
concerned.” 

This report, in the apparent absence of grass, agrees =n the 
field observations made by Dr. Christy. He says that the animal 
‘‘feeds chiefly on the small leaves and twigs of trees.” ‘It does 
not’ eat grass, but does browse on the coarse herbage on the 
outskirts of the forest” *. 


* Quoted from ‘The Field,’ July 10, 1915, by Sir H. H. Johnston, ‘ Nature,’ 
Aug. 16,1915, p. 714; see also to the same effect, Wilmet, C.R. Acad. Sci. Paris, 
1913, vol. 156, p. 2007. 


208 ON THE VISCERA OF AN OKAPI. 


The Spleen. 


The spleen (text-fig. 19, spd.) is massive and wedge-shaped, and 
is situated, as in other Ruminants, upon the left anterior surface of 
the paunch, adherent by its thick end to the diaphragm and with 
‘its narrow edge projecting freely to the left into the abdominal 
cavity. 


The Colic Coil. 


The colic coil is flat, not conical as in the Giraffe, and is almost 
circular, measuring 190 mm. in diameter. The gut of which it 
is constituted is relatively narrow (20 mm. in diameter), and is 
arranged in 33 concentric and 33 excentric coils. This number 
appears to be high. In Antelopes of which I have seen records * 
it apparently varies from 1 to 3, in the Ox it is 14, in Antdlo- 
capra 37, and in the Giraffe may be as many as 4. 


In judging of the bearing of the above anatomical facts, it must 
be borne in mind that they are records of one individual only, 
but assuming that they fairly represent the condition normal for 
the species, the following points seem to be most worthy of 
note :— 


A. In comparison with the Giraffe : 


1. The form and superficial anatomy of the tongue. 
2. The condition of the geniohyoid and _ genioglossus 
muscles. 

3. The occipito-styloideus muscle and the form of the 
styloid bone. 

The structure and form of the larynx. 

. The arrangement of the lingual arteries. 

The general form of the stomach, and particularly the 
character of the reticulum. 


OO 


B. As more or less characteristic of the Okapi: 


1. The papille of the sublingual fringe. 

2. The cartilaginous os cordis. 

3. The elongation of the upper angle of the left lower lobe 
of the lungs. 

4, The specialised papillee in the cesophageal groove of the 
stomach, 


* Beddard, Proc. Zool. Soc. 1909, i. p. 181; Mitchell, Trans. Zool. Soc. xvii. 1905, 
p. 473. 

+ Beddard, JZ. e. 

t Owen, Trans. Zool. Soc. vol. ii. 1841, p. 227. 


ON THE ‘* DEATH-WATCH” BEETLE. 209 


EXHIBITIONS AND NOTICES. 
March 20th, 1917. 


Dr. A. SmirH Woopwarb, F.R.S., Vice-President, 
in the Chair. 


The Srecrerary read the following report on the Additions 
made to the Society’s Menagerie during the month of February 
1917 :— : 

The registered additions to the Society’s Menagerie during the 
month of February were 101 in number. Of these 36 were 
acquired by presentation, 44 were received on deposit, 19 were 
purchased, and 2 were born in the Gardens. 

The number of departures during the same period, by death 

. and removals, was 115. 

Amongst the additions special attention may be directed 
Oh 

1 Allamand’s Grison (Grison allamandi) from Le Quiacea, 
Bolivia, presented by A. 8S. Hume on February 10th. 

1 European Bison (Bison bonasus) 3, born in the Menagerie 
on February 22nd. 

2 Golden Eagles (Aquila chrysaétus) from Ross-shire, presented 
by Lady Bignold of Rosque on February 16th. 


Mr. R. I. Pocock, F.R.S., F.Z.S., Curator of Mammals, ex- 
hibited a young Lion Cub recently deposited in the Gardens, and 
also a Galago from German East Africa presented by Commander 


R. J. N. Watson. 


April 3rd, 1917. 


Prof. E, W. MacBripg, M.A., D.Sc., F.R.S., Vice-President, 
in the Chair. 


Mr. C. J. GAHAN exhibited a living example of the “ Death- 
Watch” Beetle (Anobiwm tesselatwm), and demonstrated its 
response to sharp taps, such as could be made with the point 
of a pencil on the table. 


210 MR. ALFRED EZRA ON BIG-GAME SHOOTING IN INDIA. 


Mr. C. Davies SHERBORN, F.Z.S., exhibited an autograph of 
Captain Bligh of ‘The Bounty.’ 


Mr. C. Tare Reean, M.A., F.Z.8S., exhibited a Piraya (Serra- 
salmo piraya) from the Amazon, and gave a short account of 
the habits of this ferocious fish. 


Big-Game shooting in Cooch Behar, Assam, and the Bhutan 
Duars, India. 


Mr. Aurrep Ezra, F.Z.S., gave a lantern exhibition illustrating 
Big-Game Shooting in Cooch Behar, Assam, and the Bhutan 
Duars, India, and made the following remarks :— 

On account of the heavy grass- and reed-jungle, sometimes well 
over 20 feet high, so common in these districts, it is impossible 
to do any shooting on foot. All big-game shooting is done here 
with the aid of elephants, which are thoroughly trained for the 
purpose. During the shooting-season the country is generally 
burnt in patches, and here one follows the method adopted in 
covert-shooting in England. Two or more guns are sent in with 
the line of elephants, while some guns are posted forward as 
‘“‘stops,” and the patches are beaten up to them. The elephants 
used for the ‘ stops” are generally very steady and well-trained 
animals and are most valuable. The shooting arrangements at 
Cooch Behar are reduced to a science, and the men in charge of 
the line are so well trained that a tiger can invariably be beaten 
out to any particular “stop,” should the Maharaja wish a special 
guest to have the first shot. My favourite way of shooting in 
this country is to go alone on a pad-elephant with an intelligent 
mahout (driver) who is a good tracker. In this way you go 
through the thickest parts of the jungle without disturbing the 
game, and you see the animals in their natural haunts, sleeping, 
grazing, or w allowing in the muddy streams. Being accustomed 
to always seeing wild elephants, the animals do not take the 
slightest notice of your mount, and you are able to get within a 
few yards of them. At such close quarters the sportsman can 
pick and choose his head, only shooting something that is worth 
having as a trophy. The game found in this country consists of 
tiger, leopard, bear, rhino, buffalo, bison (gaur), sambur, swamp- 
deer, wild pig, and hog deer, 


ON THE PLUMAGE OF THE FEMALE JAPANESE PHEASANT, Date 


April 17th, 1917. 


Dr. A. SmirH Woopwarp, F.R.S8., Vice-President, 
in the Chair. 


The Srcrerary read the following report on the Additions 
made to the Society’s Menagerie during the month of March 
1917 :— 

The number of registered additions to the Society’s Menagerie 
during the month of March was 149. Of these 21 were acquired 
by presentation, 125 (chiefly small Finches) were received on 
deposit, 2 by purchase, and 1 was born in the Gardens. 

The number of departures during the same period, by death 
and removals, was 101. 

Amongst the additions special attention may be directed 
tO :-— 

1 White-handed Gibbon (Hylobates lav), from Pahang, deposited 
on March 29th. 

1 Bushy-tailed Galago (Galago crassicwudata), from Mafia I., 
Kast Africa, presented by Commander R. J. N. Watson, R.N., on 
March doth. 

1 White-toothed Shrew, from Albert, France, new to the 
Collection, deposited on March 23rd. 


\ 


Mr.C. Davies SuErgory, F.Z.S., exhibited the piece of hairy skin 
of a Gorilla, which specimen was torn off the freshly killed beast, 
and posted to Richard Owen, by P. B. Du Chaillu, from the interior 
of the Gaboon. ,The envelope is postmarked ‘+ Etablissements du 
Gabon, 1 Sept 1864,” and was the precursor of Du Chaillu’s long 
letter of August, 1864, from the Fernand Vaz river on the coast, 
announcing that six skins were being shipped to London among 
other specimens. These were registered at the British Museum 


1 Dec. 1864. 


My. D. Sers-Snrtu, F.Z.S., Curator of Birds, exhibited a female 
Japanese Pheasant (Phasianus versicolor) which had assumed 
the very nearly complete plumage of the male. The ovaries 
were found on dissection to be very atrophied. 

A male specimen of the Carolina Dueck (#w sponsa) was also 
shown in which the colour of the soft parts were similar to 
those of the female. When living, this bird was thought to be a 
female in male plumage, but on dissection proved to be a male 
with atrophied testes. 

Mr. Seth-Smith also showed some specimens of the trachea of 
various species of Ducks. 


ANI THE SECRETARY ON AN ABNORMAL CAPE LOURIE. 


My. EK. Heron-Auen, F.L.S., P.R.M.S., F.Z.S., described the 
Mussel-fishery and Foraminifera of Esnandes (La Rochelle), and 
the early work of Alcide d’Orbigny. . 

A series of slides was exhibited illustrative of the early studies 
of Alcide d’Orbigny at Esnandes (near La Rochelle), and the 
Mussel-fisheries established there since the year 1035. The 
experiments of Prof. W. A. Herdman on the West Coast of 
England were referred to, and those of Prof. A. Meek at Holy 
Island on the East Coast. A further series was shown illust- 
rating some of the notable d’Orbignyan species found in the 
neighbourhood, not recorded from there by d’Orbigny in 1826, 
but recorded from other localities at that date, and from distant 
seas between 1839 and 1846. A third series of slides illustrated 
well-known species from the locality which had been recorded and 
described by earlier authors, but were not apparently identified 
by d’Orbigny from the neighbourhood of La Rochelle. 


May 1st, 1917. 


Dr. A. SmrirH Woopwarp, F.R.S., Vice-President, 
in the Chair. 


The Srcrerary, Dr. P. Coatmers Mircue t, F.R.§., announced 
with the deepest regret that Mr. Henry Peavot, the Society’s 
Librarian and Clerk of Publications, had been killed in action. 
Mr. Peavot had entered the service of the Society in 1896, and, 
after passing through various departments, was appointed 
Assistant Librarian and Clerk of Publications in 1908, and was 
promoted to the post of Librarian and Clerk of Publications in 
1912. In every way he had gained the esteem and regard of 
the Scientific Fellows of the Society, and was one of the most 
valuable and competent members of the Society’s staff. 


The Srecrerary exhibited a coloured sketch of an abnormal 
Cape Lourie (Z'uracus corythaix Wagl.), kindly sent to him by 
Mr. E. C. Chubb, F.Z.S., the Curator of the Durban Museum, 
Natal. The specimen, which was the property of the Durban 
Museum, was normally coloured on the head and neck, but the 
rest of the plumage was pale grey or dirty white, with the 
exception of a portion of the wings, which retained the normal 
red colour. The example could not be exactly described as an 
albino, inasmuch as the red colour was known to be due to a 
pigment, the ‘‘ Turacin” originally described by Professor Church, 
and the green colour of the head and neck, unlike that in most 
species of birds with green feathers, was due also to a green 
pigment. 


THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 203 


A Gynandromorphic Specimen of the Karwig ( Forficula 
auricularia). - 


Prof. J. P. Hitu, F.R.S., F.Z.S., remarked as follows :— 

The specimen (a photograph of which was exhibited) attracted 
attention by reason of the asymmetry of its forceps. The right 
limb of the latter measured 3°25 mm. in length, and is unmistake- 
ably of the male type, being stout and distinctly curved. ‘The 
left limb measured 2°75 mim., and is just as unmistakeably of the 
female type, being slender and only slightly curved. 

Dissection of the specimen revealed the presence of a well- 
developed testes, containing fully formed sperms, on the right 
side. The left gonad was not observed and was probably either 
absent or very rudimentary. It is certain that a well-developed 
ovary was not present on this side. 


Dr. A. Smira Woopwarp, F.R.S., Vice-President, exhibited 
enlarged models, made by Mr. Frank O. Barlow, of the first 
lower molar tooth of a large Chimpanzee, Melanesian Man, and 
Piltdown Man (Loanthropus dawsoni). He maintained that the 
latter tooth was of the megadont human type. 


Mr. D. M. 8S. Warson, F.Z.8., exhibited and explained by 


diagrams, models of the teeth of fossil horses. 


Professor H. Maxwextui Lerroy, M.A., F.Z.S., described the 
Silkworms and Silk industry of India, illustrating his account 


with lantern-slides and specimens of the Silk Moths, Cocoons, 
and Silk. 


May 15th, 1917. 


Dr. 8S. F. Harmer, M.A., F.R.S., Vice-President, 
in the Chair. 


The Srcrerary read the following Report on the Additions 
made to the Society’s Menagerie during the month of April 
1917 :— 

The registered additions to the Society’s Menagerie during the 
month of April were 130 in number. Of these 46 were acquired 
by presentation, 33 were received on deposit, 48 by purchase, 
1 in exchange, and 2 were born in the Gardens. 


214 THE EVANESCENCE OF PATTERN IN THE SKINS OF ZEBRAS, 


The number of departures during the same period, by death 
and removals, was 93. 

Amongst the additions special attention may be directed 
to: 

1 Chimpanzee (Anthropopithecus troglodytes) 9, from West 
Africa, presented by Lieut.-Comm. J. W. Rainier, R.N., on 
April 16th. 

1 Demidoff’s Galago (Hemigalago demidoffi) 3, from Obuassi, 
Ashanti, presented by Dr. G. H. F. Spurrell on April 5th. 

2 Pandas (“lurus fulgens) $ 9, from Nepal, purchased on 
April 12th. 


Dr. P. CHatmers MircuHe.y, M.A., D.Sc., F.R.S., Secretary to 
the Society, showed the behaviour of a large series of Birds and 
Mammals in the presence of living Serpents, and demonstrated 
that, except in the case of the higher Monkeys and a few kinds 
of Birds, there was no trace of instinctive dread of Serpents. 


Mr. D. Seru-Smitu, F.Z.8., Curator of Birds, gave a lantern 
exhibition of Birds now. or recently living in the Society's 
Gardens. 


June 5th, 1917. 


Dr. 8S. F. Harmer, M.A., F.R.S., Vice-President, 
in the Chair. 


My. R. I. Pocock, F.R.S., Curator of Mammals, exhibited on 
behalf of Messrs. Rowland Ward Ltd. the skins of two Zebras, 
shot by Mr. David Mackenzie at Gil-Gil in British East Africa, 
which showed remarkable evanescence of pattern affecting the 
body, neck, and head and to a less extent the legs. The oblitera- 
tion of pattern was carried to a much greater extent than in the 
skin described by Prof. Ridgeway as representing a new variety 
named Hguus quagga goldfincha. 

The two specimens were secured from a herd of normally 
coloured Zebras of the East African race Equus quagga granti. 


ON EGG-PRODUCTION AT THE POULTRY EXHIBITION. 215 


Mr. R. E. Hotpine exhibited and made remarks on a Cirriped 
(Coronula diadema) which occurs in considerable numbers on the 
Humpbacked Whale (Megaptera nodosa Bonn. ). 

Mr. Holding also exhibited two curiously malformed eggs in 
which the inner membrane only remained, the shell and yolk 
being absent. 


Mr. D. Sera-Situ, F.Z.8., Curator of Birds, gave an account 
of the Poultry Exhibition which the Society is now holding for 
the second season with a view to the encouragement of poultry- 
keeping for egg-production. The method of keeping hens on the 
intensive system was explained, as well as the lines to be followed 
in the breeding and selection of heavy-laying strains. Mr. Seth- 
Smith called attention to the value of certain strains of ducks, 
especially those known as ‘** white runners,” for egg-production. 


eu * 


FY 


<@ 


, 


INDEX. 
1917.—Pages 1-215. 


[New names in clarendon type. 
(z.8.1.) indicates additions to the Society’s Menagerie. | 


Systematic references in italics, 


Abraxas grossulariata (Zz, 8. L.): 


logy, 96. 
Ailurus fulgens (4, 8. u.), 214. 


Aéschna grandis (Z. 8. L.) : ethology, 97. 


juncea (4. 8. L.): ethology, 
/Ex sponsa: exhibited, 211. 


etho- 


lord 


D1. 


Agrion puella (z. s. u.): ethology, 97. 


Anatomy. See STRUCTURE. 


Anobiuin tesselatum ; exhibited, 209. 


Anseranus semipaliuata: variat 


ion, 96. 


Anthropopithecus troglodytes (z. s. L.), 


214. ; 
Aquila chrysaétus (z. s. u.), 209. 
AVES: 


Archeopteryx: (fossil) structure, 1 ; 


Hybrid between male 


Thrush 


and female Blackbird, 95; Trachea 
of a male Anseranus semipalmata, 
96; Migration of Woodcock, 159; 
Plumage of Japanese Pheasant, 


211; Soft parts of male specimen | 
of Carolina Duck, 211; Trachea of’ | 


various Species of Ducks, 211; 


Sketch of an abnormal 


Cape 


Lourie in the Durban Museum,. 


212; Behavicur of Birds 
presence of living Serpent 


in the 


s, 214; 


Lantern-exhibition of birds from 
the Society’s Gardens, 214; Mal- 


formed eggs: structure, 


215 ; 


Poultry exhibition in the Society’s 


Gardens, 215. 


Proc. Zoou. Soc.—1917, No. XV. 


Bison bonasus‘(z. 8. u.), 209. 


. * rey + cA ‘y . 
c =) ° e . ° 
Brachytron pratense (z. 8. L.) 


logy, 97. 


Capricornis argyrochetes, 22, 
—  collasinus, 23. 


milne-cdwardst, 22. 


rocherianus, 23. 


vidianus, 22. 


Cervus dugennianus, 19. 


hanglu (z. 8. L.). 99. 
Cinnyris afer (z: Ss. L.), 96. 
Cis alni, 89. 

— bidentatus, 89. 

—- hilamellatus, 87. 
boleti, 88. 

castaneus, 91. 

—- dentatus, 89. 

—— elongatulus, 91. 
JSestivus, 91, 


—— hispidus, 89. 

—— jacquemarti, 87. 

-—— latifrons, sp. n., 89. 
lineatocribratus, 86. 


--— lineatosetosus, sp. n., 87. 


—— jicans, 88. 
nitidus, 87. 


oblongus, 90. 


—— punclulalus, 90. 
—— setiyer, 88. 
vestitus, 91, 


15 


etho- 


X1V 


CLASSIFICATION: 
Pre-Jurassic Tetrapods, 167. 
Cobus defassa (z. s. u.), 95. 
Coronuia diadema, exhibition, 215. 
CRUSTACEA: 
Palamonide : mouth-parts, 37 ; Coro- 
nula diadema (exhibited), 215. 
Cryptorrhynchus harrisoni, 
sp. n., 93. 


Donacia linearis (z. 8. L.): 
97. 

Duthiersia elegans, 79. 

expansa, 80. 

— finbriata, 79. 


ethology, | 


Ennearthron affine, 92. 
-— cornutum, 92. 
EKoanthropus dawsoni: exhibition, 213. 
Equus quagga granti: exhibition, 214. 
Eremias heterolepis, 154. 
Erinaceus albiventris (z. 8. 1.), 96, 
ErnoLoey. 

Mammauta: Behaviour in presence of | 

living Serpents, 214; Pattern on 


skins of Zebras from British East 
Africa, 214. 

AVES: Woodcock, | 
159; Behaviour of Birds in pre- | 
sence of living Serpents, 214. 

Insecta: British Cissid Beetles, 
83; species bred in the Society’s | 
Gardens, 96. 


Migration of 


Forficula auricularia : photograph, 213. 


Galago crassicaudata (z. s. L.), 211. 
GEOGRAPHICAL: 


MammautA: Heude’s collection of 


pigs &c. in Sikawei 
SHanghai, 7; Big Game-shooting 
in India, 210; Zebras from British 
East Africa (skins), 214. | 

Aves: Migration of Woodcock breed- | 
ing in the West of Ireland, 159. 

Piscus : Exhibition of a Piraya from | 
the Amazon, 210. 

Moutusea : Mussel-fishery &c. of Es- 
nandes, 212, 


Museum, 


INDEX. 


Grison allamandi (z, s. L.), 209. 


Hemigalago demidoffi (z. s. u.), 214. 
Hyla pulchella (z. s. 1.), 96. 
Hylobates lar (z. s. t.), 211. 


InsEcTa: 
British Cissid Beetles, 83; Insects 
bred in the Caird Insect House, 
96. 
Coleoptera : Anobium 
exhibition of living example, 209. 
Coleoptera: Cissidve found in Britain : 


tesselatum : 


systematic, 83. 
Dermayptera : Forficula auricularia, 
213. 
Lepidoptera: Silk Moths, Cocoons 
and Silk (exhibition), 213. 


Lacerta spinalis, 146. 

Latastia degeni, 154. 

hardeggeri, 150, 152, 154. 

—— neumanni, 150. 

-— phillipsit, 148. 

spinalis, 146, 

Libellula quadrimaculata (z. s. L.); 
ethology, 97. 


Mammalia. 

Chinese: systematic, 7; work of 
Beavers in the Society’s Gardens, 
100; Okapia johnstoni (viscera) : 
structure, 187; Pre-Jurassic Te- 
trapods: classification, 167; Lion 
Cub, 209; Galago from German 
East Africa, 209; Big Game- 
shooting in India, 210; Piece of 
hairy skin of a Gorilla, 211; Molar 
tooth of a Chimpanzee &c.: struc- 
ture, 213; Teeth of horses (fossil) : 
structure, 2138; Behaviour in the 
presence of living Serpents, 214 ; 
Pattern on sking of Zebras from 
British East Africa, 214. 

Merula merula: variation, 95. 
Moxuusca: Mussel-fishery of Esnaudes, 
exhibition of slides, 212. 
MorrPuo_oey. 
Nemorhedus arnouxianus, 2d. 


a 


See STRUCTURE, 


INDEX. | XV 


Nemorhedus caudatus, 26. 


cinereus, 25. 


griseus, 25. 


henryanis, 25. 

—— niger, 25. 

raddeanus, 26. 

Octotemnus glabriculus 93. 

Orthetrum ecerulescens (z. s. L.): etho- 


logy, 97. 


PaTHOLOGY. 
Animals in the Society’s Gardens, 
2: 
Phasianus versicolor, exhibited, 211. 
Philochorius hardeygeri, 154. 
intermedius, sp. n., 152. 
— neumanni, 150. 
— phillipsii, 148. 
spinalis, 146. 
PISCEs : 
Chimera colliei: L055 
Serrasalino piraya, exhibited, 210. 


structure, 


Reptiia: 
Lizards of the Genus Philochortus: 
systematic, 145. 
Rhopalodontus fronticornis, 92. 
—— perforatus, 92. 


Serrasalmo piraya: exhibited, 210. 
Solenophorus fimbriatus, 79. 
STRUCTURE, 

Mamnauia: Chinese, 7; Work of 
Beavers in the Society’s Gardens, 
100; Chimera colliei (skull), 105; 
Okapia johnstoni (viscera), 187; 
Mcdels of molar tooth of Chim- 
panzee &c., 213; Models of teeth 
of fossil horses, 213. 


Aves: Archxopteryx (fossil), 1; 
Ducks (trachea), 211; Malformed 
egos, 215, 

Insecta: Forficula auricularia (Gy- 
nandromorphic specimen), 213; 
Silkworms and Silk industry of 
India, 213. 

CrustacHa : Palemonidee (mouth- 
parts), 37. 

Vermipea: Duthiersia, 73. 

Sus coreanus, 15. 


gigas, 14. 

—- meles, 1d. 

—— moupinensis, 138. 
paludosus, 12. 


Tetropium gabrieli (z. s. L.): ethology, 
SH 

Trimerorhinus triteniatus (z. s. b.), 96. 

Turacus corythaix: sketch exhibited, 
22, 

Turdus musicus: variation, 9d. 


Ursus horribilis (z. s. u.), 95. 


VARIATION, 

Mammaria: Pattern on skins of 
Zebras from British East Africa, 
214, 

Aves: Hybrid between Thrush and 
Blackbird, 95 ; Trachea from An- 
seranus semipalmata, 96; Plumage 

hile 
Colour of soft parts of male Caro- 
lina Duck, 211; Cape Lourie in 
the Durban Museum, 212. 

VERMIDEA: 

Duthiersia : 
73. 


of Phasianus _ versicolor, 


structure, systematic, 


PRINTED BY TAYLOR AND FRANCIS, 


RED LION COURT, FLEET STREKT, 


PROCEEDINGS 


OF THE 
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS 


OF THE 


ZOOLOGICAL SOCIETY 


OF 1-0 NON, 


1917, pp. 217-338, 


witH 5D PLATES and 33 TEX't-FIGURES. 


PRINTED FOR THE SOCIETY, 
SOLD AT ITS HOUSE IN REGENT’S PARK. 
LONDON: 
MESSRS. LONGMANS, GREEN, AND CO,, 
PATERNOSTER ROW. 


kes 


OF THE 


COUNCIL AND OFFICERS 


OF THE 


ZOOLOGICAL 


SOCIETY OF LONDON. 


WM ee 


Patron. 
. His Masesry THe Kina. 


COUNCIL. 


His Grace THe Duxe or Beprorp, K.G., F.R.S., President. 


THe Hon. Crecit Barina, M.A. 
R. H. Burne, Esq., M.A. 
ALFRED H. Cocks, Hisq., M.A. 


Li.-Cout. S. Moncxron Copes- 
MAN, M.D., F.R.S. 

CHARLES Drummonb, KEsq., 
Treasurer. 

ALFRED Ezra, Esq., Vice- 
President. 

Cart. Huau S. GLADSTONE, 
M.A. 


Tue Viscount Harcourt, P.C. 


SIDNEY FrepERIC HArRMER, KsqQ., | 
Vice- | 


MrAy Seb oR. 
President. 

Proressor J. P..H1t1, D.Sc., 
EBS: 

WiLiiAM Huntsman, Esa. 


Sik HoMunp Gites Lopmr, Br., 
Vice-President. ; 

Con. Sir A. Henry McManon, 
G.C.M.G., C.8.1. 

E.:G. B. MerApE-WaAtzpo, 
Esq., Vice-President. 

P. CuatmMers MircHey, Esq., 
M.A., D.Se., Li, a 
Secretary. 

THe Marquis oF Suiao, F.S.A. 


_ OLpFrELD Tuomas, Esq., F.R.S. 


AUBYN Trevor-Batrys, Esq., 
M.A. 

ARTHUR SMITH WoopwarD, 
Ksq., LL.D. F.R.S., Vice- 
President. 

Henry Woopward, Esq., LL.D., 
F.R.S., Vice-President. 


PRINCIPAL OFFICERS. 
P. Caatmers Mircurtt, M.A., D.Sc., LL.D., F.R.S., 


Secretary. 


R. I. Pocock, F.RS., F.L8., Curator of Mammals and 
Resident Superintendent of the Gardens. ~ 

D, Seru-Smiru, Curator of Lirds and Inspector of Works. 

Lieut. Epwarp G. BouLencer, Curator of Reptiles. 

Prof. H. Maxwetu Lerroy, Curator of Insects. 


JOHN Barrow, Accountant. 
W. 4H. Corn, Chief Clerk. 


g 


LIST OF CONTENTS. 


1917, pp. 217-338.* 


EXHIBITIONS AND NOTICES. 


The Secrerary. Report on the Additions to the Society’s 
Menagerie during the months of May, June, July, 
August, and September, 1917....... SA Une err e e cir 


Prof. F. Woop Jonss, M.B., D.Se., F.Z.S. Exhibition 
of the skull of a Lioness, showing the effect of 
Seay HIS CN RR Nn Pers a Rana a esau la oa ihasue Tad edousledinnss 


Miss E. CHEEsMAN. Exhibition of, and remarks upon, the 
feeding habits of the Long Water -Scorpion, on 
the burrows made by the Mole-Cricket, and on the 
formation of the case by the Caddis-worm Phryganea 
grandiosa, based on observations made in the Caird 
Imeect Itouse:. (Vexb-tigure: Lo iiewwesssnecceden sen sneene 


Mr. C. Tare Reean, M.A., F.R.S., F.Z.S. Description of 
the present state of knowledge regarding the life- 
history of the Common Eel, illustrated by means of 
PMMA D Ge oh exes iae icin tonics takeie Mieaels teeta 345 


Mr. Rk. H. Burne, M.A., F.Z.8. Exhibition of a Common 
Frog showing the Spalteholz method of preparation . 


Prof. F. Woop Jones, M.B., D.Sc., F.Z.S. Exhibition of 


a series of brains prepared in the Prosectorium ...... 


Dr. P. Coatmers Mircue tt, F.R.S., F.Z.8., Secretary to the 
Society. Exhibition on behalf of Mr. D. Seru- 
SmirH, F.Z.8., Curator of Birds, of a set of lantern- 
slides prepared from photographs of mammals......... 


* (Parts III. & IV. (P. Z.S. 1917, pp. 217-388) were published 
together. | 


Page 


331 


333 


339 


336 


iv 


Page 
The Secrerary. Report on the Additions to the Society’s 
Menagerie during the month of October, 1917......... 336 
Mr. R,- I. Pocock, F.R.S., F.L:S., F.Z:8., Curatormen 
. Mammals. Exhibition of the head of a castrated 
Bushbuck, and of three successive pairs of antlers of 
an adult. Pére David's Deer 2!.......2.....0. eee Be 
Prof. J. P. Hitu, D.Se., F.R.S., F.Z.S. Lantern exhibition 
of new-born Marsupials, with an account of the mode 
of parturition and external characters .................. 330 
PAPERS. . 
12. Note on a Hermaphrodite Dogfish. By Rutan C. 
BamBer, M.Sc. (Plates 1. & ©.) 2.25 217 


13. On the Use of the Names VPlestosauria and Sauro- 


pterygia. By G. A. Boutencer, F.R.S., F.Z.S8. ...... 221 


14. Some Additions to the known Dragonfly Fauna of 
Borneo, with an Account of new Species of the Genus 
Celiccia. By F. F. Latpuaw, M.A., F.Z.8. (Text- 


foures T-2O.) 9125 a ccbeaas sat cel yo ane eee 223 
15, Notes on a Collection of Heterocera made by Mr. W. 
Feather in British East Africa, 1911-138. By Lt.- 
Col. J. Mo Fawenpie (Plate 1) er aes 9. ht eee 233 
16, Deformity of Os penis in a Seal (Phoca caspica Nilsson). 
By Srereius ALpHiRAKY. (Text-figure 1.) 23.0) 2. 251 
17. Some Observations upon Concealment by the apparent 
Disruption of Surface in a Plane at Right Angles 
to the Surface. By J. C. Morrram, M.B.(Lond.). 
(Text-feures Loa Wn; voici tess ee beexe eer eee ee 253 
18. New South-American Rhopalocera. By J. J. Joiczy, 
PLS. F.Z.S., EHS, and) “Gl SPALson, ibe: 
(Plates) 3.00 suki ete treet ie 259 


19, New South-American Arctiide.. By Joek J OICEY, . 
F.ILS., F.Z.8., F.E.S., and G. Tarsor, F.ES. 
(Platte Di ence. Re, ae nee ae 26 


and 


V 


Page 
20. -New Butterflies from Africa and the East. By J. J. 
Jorcny,. HES. h Zs: EES. and G. Tarsor, 
F.E.S. eovreereeer et POF FH HH OHHH HTC FOF SHH EHH HOHE HHH HOOT EHH HHO REO OD PG 
21. A Gynandromorph of Papilio lycophron Hbn. By 
J.J. Jorcey, F.LS., F.Z.8., F.E.S., and G. Tapor, 
He Bees 4( Plated) ...... BS Dit are cola ie citi den ipl 273 
22. Three Aberrations of Lepidoptera. By J. J. Jorcry, 
Has. F.Z.5., FES. and G. Tarsor, FES. 
(Blate wl, fig. 3.)- si..6s- Ree Sem R ee Eesisclee hers es oeaianrgatan cinta 275 
23. Ant-like Spiders from Malaya, collected by the 
Annandale - Robinson Expedition, 1901-2. By 
H. D. Bapcock, M.A. (Text-figures 1-12.) ......... 277 
24, The Structure of the Orbito-temporal Region of the 
Skull of Lemur. By F. Woop Jones, M.B., D.Sc., 
Boe (Rempneures. LD.) cccesssseisievetwessetes cx cans 323 
Alphabetical List of Contributors ................0cseccsneeseoeeee vil 


BUTS Ge ee ey sect he isn ag ives v'sluo ok WeE Sod we UW edad doxadedeoaseseeen’s xi 


= 


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iy 


wie A Db iE hCG A bal St 
OF THE 
CONTRIBUTORS, 


With References to the several Articles contributed by each. 


(1917, pp. 217-338.) 


Page 


ALPHERAKY, SERGIUS. 


Deformity of Os penis in a Seal (Phoca caspica Nilsson). 


(lextstioure’ 1.) 3.05... ER Ore er een Cee 251 


Bavcock, H. D., M.A. 


Ant-like Spiders from Malaya, collected by the 
Annandale-Robinson Expedition, 1901-2. (Text-figures 


L212 eon eee ne Set ae aCe rca tn a eee ena Pre 


BamsBer, Ruts C., M.Sc. 
' 


Note on a Hermaphrodite Dogfish. (PlatesI.&II.)... 217 


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


On the Use of the Names Plesiosauria and Sauro- 
TOSELED TG CRP seh at OBE A EEL eee ene ee ee eee 22 


Burne, R. H., M.A., F.Z.S. 


Exhibition of a Common Frog showing the Spalteholz 
method of preparation 


vill 
} Page 
CHEESMAN, Miss EK. 

Exhibition of, and remarks upon, the feeding habits 
of the Long Water-Scorpion, on the burrows made by 
the Mole-Cricket, and on the formation of the case 
by the Caddis-worm Phryganea grandiosa, based on 
observations made in the Caird Insect House. (Text- 


figure 1.) wi sassi canes causes esuinn e> delncleleesiese see see see eee 333 


Fawcett, Lt.-Col. J. M. 


Notes on a Collection of Heterocera made by Mr. W. 
Feather in British East Africa, 1911-13. (PlateT.) ... 233 


Hint, Prof. J. P., D.Sc., F.B.S., F.Z.S. 


Lantern exhibition of new-born Marsupials, with 


an account of the mode of parturition and external 


Characters. scot. cbecy sees cs de ole cesar eee eee 337 
Jorcey, J. J., F.LS., F.Z58., F.E.S., and Tatsot, G,, FBS: 
New South-American Rhopalocera, (Plate I.)......... 259 
New South-American Arctiide. (Plate I.) ............ 265 
New Butterflies from Africa and the East............... 271 
A Gynandromorph of Papilio lycophron Hbn. 
(Plates i c5o ist A ae te ue e eee APA Be te eee 273 


Three Aberrations of Lepidoptera. (Plate I. fig. 3.). 275 


Jonges, F. Woop, M.B., D.Sc., F.Z.S. 


The Structure of the Orbito-temporal Region of the 
Skull‘of Lemur. o(lext-tigures 1=5;) 0). e eee 323 


Exhibition of the skull of a Lioness, showing the 
efiect of captivity: is.2501...c:caaseaseoueean eh eee ae tee a eee 333. 


Exhibition of a series of brains prepared in the 
Prosectorium 


1X 


LAIDLAW, F. F., M.A., F.Z.S. 


Some Additions to the known Dragonfly Fauna of 
Borneo, with an Account of new Species of the Genus 
Colca. WWext-houres! 10.) cose. cect eres een uote eae 


MircHei, P. Cuatmers, M.A., D.Sc., LL.D., F.R.S., F.Z.8., 
Secretary to the Society. 

Report on the Additions to the Society’s Menagerie 

during the months of May, June, July, August, and 


EC DUCINIO CI MMO -guharns ia atomasn Setter nce soagteas ates 

Exhibition, on behalf of Mr. D.’Sern-Smira, F.Z.S., 
Curator of Birds, of a set of lantern-slides prepared 
frome PhoOtvopraphs OL MAM MGS i656 s.0c.sscateeccaeaneeassaees 


Report on the Additions to the Society’s Menagerie 


during the month of October, L917 .....0c..-...ecscssevess eee 


Morrram, J. C., M.B. (Lond.). 


Some Observations upon Concealment by the apparent 
Disruption of Surface in a Plane at Right Angles to the 
murmiaeer o(Text-feures 1-4.) (crise. cccceseecseossincrsosceess 


Pocock, RecinauD I., F.R.S., F.LS., F.Z.8., Curator of 
Mammals. 


Exhibition of the head of a castrated Bushbuck, and 
of three successive pairs of antlers of an adult Pere 


David’s Deer .......... Be er ee era SMM ei cinterg 4 Vit Aa tact eB tae 


Re@an, C. Tats, M.A., F.R.S., F.Z.S. 


Description of the present state of knowledge regarding 
the life-history of the Common Eel, illustrated by means 


WimlaMiberMeslIGOe es 7. .c cadet coud cics ub ad dabwsdedsce vaceereaeses 


Taupot, G. See Jorcey, J. J. 


Proc. Zoou. Soc,—1917, 6 


Page 


223 


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336 


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SCYLLIUM CANICULA. HRRMAPHRODITE CONDIMGR 


P.Z.8.1917.BAMBER PLII. 


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CANI CUuLA. HERMAPHRODITE CONDITION. 


SCYLLIUM 


PROCEEDINGS 


OF THE 


GENERAL MEETINGS FOR SCIENTIFIC BUSINESS 


OF THE 


ZOOLOGICAL SOCIETY OF LONDON. 


PAPERS. 


12. Note on a Hermaphrodite Dogfish. 
By Ruru ©. Bamser, M.Se.* 


[Received August 8, 1917: Read October 23, 1917.] 


(Plates I. & II.*) 


INDEX. Page 
AVG EVEN GUO) 01. Meg rcunayes cat ak! hg on Sn hee ED) Bo 


A case of hermaphroditism in Scylliwm canicula L. has recently 
come under my notice whilst demonstrating in the Zoology 
laboratories at the University of Liverpool, and, as the condition 
is rare in Selachians, it may be of interest to record the details. 

In 1914, Vayssiére and Quintaret ¢ described a hermaphrodite 
specimen of Scyllium stellare L. from the Gulf of Marseilles, but 
it was quite unlike the one I have seen. Their specimen had the 
left pelvic fin like that of a normal female, and the right like 
that of a male, with the clasper well developed. Internally an 
ovary and both oviducts were present. Also there was one 
testis, the right, united with the posterior end of the ovary, and 
the male ducts were normally developed on the right side only. 


* Communicated by C. TatE Rrean, F.R.S., F.Z.S. 

+ For explanation of the Plates see p. 219. 

f~ A. Vayssiére et G. Quintaret, “Sur un cas d’hermaphrodisme dun Seylliwm 
stellare L.,” C. R. Ac. Sci. Paris, 158. pp. 2013-2014, : 


Proc. Zoou. Soc.—1917, No. XVI. 16 


218 MISS RUTH C. BAMBER ON 


The urinary system was not connected in any way with the 
genital system, the right and left ureters joining together to 
form a short common canal which opened separately into the 
cloaca, on a papilla behind the female genital opening. 

There does not appear to be any other record of hermaphro- 
ditism in Seyllium; but Semper*, in 1875, recorded a rudimentary 
testis in a female Hexanchus. 

The specimen which is described below came from Port Erin, 
Isle of Man. It was unfortunately so badly damaged that the 
description 1s necessarily incomplete. 

Externally the animal was a typical male; the claspers were 
well developed, and the pelvic fins were united together along 
almost the whole length of their inner edges. 

Internally, both male and female organs were present. Both 
testes were developed, and except for the lobate character of the 
left testis, looked normal when seen from the ventral surface 
(Pl. I. fig. 1). On the dorsal surface, however, the right testis 
showed a small mass of ova developed at the anterior end, on 
the inner side. Dissection showed that the ovarian part formed 
about half the thickness of the right gonad, and was confined to 
the anterior end (Pl. II. fig. 4). The oviducts were normal, and 
each had an oviducal gland. The right duct and gland were con- 
siderably larger than the left. The ducts opened together into 
the coelom anteriorly as in a normal female. Posteriorly they 
were united together by their adjacent walls for about 2 cm., 
but their cavities remained distinct (Pl. I. fig. 2). The external 
opening was not seen. 

The male ducts were well developed. The vasa efferentia 
were not seen because of the damage to the mesentery, but on 
each side there was a well-defined epididymis, and a long, narrow, 
convoluted vas deferens, increasing in diameter posteriorly to form 
the straight vesicula seminalis. Sperm-sacs were not present 
(fig. 2). Posteriorly each vesicula seminalis united with the dorsal 
wall of the oviduct of its own side, and for a short distance 
after uniting, the right one ran parallel with the oviduct and 
then opened into it. The left one, however, ran obliquely in the 
wall of the left oviduct towards the median line of the body until 
it reached the wall separating the two oviducts, and then opened 
into the right one, side by side with the opening of the right 
vesicula seminalis (PI. II. fig. 3). No papilla was formed, but 
a little fold of the wall of the oviduct surrounded the two male 
genital openings, so that they lay in a slight depression. 

The interrelationship of the genital ducts, as also the complete 
development of the oviducts coupled with the absence of sperm- 
sacs, might suggest the possibility that the sperm-sacs are 
normally developed from the posterior ends of the degenerate 
oviducts. The specimen deseribed by Vayssiére and Quintaret, 


* Semper, “ Das Urogenitalsystem der Plagiostomen und seine Bedeutung fiir das 
der titrigen Wirbelthiere,” Arb. Inst..Witizburg, Bd. 2, 1875, pp. 195-509 (see 
p. 278 & Taf. xiv.). 


A HERMAPHRODITE DOGLISH. 219 


however, conclusively disproves any such homology, for it had 
both oviducts well developed and also the sperm-sac of the 
right side. 

The ureter of each side was attached to the inner side of the 
vesicula seminalis, slightly in front of the point where the latter 
became united with the wall of the oviduct; behind this point 
it was united also with the wall of the oviduct. The ureter 
opened into the vesicula seminalis of its own side immediately 
before the latter opened into the oviduct. The small duct from 
the posterior end of the kidney opened separately into the vesicula 
seminalis on a level with the opening of the ureter. 

It will be seen that, except for the absence of sperm-sacs, this 
animal had the complete genital systems of both male and female, 
and thus showed the completion in the adult of the primitive 
embryonic condition in which both male and female genital ducts 
are potentially present. 


EXPLANATION OF THE PLATES. 


Pruate I, 


Fig. 1. Gonads, from ventral surface: about natural size. 
2. Urinogenital ducts: semi-diagrammatic: about natural size. 


Puate [T, 


Fig. 3. Diagram of posterior part of urinogenital ducts. Ventral view. 

Figs.3a,36, & 3c. Diagrams of transverse sections of urinogenital ducts at a, 5, 
& c respectively. 

Fig. 4. Anterior end of right gonad, dissected to show hermaphrodite character of 
that region. Ventral view. The gonad has been split along the inner 
edge, and the ventral half turned back to show tl.e ovarian region: about 
1} natural size. : 


16 


. ON THE NAMES PLESIOSAURIA AND SAUROPTERYGIA. 220 


13. On the Use of the Names Plesiosauria and Sauropterygia. 
By G. A. Boutenesr, F.R.S., F.Z.S. 


[ Received October 6, 1917: Read November 6, 1917.] 


INDEX. 
SYSTEMATIC: Page 


On the names Plesiosauria and Sauropterygia ......... 221 


I wish to point out the misapplication, or rather inverted 
application, of the names Plesiosauria and Sauropterygia by some 
of the most recent writers on the classification of fossil Reptiles. 

The order Plesiosauria was so named, and properly defined, by 
de Blainville in 1835 *; the genus Plesiosaurus was then its only 
representative. 

When, in 1839, Owent accepted the name Hnaliosauria 
(Conybeare, 1821) for the marine Reptiles known as Plesiosaurs 
and Ichthyosaurs, which were associated in one order, he desig- 
nated the former as Plesiosawit and the latter as Jchthyosaurt. 
After the relationship of the Nothosaurs to the Plesiosaurs had 
been recognised by Hermann von Meyer, they were placed 
together as Plesiosauri by Quenstedt in 1852 f. 

Plesiosauria (1835-1852) is the earliest name for the order in 
question, it is open to no objection, and it should therefore be 
used, as it has been by Huxley, Gegenbaur, Cope, Baur, Hay, 
and myself, 

In 1859, Owen§, dropping the artificial group Enaliosauria, 
proposed to call Sawropterygia and Ichthyopterygia the two orders 
on which he had already bestowed names which there was no 
need to change. The Sauropterygia were defined as long-necked 
marine Reptiles with fin-like limbs with not more than fiv 
digits. Owen insisted on the character of the limbs as distinctive 
of the order and, although accepting the proposition that the 
Nothosaurs should be included, remarked, rather inconsistently :— 
‘““T continue, as in my former Report of 1841, to regard the fin- 
like modification of the limbs as a better ordinal character than 
the number of vertebre in any particular region of the spine 
, ee The Plesiosaurus, with its very numerous cervical 
vertebra, sometimes thirty 1n number, may be regarded as the 
type of the Sauropterygia or pentadactyle sea-lizards.” 

It is therefore perfectly clear, and beyond discussion, that the 


* Ann Mus. Paris (3) iv. p. 241.—Reference to this important contribution to 
the classification of Reptiles has unfortunately been omitted from O. P. Hay’s most 
useful bibliography, Bull. U.S. Geol. Surv. no. 179, 1902. 

+ Rep. Brit. Assoc. 1839, p. 45; also 1841, p. 60. 

{ Handbuch der Petret faktenkunde, pe 130: 

§ Rep. Brit Assoc, 1859, p. 159. 


222 ON THE NAMES PLESTIOSAURIA AND SAUROPTERYGIA, 


earlier name Plesiosauria, also the better from its non-committal 
meaning, should stand for the order in its wide sense; and that 
the later name Sauropterygia should only be applied, in accordance 
with Owen's definition and express designation of the type, as 
well as with its etymology, to the suborder including the 
Plesiosauride, Pliosauride, and Hlasmosauride, in which the 
limbs are transformed into hyperphalangeal fin-like paddles. 


ON BORNEAN DRAGONFLIES. Dire 


tel hl 


14. Some Additions to the known Dragonfly Fauna of 
Borneo, with an Account of new Species of the Genus 


Celiccia. By F. F. Latpiaw, M.A., F.Z8: 
[Received September 4, 1917: Read November 6, 1917.] 


(Text-figures 1-10.) 


INDEX. 
MorPHoLocy : Page 
A feature in the structure of the prothorax of Celiccia sp., 2 ... 229 
SYSTEMATIC: 
COMCCUG {LADOSLIIGTA, SP. Dy. saiciy22c55 ska vee ess ava edits wasann evs syn caesea, 2D 
of CONUPUOMU SIs Us. Vice nntncees ts vancetm sens vat cbs soap sater ares eM 
55 RUA CTOSELGIN Gs SP Wie 2 0864.5 x95 ee seas talesdwatennetdennnoveeccss S20 
4 TWUOCONCMALE ASD Ms: kia tusks Saks tie a SO 
ZYGOPTERA. 
Family AGRIONID& (=Ca@NAGRIONINE Kirby), 
Legion PLATYCNEMIS. 
CaELICCIA FLAVOSTRIATA, sp. n. (Text-figs. 1, 2.) ‘ 


25 $. Mt. Merinjak, 21-28.v.14, (The second specimen is 
the type.) 
1g. Mt. Matang, 4. xii. 13. 
Length of abdomen 35 mm., of hind wing 21°5 mm. 
M, rises at level of subnodus, Rs a little after *. Three cells 
between quadrilateral and subnodus. Costal margin of quadri- 


lateral three-fifths the length of anal margin in fore wing ; three- 
fourths in hind wing. 


Text-figure 1, 


Wings of Celiccia flavostriata Laidlaw. Type g. 


Pterostigma dark brown, sides approximately parallel, covering 
rather more than one cell. 12-13 postnodals on fore wing. 
(Text-fig. 1.) 


* The veins are indicated by the Comstock-Needham notation, 


294 MR. F. F. LAIDLAW ON 


Head: Lower lip yeliowish white, otherwise the head is black 
except the anteclypeus and extreme base of upper lip, which are 
white, and a pair of transverse marks, cuneiform in shape, on 
either side ef the hinder ocelli, these marks being yellowish 
white. 

Prothorax: Anterior and posterior lobes black, middle lobe 
yellow, with a very fine median black line. 

Thorax black above, yellow at the sides and underneath, a 
pair of orange-yellow antehumeral stripes, and at the top of 
either shoulder, just outside the stripes, a fine yellow spot. On 
the side of the thorax a black line on the second lateral suture, 
incomplete below. 

- Abdomen: Segment 1 yellow, with median line of black. 
Segments 2—5 brown, paler below, each with a black terminal 
ring. The remaining segments progressively darker, the last 
two bronze-black. 

‘The youngest of the three males has a whitish, diamond-shaped 
spot on the dorsum of segment 10. 


Text-figure 2 


Celiccia flavostriata, 3g. 


Anal appendages, seen from above. 


Anal appendages black, upper pair longer than segment 10, 
lower pair one-third: longer than upper pair. Upper pair mode- 
rately stout, incurved, flattened dorso-ventrally towards the free 
extremity; each carries an internal tooth at the end of its basal 
third. In the youngest specimen the flattened part of the 
appendage is yellow. The lower pair are slender, cylindr ical and 
incurved, (Text-fig. 2 

Legs: Femora sllon with black stripes ; tibie brown, tarsi 
and spines black. 

Type 3 in the'British Museum. Paratypes, one in my own 
collection, and one to be returned to the Sarawak Museum. 


C@LICCIA CAMPIONI, sp. n. (Text-figs. 3, 4.) 
1g: lio Matu;, 31. x54, 


Length of abdomen 37 mm., of hind wing 21 mm. 

Postnodals on fore wing 13. 

Closely allied to the preceding species (C. flavostriata). It 
differs as follows :— 

Colour: No cuneiform markings on vertex.. Markings on 


BORNEAN DRAGONFLIES. Pia, 


prothorax and thorax bluish white, possibly blue in life. The 
black lateral line on thorax relatively longer. 

Venation: The pterostigma is more oblique, its inner and outer 
margins more definitely parallel than is the case in C. flavostriata. 
(Text-fig. 3.) : 

Prothorax: The posterior lobe carries a very fine projecting 
point at either lateral angle, not present in C. flavostriata. 


Text-figure 3. 


Wings of Celiccia campioni Laidlaw. Type. 


Text-figure 4. 


See ee 
Coliccia campioni, g. 
Anal appendages, seen from above. 


Anal appendages (see text-fig. 4): Upper pair a trifle longer 
than lower, stout and a little incurved; hooked sharply down- 
wards at the extremity ; each carries at its middle an inwardly 
directed spur. Lower pair stouter and relatively much shorter 
than in C. flavostriata ; hooked in towards each other distally. 

Type ¢ in the British Museum. 

These two small species, characterized by their open reticula- 
tion and antehumeral stripe, would appear to form an independent 
section of the genus. ‘ 


CG@LICCIA MACROSTIGMA, sp.n. (Text-figs. 5, 6.) 
I-¢- Baram; Borneo, 20.x.10 (1 2, Baram, 19.x. 10). 


Length of abdomen, ¢ 35 mm., of hind wing 21 mm. 


” ” » (¢@ 33 mm., - 22) mm.) 
; ace Louie 14 | 14. 
Postnodals it) 15" ( e) ae) 


226 MR. F. F. LAIDLAW ON 


M, rises at subnodus, Rs a little distal. Three cells between 
quadrilateral and subnodus. Costal margin of quadrilateral 
three-fourths the length of anal margin in the fore wing, four- 
fifths in the hind wing. : 

Pterostigma brown, with fine pale margin, rather large, dis- 
tinctly broader than the cells of the postnodal costal area. In 
the front wing it lies over a single long and narrow cell, in the 
hind wing it covers one cell and a half. (Text-fig. 5.) 

Head: Upper lip and anteclypeus black, gene and lateral parts 
of postclypeus yellow, the latter black in the centre. rons 
and vertex black, base of antenne and a minute spot on either 
side of the posterior ocelli yellow. Occiput black, with yellow 
postocular mark. 


Text-figure 5. 


Wings of Celiccia macrostigma Laidlaw, . Type. 


Text figure 6. 


Caliccia macrostigma, 6. 


Anal appendages. 


Prothorax: Anterior and posterior lobes black; middle lobe, 
ventral and lateral surfaces whitish (probably blue in life). 

Thorax: Dovsum black, with broad (biue ?) antehumeral bands. 
These are indented at the middle by a black mark projecting 
from the inner side. Laterally the thorax is bluish white, . 
ventrally yellowish whlate ; a fine black line runs along the second 
lateral suture. 


BORNEAN DRAGONFLIES. PAE 


Legs yellowish white, the joints, spines, and a line on the 
femora black. 

Abdomen: Segment 1 yellowish white with dorsal brown spot ; 
segments 2-7 brown, becoming darker posteriorly, with narrow 
black apical rings, 2 has a small pair of basal lateral blue (?) 
spots; segment 8 is black above, yellowish white below, 9 and 10 
yellowish white. During life the light colouring on these seg- 
ments is probably blue; 9 and 10 have fine black basal rings. 

Anal appendages yellowish brown (? blue) tipped with black. 
Upper pair a trifle longer than lower pair, each with a large 
triangular projection inwards and downwards, scarcely visible in 
profile. Lower pair cylindrical, slightly incurved, and with a 
distinct elbow at the middle. (Text-fig. 6.) 

The presumed female of this interesting species has most un- 
fortunately been completely destroyed whilst in my possession. 
I have, however, in my possession an account of the specimen 
which I append here. 

Colouring in general very similar to that of the male. It 
differs as follows :— 


1, Antehumeral band of thorax completely divided by the 
black projecting mark into a broader lower part, which is 
rounded below and pointed above, and a narrower upper 
part, which is linear. 

2. Abdomen: Segment 1 entirely yellowish brown above; 8 
pale above (probably blue during life); 9 black, with a 
pair of pale marks above on either side of the middle line ; 


10 black. 


The structure of the prothorax is very remarkable. A curious 
crescentic projection stands out on either side of the middle lobe 
attached to the prothorax by its convex border. When looked 
at obliquely from above, each of these projections shows like the 
moon at the end of the first quarter, but when viewed directly 
from above each shows as a single outstanding spur. 

In addition the black posterior margin of the prothorax carries 
a fine black spur mid-dorsally. This is nearly -5 mm. in length, 
and projects forwards and upwards at an angle of about 45° to 
the long axis of the body. Oni either side of this projection lies 
a small lappel, also black, directed backwards, 

I do not know of any structure in the Legion Platycnemis 
that can be compared to the crescentic structure on the pro- 
thorax, except in the females noted below. 

This female was taken in the same locality as the male, and 
the two specimens were captured on two consecutive days. ‘They 
agree In size and wing-characters, and closely in colouring, though 
in this respect they are by no means identical. The evidence 
that they belong to the same species is not conclusive, though 
enough to justify the assumption that they may do so. 

Type 3 in the British Museum. 


228 MR, F. F. LAIDLAW ON 


CG@LICCIA NIGROHAMATA, sp. n. (Text-figs. 7, 8.) 


6 gd. Mt. Merinjak, Borneo, 21-28. v. 14. 


Length of abdomen 38 mm., of hind wing 24 mm. 

M, rises from subnodus, Rs a little distal. Three cells between 
quadrilateral and subnodus. Costal margin of quadrilateral 
about two-thirds the length of anal margin in fore wing ; four- 
fifths in hind wing. Postnodals 17-18. 

Pterostigma black, covering 14-2 cells. (Text-fig. 7.) 

Head: Upper surfaces black, except for a pair of yellow marks 
on the gene, and a pair of minute yellow spots on the vertex on 
either side of the hinder ocelli. 

Prothorax black, with lateral and ventral parts yellow. 


Text-figure 7. 


Wings of Celiccia nigrohamata Laidlaw, g. Type. 


Thorax: Dorsum bronze-black; on either side of the mid- 
dorsal carina are two silver-blue spots, the lower large and 
elliptical, the upper small and linear. The black colour extends 
to the level of the first lateral suture, beyond this as far as the 
second lateral suture the side of the thorax is of a dull purple- 
blue colour. Below this the thorax is yellowish white, delimited 
by a black line along the second lateral suture. 

Legs yellowish brown; the articulations and spines black, as 
are a line on the posterior surface of the femora and on the 
anterior surface of the tibie. 

Abdomen: Segments 1-7 black, 1 with yellow on the sides; 
segment 8 is black dorsally and blue below; segments 9 and 10 
are blue. 

Anal appendages black; the upper pair shorter than the lower 
pair, obliquely truncate, with two internal ventral teeth on each, 
Lower pair slender, incurved, pincer-like (see text-fig. 8), 

Type o in the British Museum. 


BORNEAN DRAGONFLIES. 229 


Cauiccia sp. (Text-fig. 9.) 
3 9 2. Kuching, Sarawak. 


These specimens belong to a species probably closely allied 
both to C. macrostigma and to C..nigrohamata. ‘They may even 
belong to the latter species, but as there is no definite evidence 
on the subject I note them here separately. 

Length of abdomen 22 mm., of hind wing 34 mm. 

M3; rises at or immediately before subnodus, Rs distal; costal 
margin of quadrilateral two-thirds length of anal margin in the 
fore wing, four-fifths in the hind wing. Three cells between 
quadrilateral and subnodus. 

Head black, upper hp with paired yellow spot at its base; 
frons with yellow line from eye to eye. Yellow marks between 
ocelli, antenne, and eyes, small linear postocular mark of the 
same colour. 

Prothorax: Anterior and posterior lobes black, middle lobe 
yellow, with median longitudinal black stripe. On either side of 


Text-figure 8. Text-figure 9. 


Text-fig. 8—Celiccia nigrohamata, $. Anal appendage. 
Text-fig. 9.—Ceeliccia sp., 2. Prothorax, seen from above. 


the lateral lobe is developed a small projecting spur, similar: to 
that found in the presumed female of C. macrostigma, but not so 
largely developed (see text-fig. 9). Posterior margin with fine 
median spine about -25 mm. long. 

Thorax black above, with large concentric antehumeral bands, 
concave inwards, blue in colour. Sides and under surfaces yellow ; 
a fine black band on the second lateral suture, incomplete below. 

Legs yellow; spines and articulations, and a band on the 
posterior surface of femora, black. 

Abdomen brownish black above; segment 1 yellow; segments 
9 and 10 black. Ventral side brownish yellow. Pale, small 
antero-lateral lunules on segments 3 and 4. 


Ca@LICCIA sp. 
1 $. Platang Rd. 


There remains to be noted this specimen, which is unfortunately 
: a 


230 MR. F. F. LAIDLAW ON 


incomplete, being without the head. It is evidently very closely 
allied to C. mgrohamata, but differs in the following respects. 
Segment 8 of the abdomen bas its terminal quarter ringed with 
blue, the blue mid-dorsally has its margin indented with black. 
The upper anal appendages are brownish black, more abruptly 
truncate than is the case in C. nigrohamata, and distinctly 
though very little longer than the lower pair. The lower pair 
are relatively stouter than in C. nigrohamata, and are brownish 
yellow (probably blue in life). 


The three species, C. macrostigma, nigrohamata, and that to 
which this male belongs will probably constitute another distinct 
section of the genus Cliccia, characterized by small size, rather 
dense reticulation of wings, as opposed to the open venation of 
C'. flavostriata; by the shape of the upper anal appendages of the 
male; and probably by the prothoracie structure of the female. 

To this group I believe it will be found that the species called 
Trichocnenis octogesima albicauda by Forster (which I suggest 
should. be named C. albicauda (Forst.)), also belongs. 


C@LICCIA MEMBRANIPES (Ramb.), race NEMORICOLA Laidlaw. 


CO. nemoricola Laidlaw, Journ. R. Asiat. Soc. Str. Br. no. 63, 
p. 95 (1912); id. Proc. Zool. Soc. London, 1915, p. 37. 

L have re-examined the examples of this form from Mt. Kina 
Balu, and have come to the conclusion that they may fairly be 
taken as representing a local race of the widely-spread C. mem- 
branipes (Ramb.). The character most readily empioyed for 
separating males of nemoricola from the typical membranipes is 
afforded by the colour of the upper anal appendages, which are 
blue in membranipes, black in nemoricola. 

The Kina Balu specimens show marked variability in size and 
in venation. 

Measurements of 5 males taken at random from amongst the 
15 male specimens that I have received from Mr. Moulton are as 
follows :— 

Length of abdomen 47 mm., 43 mm., 41 mm., 45 mm., 45 mm. 

- hind wing 30 mm., 28 mm., "26 mm. 30 mm. 30 mm. 

The number of postnodal costal nerves ranges from 16 to 21. 

The point of origin of M, and Rs varies greatly, as shown in 
the following table taken from the same males. 


M,, rises at subnodus. 

¢=") Re distal. 

is M, rises proximal to subnodus. 
Rs distal. 

a bia o.- ala 

ieee la” i b |b 


BORNEAN DRAGONFLIES, 23 


C@LICCIA OCTOGESIMA (Selys). | 

C. octogesima Kirby, Cat. Odonata, p. 128 (1890). 

TO. Matang Rd:, 3.1. 10. 

This specimen, unfortunately much damaged, agrees well with 
the description of the type female in the Synopsis. 

It differs from other females of the genus I have seen in the 
ereater length of the spine developed on the posterior margin of 
the prothorax, which projects forward at an angle of about 45° to 
the main axis of the body and is approximately 1 mm. in length. 

I have not been able to identify any of the species I have seen 
with (. borneensis Selys, and as the Bornean species are evidently 
numerous, and often very similar in appearance, it will be difficult 
to redetermine Selys’ species. 


Legion AGRION. 

SrENAGRION DuBIUM (Laidlaw). (Text-fig. 10.) 

Pseudagrion ? dubium Laidlaw, Journ. R. Asiat. Soc. Str. Br. 
Noe Oo, 1912. p. 97, pl. fig. d. 

Stenagrion dubiwm Laidlaw, Proc. Zool. Soe, London, 1915, 
Pp: ov. ; 

I take the opportunity of figuring here the venation of the 
type maie of this interesting species (text-fig. 10). It shows a 
development which parallels with remarkable exactitude that 
of Celiccia. Iam indebted for the photograph of the wings of 
this species, and of all the new species of Caliccia described in 
this note, to Messrs. H. and F. E. Campion, to whom I desire to 
tender my best thanks. 


Text-figure 10. 


Wings of Stenagrion dubium Laidlaw. Type g. 
Mt. Batu Lawi, Borneo, 27. v.11. 


‘TEINOBASIS SUPERBA (Selys). 

Teinobasis superba Kirby, Cat. Odonata, p. 157 (1890); Ris, 
Nova Guinea, xiil. Zool. pp. 100, 101, 122 (1915). 

1 S$. Lio Matu. 

The specimen belongs, I think, to the typical race of this species. 


2.32 ON BORNEAN DRAGONFLIES. 


ANISOPTERA. 
AVSCHNIDA. 


GOMPHIN#. 


HETEROGOMPHUS SUMATRANUS Kriger. 


Heterogomphus sumatranus Kriiger, Stett. Entom. Zeit. 1898, 
p. 294. 

5 $6 d. Retuh, 15-16.v. 14. 

Mi. J. C. Moulton tells me that this species has blue eyes. 
It is otherwise remarkable for the rich chestnut-brown colouring 
of the thorax and abdomen. ‘The latter has the apices of seg- 
ments 2—9 ringed with black. 

Lhe species is quite distinct from H. icterops Martin. 


| Nore.—Owing to the great difficulty of setting the wings of 
these very fragile insects, some distortion of the wing-base has 
occurred in the specimens photographed. ‘This is especially the 
case with the types of C. macrostigma and C. nigrohamata, con- 
sequently the figures of the wings of these species (text-figs. 5 
and 7) do not exhibit clearly the course of the “anal bridge” 
vein, which is, however, that normally found in the genus. | 


eo She SUSIE Fawcett. Pia 


J. M. Fawcett, del. 


HETEROCERA FROM BRITISH BAST ABNICA 
AND SOME OTHER EOCALITIES: 


MENPES PRESS, WATFORD. 


ON MOTHS FROM B. E. AFRICA. Doe 


15. Notes on a Collection of Heterocera made by M-. W. 
Feather in British East Africa, 1911-13. By Lt.-Col. 
J. M. FAwWceErTtT *. 


"Received October 10, 1917: Read November 20, 1917. | 


(Plate I.+) 


INDEX. . 
GEOGRAPHICAL : Page 
Localities at which collections were made .............ceeceseeeeeeeeene ene 23 
SYSTEMATIC: 
era COm COMa Mem we ans coutsns ccateh staat. etareaanes ish sab seb esieddi'stsen's 240 
Page Page 
Syntomis polyxo, sp. UN. ............6.. 284 | Lacipa albula, sp. ne... cee. BAZ 
PLUTO AGANACE, SP. Me... 0. ose. cc ees ves 235 | Hylemera lepta, sp. n. ............... 248 
Cladocerotis ceneus, Sp. Ne... .....6 +. 230 |) Eheterocrita Metis, Sp... 7 icis.sss30. 244 
Odontestra avitta, sp. Mn. ............ 236 DPROSEAUNOCSH, SP Wyo iecceess.eceesten GAN 
HAVESUICRLIS OFY1US, SP». N....... 0. 237 LPhOSCA Pervsets, SP. Ne... .c0ecceerses+. BAA 
Porastichtis lysis, sp.N. «........... 288 | Thosea chloris, sp. un. ...............-.. 245 
Giaura arethusa, sp. N................ 238 | Rhodoneura bryavis, sp. n. ......... 246 
Brevipecten icarus, sp.u............. 240 | Surattha rufistrigalis, sp.n. ...... 247 
Cynisca thisbe sp.n............. ..... 240 | Ancylolomia chrysolinealis, sp. n.... 247 
Pteronycta cervicornis, sp. n. ...... 240 | Ancylolomia gracilis, sp. n. ......... 243 
Rhodesana mintha, sp. n. ............ 241 Anthela nigristigma, sp. Me ...... 60 248 
PAleteronygmia aurelia, sp. un. ...... 242 | Nyctipao acuta, sp. 0. .......0...... 249 
EHuproctis rubricosta, sp. n. ......... 242 d 
et 


The object of the third memoir on this subject is to describe 
a few more of the remaining forms of this collection in my 
possession, and to figure some of the forms described in my 
second memoir, published in 1916, of which I was unable to give 
figures from want of space on the plate. 

The forms dealt with in the present memoir were taken at 
Kedai (altitude 2500 feet) and Masongaleni (altitude 3000 feet) 
in B. E. Africa, with the exception of a few specimens taken by 
Lady Colvile at Meru, B. H. Africa, At the end I have described 
two new forms from India and Australia, which have been some 
time in my possession awaiting an opportunity of publication. 

It will be noticed that among the forms from B. EK. Africa 
there are a good many which are typical of the desert, of which 
Euphiusa hermione, Ctenusa psamatha, Galactomoia berenice, 
described in my former memoir, and Cynisca thisbe, in this one, 
are notable examples, which seem to occur along with darker, 
richer-coloured forms such as we should expect to meet with 
in a country of considerable rainfall. Not being acquainted 
with the district myself, and having no notes on the subject 
by Mr. Feather, I regret [ cannot offer an explanation of this 
curious phenomenon. 

I notice that my first memow on this subject, published in 


* Communicated by the SECRETARY. 
+ For explanation of the Plate see p. 250. 


Proc. Zoot. Soc.—1917, No. X VII. Ih 


234 LT.-COL. J. M. FAWCETT ON 


P. Z.S. 1915, has evoked a certain amount of criticism, and 
that from a quarter from which I did not expect it, as one of the 
writers was good enough to help me in the thankless task of 
working out my specimens amongst the vast material in the 
British Museum Collection, and to agree with me at the time in 
the conclusions I arrived at. 

These criticisms refer to my descriptions of what I consider as 
new forms of Sphingide, and I would remark that until the pre- 
liminary stages and life-history of the specimens are known (and 
we are still in ignorance of them), I do not see how anyone, 
even the highest ‘author ity on the subject, can say as an absolute 
fact that such a specific name is a synonym of one already 
described. Such an assertion is merely a matter of opinion. 
In any case, it is desirable that undescribed forms from an 
extensive country like British East Africa, which has not been 
worked out to any great extent, should receive names in these 
days, when the Lepidoptera of each separate island in the 
Eastern Archipelago are being given distinct names as geo- 
graphical forms—even when obviously belonging to well-known 
previously described species. The reader has only to refer 
to Seitz, ‘Macrolepidoptera of the World,’ to verify what I say. 
In making the above remarks I do not, for a moment, deprecate 
criticism ; on the contrary, it is most interesting to me to read 
it, and, as I am a correspondent to the press on sporting subjects 
during the winter months, I get my fair share of it. 

The figures in the accompanying plate are drawn by myself to 
represent the exact size. In this memoir B.M. stands for 
British Museum, and C. L. P. for ‘ Catalogue of the Lepidoptera 
Phalenz,’ by Sir George Hampson. 

My best thanks are due to Lord Rothschild, Mr. Louis Prout, 
and Mr. J. H. Durrant, for their kindness and ever ready help 
in the British Mnseum. 


HETEROCERA 
Family SYNTOMID&. 


249. SyNTOMIS PoLYxo, sp.n. (PI. I. fig. 10.) 


Description.— $. Black, shot with coppery suffusion in certain 
lights. Antenne black with white tips; frons orange-yellow ; 
collar biack; patagia with orange-yellow stripes ; metathorax 
with an orange patch ; abdomen nee with six yellow bands. 
Fore wing with the following white diaphanous spots :—one 
below base of cell; an oblique diamond-shaped spot below middle 
of cell and vein 2; a spot in end of cell; an elongate spot above 
vein 6, and two spots between veins 3 and 5. Hind wing with a 
large basal spot between median nervure and inner margin, and 
a smaller one beyond end of cell, above vein 3. 

The plan of markings is similar to that of S. humeralis Butler, 
from North Australia (C. L. P. i. pl. it. fig. 15, p. 63); but this 


MOTHS FROM B. BE. AFRICA. 2935 


form is much smaller, and the spots are white, and not yellow 
as in that species. 

This is the smallest form of Syntomis that I am acquainted 
with, and the figure is slightly larger than natural size. 

Hahitat Kedai, 3lst December, 1911, 2 9; 15th March, 
Pies. | xpanse, ¢ 13, 9° 15 mm. 


Family ARCTIADAZ, 
Subfamily ARCTIANZ, 


250. ESTIGMENE TENUJSTRIGATA Hampson, Ann. S. Afr. Mus. 
i. peo (1900); C. L./P. pl. xiviil. fig. 16. 
Habitat. Masongaleni, 18th April, 1911, 1° 


250 a. Estig@MENE LeMNiscata Distant, A. M. N. H.° (7) i. 
pe Vi6(1898)3) C.L. P. pl. xivii. fig: 20: 

Habitat. Meru, B. E. Africa, taken by Lady Colvile, 1 ¢. 

This specimen is much more heavily marked, as regards the 
black bands, than the figure above quoted, and, moreover, has a 
black band along the costa. 


250 6. EstigMENE LINEA Walker, Cat. il. p.671 (1855); C. L. P. 
ple xlvil. fig. 2. 

Hubtiat. Meru, B. BE. Africa, taken by Lady Colvile, 2 3. 

These specimens agree fairly well with the figure in C. L. P., 
but one example is much lighter in tone than the other, whilst 
both have more heavy black : markings. 


251. TERACOTONA SUBTERMINATA Hampson, C. L. P. iil. p. 471, 
ple. ie. 16 (1901), 
fabitat. Masongaleni, 18th April, 1911, 1 ¢. 


Family Hy psip@. 


252. BITHRA AGANICE, sp. n. (PI. I. fig. 11.) 

Description.—-9 . Head and thorax pale grey-brown ; antenne 
black, and pectinate for half the length, filamentous thence to 
tip. ali black spots on collar, two on euch of the patagia, and 
one on metathorax. Abdomen pale orange, with a dorsal row of 
black spots on first three somites, and a interval series on each 
somite. Fore wing pale grey- on with the following black 
points: two basal; two subbasal, one of which is near the base 
of costa, and one below base of cell; two antemedial one of 
which is below cell, and one above vein 1; one in cell before 
middle, and one in interspaces 3, 4, and 6, near base, and beyond 
cell. Hind wing dull yellow. Underside similar to upperside, 
uninarked. 

This form is nearest to Bithra aganois Felder, which has only 
one basal black spot and one in cell. 


Habitat. Kedai, 7th January, 1912,1 29. Expanse 86 mm. * 
Lie 


236 LT.-COL. J. M. FAWCETT ON 


Family NocTUID.. 
Subfamily AGROTINE, 


253. CLADOCEROTIS GNEUS, sp. n. (PI. I. fig. 12.) 


Description.— $. Head, thorax, abdomen, and fore wing pale 
red-brown. Head and thorax densely covered with thick long 
hair ; abdomen with lateral tufts of hair. Fore wing with the 
veins, orbicular and reniform pale ochreous; reniform defined 
outwardly by black lines. Interior of cell dark red-brown, with 
a black line below costa from base to middle. Claviform very 
elongated and black, from base to beyond middle. An irregular 
series of black wedge-shaped spots subterminally between veins 4, 
5, and 6, double the size of those between the other veins. A 
fine black terminal line. Ciba ochreous on both wings. Hind 
‘wing with the basal area ochreous and the terminal area pale 
fuscous. 

This form seems fc be allied to C. optabilis Boisd., from Europe, 
but is larger than the figure in C. L. P. and Sawek darker, the 
claviform is black, not ochreous defined by black, the marginal 
spots are irregular, and the hind wings ochreous and pale fuscous 
instead of white as in that species. 

Habltat. Meru, taken by Lady Colvile, 2 ¢, 19. Expanse 
36 mm. 

The 2 specimen is darker in colour, the markings are more ill- 
defined, and the antenne are not so highly pectinated. 


254, PorosacGroris CAFFRA Hampson, ©. L. P. iv. p. 658, 
pl. Ixxvul. fig. 29 (1903). 
Habitat. Meru, taken by Lady Colvile, 1 9. 


255. ADISURA ATCINSONI Moore, P. Z.S. 1881, p. 368, pl. xxxvii. 
fiee6; 

Habitat. Kedai, 6th March, 1912,1 

This form is very variable, and the specimen under notice has 
darker hind wings than some specimens in the B.M. from 
British East Africa. 


256, CHLORIDEA OBSOLETA Fabricius, Ent, Syst. 3, 1. p. 546 
(1793). 


Habitat. Kedai, 16th March, 1912,1 3. 
Formerly known as armigera Hiibner, Samui, is Schmett. 


fig. 370 (1827). 
Subfamily HapvEniInz&, 


257, ODONTESTRA AviTTA, sp, n. (PI. I. fig. 15.) 


Description.— 8. Head, thorax, and abdomen pinkish brown. 
Fore wing pinkish brown, chestnut in the medial area, Sub- 
basal and antemedial lines represented by a single thick black 


MOTHS FROM B. E. AFRICA, yaae 


line, Claviform short and thick, black. A black line on inner 
margin from base below vein | to antemedial line. No ochreous 
white fascia on medial part of inner margin. Orbicular and 
reniform large, extending over costa, pinkish defined outwardly 
by indistinct ochreous. Postmedial line excurved round reni- 
form and thence straight to inner margin—a thick black line. 
An indistinct subapical spot below costa. Hind wing white with 
some ochreous irroration near the apex. Underside pale ochreous, 
with discocellular spots and diffused postmedial lines. 

This form differs from vittigera Hampson, from Mashonaland 
and Natal, in being without the ochreous-white fascia on medial 
part of inner margin, that part of the wing being concolorous 
with the remainder; also in the following characters :—-the 
orbicular and reniform are larger and spread out over the costa, 
and are pinkish in colour; the ante- and postmedial lines are 
thick black single lines, not double lines filled in with ochreous ; 
the subterminal area beyond the postmedial Jine is plain pinkish 
brown, without a trace of any marks save a small subapical spot 
below the costa. 

It also differs from albivitta Hampson (C. L. P. v. p. 206) in 
being without the whitish fascia above the claviform, which 
species is also recorded from British East Africa. 

Habitat. Kedai, 8th December, 1911, 1 ¢. Expanse 30 mm. 


Subfamily AcRoNYCTINA. 


258, PERIGEA CAPENSIS Guén. Noct. 1. p. 213 (1852), 
ITabitat. Masongaleni, 22nd June, 1911,1 ¢. 


259, LapHyema ExempTa Walker, Cat. x. p. 855 (1856). 


Habitat. Kedai, 1911, 24th November, 1 ¢ ; 25th November, 
2G 


260. PARASTICHTIS OXYLUS, sp. n. (PI. I. fig. 22.) 


Description.— 3. Head, thorax, abdomen, and fore wing dark 
red-brown with black-brown markings. Fore wing: subbasal 
and antemedial lines only represented by strigeze on the costa. 
Orbicular and reniform defined by fine black lines with a dark 
shade on and below the costa between them; an irregular dentate 
postmedial line angled inwardly above veins 4 and 6, and out- 
wardly at veins 4 and 3, with some black lines running from it 
to the outer margin. A black band starting from base under 
vein 1, angled up to vein 2, and running below it to postimedial 
line, and thence on to outer margin. Hind wing pale ochreous 
basally and fuscous towards the apex. Cilia of fore wing black 
and ochreous chequered, of hind wing whitish. Underside pale 
ochreous. 


Habitat. Kedai, 19th November, 1911,1 ¢. Expanse 34 mm. 


238 LY.-COL. J. M. FAWCETT ON 


261. PARASTICHTIS Lysis, sp. n. (PI. I. fig. 25.) 

Description.— 3. Head and thorax bright ferruginous, abdo- 
men reddish ochreous. Fore wing bright ferruginous, the sub- 
terminal area beyond the postmedial line pale ochreous. Orbicular 
and reniform defined outwardly by fine black lines and with a 
dark shade below the costa between them. Subbasal and ante- 
medial lines obsolete. A black line from the base of interspace 1 
to middle, where it is recurved upwards and then inwards. 
Another black line from base below vein 1, with a small black 
spot on inner margin below it. Some black streaks with bright 
red edges between the veins on the subterminal area. Post- 
medial line highly dentate. Hind wing pale ochreous, darker 
towards the termen. Underside pale ochreous, with fuscous 
discocellular spots and diffased postmedial bands. 

Habitat. Kedai, 1911, 10th November, 1 ¢ ; 18th November, 
1 g. Expanse 34 mm. 

Of the large number of species (39) of the genus Parastichtis 
given in C. L. P. vil., only one is recorded from Africa, nigri- 
costata Hampson, which is a totally differently-coloured insect. 

The first of the forms under notice may always be known by 
the prominent curved black band below vein 1,and the second by 
its bright ferruginous colour. 


Subfamily SARROTHRIPIN &. 


262. GIAURA ARETHUSA, sp. n. (PI. I. fig. 14.) 
_ Description.— $. Head, thorax, and abdomen grey. Fore 
wing grey, minutely irrorated and striated with fuscous. Subbasal 
line represented by a V-shaped striga on the costa, Antemedial 
line represonted by another costal striga which gradually becomes 
obsolescent in the cell, with a dark shade beyond it. Postmedial 
line outwardly oblique to vein 4, then bent inward, and becoming 
obsolete before reaching the inner margin. A highly dentate 
submarginal line with diffused fuscous shading beyond it, from 
apex to near tornal angle, where there is a black spot before it. 
A fine marginal black line. Cilia grey. Hind wing pale ochreous 
at the base, terminal area fuscous black. Cilia ochreous. Under- 
side of fore wing ochreous, thickly irrorated with fuscous towards 
the margin ; hind wing white with a black subterminal band. 

Habitat. Kedai (date not recorded), 1 g¢. Expanse 34 mm. 


Subfamily HRAsTRIANz. 
263. OZARBA SUBTERMINALIS Hampson, P. Z. 8. 1910, p. 407; 
Cie. x. pl. clayton: 
Habitat. Kedai, 27th November, 1912, 1 3. 


264. OzarBa vARIA’ Walker, Cat. xxxill. p. 772 (1865). 


Habitat. Kedai, 5th January, 1912, 1 ¢. 
This specimen has orange hind wings in contradistinction to 


MOTUS FROM B. E. AFRICA, 239 


some specimens in the B. M. Collection ; but the species is exceed- 
ingly variable. 

265. OZARBA SINUA Hampson, C. L. P. x. p. 414, pl. clx. fig. 31 
OES LT). 


Habitat. Kedai, 18th November, 1911,1 ¢. 
‘This specimen represents a very light form of the species. 


266. OzarBA ABscissA Walker, Cat. xv. p. 1764 (1858). 
Habitat. Kedai, 9th December, 1911, 1 ¢. 


Subfamily ACONTIAN A. 


*148. Leocyma CANDACE Fawcett, P. Z. 8S. 1916, p. 712. 
GE fie, 20.) ; 

As promised in my former paper, quoted above, I am now 
able to give a figure of this species, which the reader is requested 
to compare with the woodcut figure of camila Druce, in C. L. P. 
x1. p. 663, and that of discophora Hampson, pl. exci. fig. 19. 


Subfamily CaTocaLin®. 


*152, CreNUSA RECTILINEA Fawcett, P. Z. S. 1916, p. 713. 
PBI ities 3.) 
A figure is here given of the typical form : no. 2 form psamatha 


was figured in my memoir published in 1916 (P. Z. 5S. 1916, 
Fawcett, Pl. I. fig. 21). 


*162. Cortyta minyas Fawcett, P. Z. 8. 1916, p. 715. (Pl. I. 
ie. 1%) te 
CorTYTA GRISEACEA Fawcett, P. Z. 8. 1916, p. 718. (PI. I. 

fig. 18.) 

Figures are now given of the above species. As both minyas 
and griseacea have been taken in December 1911 at Kedai, they 
can hardly be seasonal forms of the same species, and [ am now 
of the opinion that they are distinct forms. 


267. PLECOPTERODES MODERATA Wallengren, Wien. Ent. Mon. 
iv. p. 174 (1860). 
Habitat, Masongaleni, 22nd April, 1911, 1 6. 


Subfamily Nocruinz. 


*181. PLECOPTERA POLYMNIA Fawcett, P. Z.S. 1916, p. 722. 
(PL. I. fig..23.) 

A figure is here given of this form. The reader has only to 
compare it with the figure of polymorpha Hampson, P. Z. 8. 
1916, pl. 1. fig. 3, to see how it differs from Hampson’s species, 
which is clearly a desert form. 


* These numbers refer to my paper in P. Z. 8, 1916, p. 707 et seq. 


240 LT.-COL. J. M. FAWCETT ON 


268. BREVIPECTEN ICARUS, sp.n. (PI. I. fig. 24.) 


Description.— 6. Head and thorax grey; abdomen ochreous. 
Fore wing with the basal half grey, the costa tinged with ferru- 
ginous. Subbasal line represented by two black spots below 
the costa and on the median nervure. Antemedial line very 
faint and indistinct. A square black patch before end.of cell, 
with two small black spots above it on costa, separated by 
a, patch of ferruginous suffusion, and with a patch of ferru- 
ginous suffusion below it on interspaces 1, 2, and 3, before 
the postmedial line. Reniform large, represented by a 
grey patch, beyond which the postmedial line is angled out- 
wardly at vein 6, and is then evenly curved inwards to inner 
margin, A black patch on costa before apex with ferruginous 
suffusion before it. Subterminal area pale brown, with a mar- 
ginal row of indistinct wedge-shaped spots between the veins. 
Hind wing ochreous, darker towards the margin. Cilia ochreous. 
Underside: medial areas of both wings grey, subterminal area 
pale fuscous ; costa ochreous, with some black strigz on that of 
hind wing, and a discocellular spot. 


Habitat. Kedai, 28th November, 1911,1 ¢. Expanse 32 mm. 


Genus Cynisca, nov. 

Type, C. thisbe, sp. n. 

Allied to Calpe. Proboscis fully developed. Palpi long, porrect, 
the 2nd and 38rd joints fringed with long hair above and below. 
Antenne eiliate; metathorax with slight tufts of hair, which also 
appear on dorsum of abdomen on first three somites. Costa of 
fore wing straight, not arched. Apex rectangular, not acute. 
Outer margin crenulate, evenly rounded. Inner margin with 
a very pronounced lobe at middle, and a smaller lobe at tornal 
angle. Venation as in Calpe. 


269. CYNISCA THISBE, sp. n. (PI. I. fig. 13.) 


Description.— 2. Head, thorax, and abdomen ochreous white. 
Antenne red. Fore wing pale brown, shading into pale purple 
on the centre lobe and the area above it, and witha purple shade 
on outer margin. Antemedial line red-brown from costal ner- 
vure to inner edge of lobe, with ashort line beyond it. A medial 
line from end of cell to outer edge of lobe, where it is bent 
inwards and becomes double. A very indistinct dentate post- 
medial line from before apex to outer lobe. Cilia pale brown. 
Hind wing white, with a pale ochreous shade on the veins on 
outer margin. Cilia white. Underside ochreous white, marked 
only by the postmedial band of the upperside showing through. 

[fabitat. Kedai, 9th March, 19138, 1 9. Expanse 42 mm, 

This is clearly a well-marked form of the desert. 


270. PrERONYCTA CERVICORNIS, sp. n. (PI. I. fig. 21.) 


Description.— 3. Head, thorax, abdomen, and fore wing pale 
pinkish grey. Fore wing: subbasal and antemedial lines only 


MOTHS FROM B. E. AFRICA, 241 


indicated by a couple of black spots. Orbicular stigma, a fine 
black point with a fine black line round it. Reniform indistinct. 
Postmedial line represented by a double row of black points 
bent in to inner margin. <A broad red-brown band from base 
along median nervure to end of cell, where it is bent up into a 
tooth, then curving down to outer margin at vein 4. The area 
below this band reddish towards base. Some small ferruginous 
stripes between veins 3, 4, and 5 submarginally. Hind wing 
grey, the terminal area fuscous. Cilia ochreous. Underside 
pale ochreous, with a fine striga at end of cell of fore wing. 

Habitat. Kedai, 13th January, 1913, 1 6. Expanse 40 mm. 

IT have named this form cervicornis, from a fancied resemblance 
of the broad red band to a stag’s antler. 

The only species in the B. M. Collection which seems to be 
near this form is the recently described Pteronycta Jasciata 
Hampson, from Zomba in Nyassaland, which is a much larger 
insect. 

Subfamily Hypenina. 


*189,. Hyprena sussaALis Walker. 
Habitat. Kedai, 1911, 29th December, 1 ¢; 30th December, 1d. 


HYPENA STRIGATA Fabricius. 


Habitat. Kedai, 25th November, 1911, 1 6; 3rd January, 
Poe lis. 

In my former paper I mentioned that the specimens of Hypena 
in the Feather collection represent two forms, a grey and a 
brown form. I now find that the grey form is jussalis Walker, 
and the brown form strigata Fabricius. 


Family LYMANTRIAD&. 


271. CASAMA VILIS Walker. 


Habitat. Masongaleni, 25th June, 1911,1 9. 

Specimens of this species in the B, M. Collection are nearly all 
from Ceylon and Arabia, but there are a few from the Red Sea 
littoral which agree with this specimen. Clearly a desert form. 


272. RHODESANA MINTHA, sp. n. (PI. I. fig. 19.) 


Description.— 6. Head, thorax, abdomen, and fore wing 
chestnut red-brown. Antenne bipectinate. Abdomen with 
dorsal tufts. Fore wing thickly irrorated and striated with 
black. A subbasal black line from costa to vein 1. Antemedial 
line irregularly waved and bent inwards on vein 3, and thence 
more or less straight to inner margin. A line of wedge-shaped 
‘red-brown submarginal spots between the veins. before termen. 
Cilia red-brown, whitish at the extremities of the veins, Hind 
wing ochreous, irrorated with red-brown atoms, with a somewhat 
indistinct medial line across the dise. Underside ochreous, with 
indistinct brown medial band. 


242 LT.-COL. J. M. FAWCETT ON 


Habitat. Masongaleni, 27th March, 1911, 1¢; Kedai, 27th 
November, 1912, 2 ¢. Expanse 32 mm. 

This form is nearest to /t?. crenulata Bethune Baker, but is 
much smaller and redder than that species. It also differs in the 
postmedial line being angled in on vein 3, and there being 
a medial line across the disc of the hind wing. 


273. HETERONYGMIA AURELIA, sp. n. (PI. I. fig. 16.) 

Description.— 3. Head and thorax golden yellow, abdomen 
ochreous. Antenne bipectinate. Palpi long, thickly covered 
with long hair. Tore wing golden yellow ; no antemedial lines. 
Three oblique lines, red- Drown: and leading in the shape of a 
V from the costa to a dark red-brown shade at end of cell. A 
narrow waved oblique postmedial line from costa just before apex 
to middle of inner margin. A subterminal line of black points 
between the veins. Cilia reddish yellow. Hund wing ochreous ; 
a fuscous spot at tornal angle with a fuscous striga above it, and 
three fuscous discal spots between veins 3, 4, 5, and 6. 

Underside: fore wing bright ochreous, with a fuscouse band on 
discocellulars, and a thicker fuscous postmedial band than that 
on the upperside ; hind wing bright ochreous, with medial band 
and discal spots as on the upperside. 

Habitat. Meru, B: KE. Africa, taken by Lady Colvile. Expanse 
o8 mm. 

There are no specimens of this genus in the B. M. Collection, 
but there is a figure of Heteronygmia stigmatica Holland, from 
Gaboon, which appears to be near this species, but it has no 
black oblique postmedial line as in this species. 


274. EupRoctris RUBRICOSTA, sp. n. (PI. I. fig. 7.) 

Description. 3. Head, thorax, and wings pure shining white; 
antenne bipectinate and ochreous, base of shaft white; legs 
thickly covered with long white hair. Abdomen with the first 
three somites white and the terminal three golden yellow. 
Underside white, with a red-brown band along the costa from 
base to Just before apex. 

Habitat. Masongaleni, 1911, 28th May, 1 ¢ ; 25th December, 
12. Expanse 32 mm. 


274 a. LACIPA ALBULA, sp. n. (PI. I. fig. 8.) 

Description.— 3. Head, thorax, and wings pure shining white. 
Antenne ciliate. Abdomen golden yellow with the exception of 
the first somite and the anal extremity, which are white. Fore 
wing unmarked, save for a shade on the costa beyond cell, and 
on inner margin before tornal angle. Hight black points on the 
margin between the veins, seven of which also appear on the hind” 
wing. Underside: fore wing with the basal and internal area 
white, the costa and half of cell occupied by a red-brown band, 
and short reddish bands between the veins distally, above vein 3. 
The black marginal poimts more distinct than on the upperside. 


“MOTUS FROM B. E. AFRICA. 243 


Habitat. Masongaleni, 1911, 24th March,1 ¢; 29th December, 


1 g. Expanse 24 mm. 
This form is near to Lacipa sexpunctata Distant, but is with- 


out the black spots and yellow ante- and postmedial lines. 


Family GEOMETRID4&. 
Subfamily BOARMIANA. 


275. KYLOPTERYX ARCUATA Walker. 
Habitat. Masongaleni, 21st June, 1911,1 ¢. 


9276. HonorANA EREBARIA Guen. 
Habitat. Kedai, 30th December, 1911, 1 9. 


277. HyLEMERA LEPTA, sp.n. (PI. I. fig. 29.) 

Description.— $. Head and pectus yellow. Thorax and first 
three somites of abdomen white above, yellow below, and con- 
colorous with the terminal somites which are yellow. Fore wing 
white ; the costa fuscous, and a broad apical fuscous fascia the 
inner margin of which is angled outwardly between veins 3 and 4. 
Hind wing white, with a very narrow fuscous border, which is 
almost linear, between veins 2 and 6, 

Underside as on upperside. 

Habitat. Kedai, 1911, Ist December, 1 ¢ ; 12th December, 
19. Expanse 36 mm. 

This species differs from doleris Plotz, accra Swinhoe, and 
ansorgei Warr., from Uganda, in the black band of the hind wing 
being narrow and continuous, the above forms having a large 
black spot on the apex of the hind wing: from dexithea Druce, it 
differs in not having the inside of the fascia of the fore wing 
evenly curved, and a black spot below apex of hind wing. 


278. EURRANTHIS PENNIGERARIA Hubner. 
Habitat. Meru, B. BE. Africa, taken by Lady Colvile. 


Subfamily ACIDALIIN#. 
279. ACIDALIA SAGITTILINEA Warr. Nov. Zool. iv. p. 219 (1897). 
Habitat. Kedai, 8th April, 1912, 1 ¢. 


280.. MIcROLOXIA RUFICORNIS Warr. 
Habitat. Kedai, 29th December, 1912, 1 9. 


Subfamily GEoMErriIn #. 


231. TRIMETOPIA ZTHERARIA Guén. 

Hubitat. Kedai, 1911, 4th January, 1 9 ; 23rd November, 
1 $; 25th November, 1 2 ; 17th December, 1 3. 

One of the most beautiful forms of Geometride. 


244 LT.-COL. J. M. FAWCETT ON 


282. HetTERocrita METIS, sp.n. (PI. I. fig. 30.) 


Description.— 2. Head pinkish brown, the vertex between the 
antenne white. Antenne white above, pink beneath. Thorax 
and wings bright grass-green. Abdomen pale pinkish brown 
above, ochreous on the under surface, the first two somites 
bright pinkish. Wings unmarked except for a bright pink-. 
brown marginal line, inside which are two minute white spots 
defined inwardly by pinkish brown between veins 5 and 7, and a 
large one of the same colour at tornal angie of each wing. The 
inner margin of the hind wing also defined by a pinkish-brown 
line. Cilia pale pink, Underside glaucous grey. 

Habitat, Kedai, 20th February, 1912, 19. Expanse 30 mm. 

This form is near to, and may, perhaps, be a local race of 
Heterocrita deprensa Prout, from which it differs in (1) having 
no discoidal spots, (2) the spots at the tornal angles of both 
wings are much larger, and (3) the abdomen is much more pink- 
coloured. 


\ 


283. HURYTHECODES FLAVEDINARIA Guén. 
fTabitat. Meru, B. E. Africa, taken by Lady Colvile. 


Mr. Prout informs me that this is a common and very variable 
species, but I have only one specimen of it. 


Family LIMACODIDA. 


284. 'THOSEA GANALE Pagenstecher. 
Habitat. Kedai, lst December, 1911, 1 ¢. 


285. THosEA UNDOSA, sp. n. (PI. I: fig. 6.) 


Description.— 2. Head, thorax, and abdomen pale ochreous 
brown. Fore wing with the basal area as far as the postmedial 
line red-brown, the distal area beyond it pale ochreous brown. 
Subbasal and antemedial lines absent; a black point at end of 
cell. A broad, dark brown oblique postmedial line, waved and 
irregular, from costa to inner margin at middle. <A fine dark 
brown subterminal line from costa to tornal angle, where it 
almost meets the black marginal line. Cilia pale brown in both 
wings. Hind wing ochreous brown, with a brown marginal line. 
Underside ochreous brown, irrorated with red-brown atoms, . 

Habitat. Kedai, 4th December, 1911, 1 2. Expanse 30 mm. 

This form is nearest to 7. cana Walker (=transversata 
Walker) from India, in which the dark line is antemedial, less 
oblique, and straight without waves; the subterminal line is 
straight, and not curved, and the basal area is not so dark as in 
this species. 


286. THOSEA PERSEIS, sp. n. (PI. I. fig. 4.) 


Description.— 3. Head, thorax, abdomen, and fore wing red- 
brown minutely irrorated with black atoms. Subbasal and ante- 


MOTHS FROM B. E. AFRICA. 245 


medial linesabsent. A white pointatend of cell. A black highly-. 
waved postmedial line from costa before apex to middle of inner 
margin, which is inwardly diffused into black patches below end 
of cell, and from vein 2 to inner margin. ‘This line is outwardly 
defined by a prominent white line, with dark shades beyond it 
between veins 2 and 3 and 6and/7. An irregularly curved sub- 
terminal line followed by a black marginal line. Cilia of both 
wings red-brown. Hind wing ochreous. Underside ochreous, 
with indications of a postmedial line. 

Habitat. Masongaleni, 30th April, 1911,1 ¢. Expanse 30mm. 

This form is nearest to 7’. syrtis Schaus, from Salisbury and 
West Africa. It differs in the lnes being waved and not 
straight, and that species has not the dark patches inside the 
postmedial line. When at the B.M. I submitted the above 
specimens to the inspection of Lord Rothschild, who said the 
forms were unknown to him, and differed from described forms 
mainly in their undulating lines. 


287. THOSEA CHLORIS, sp.n. (PI. I. fig. 5.) 


Description.— $. Head and thorax red-brown beneath, frery 
orange dorsally. Abdomen red-brown, with some fiery-orange 
hairs on the dorsum. ‘Tibi fiery orange, with long red-brown 
hairs to the tarsi. Fore wing red-brown with pinkish suffusion. 
A white point at end of cell. No subbasal or antemedial lines. 
Postmedial line dark brown, from costa before apex to middle of 
inner margin, Subterminal area paler. Subterminal line from 
costa before apex to outer margin at vein 3, Two whitish spots 
between veins 2 and 3: one outside the postmedial line, and 
one on inside of subterminal line, near its junction with outer 
margin. Hind wing red-brown paling to ochreous at base. 
Underside pale red-brown unmarked. 

Habitat. Ashanti, W. Africa (4. Norris), 1g.  Expanse 
30 mm. 

This form is nearest to 7. rara Swinhoe, from Thyetmyo. 
Burma, but is darker red, and has fiery orange head and thorax 
which rara has not. In the B. M. Collection there is an unnamed 
specimen from the Gold Coast, which may or may not. be refer- 
able to this species. 


288. CoSUMA MARGINATA Holland. 


Habitat. Masongaleni, 7th April, 1911, 1 3. 
This form seems near to the following which I have from 
W. Africa. 


289. CosuMA RUGOSA Walker. 
Habitat. Ashanti, W. Africa (A. Norris), 1 3. 


290. APLUDA INCINATA Hampson. 
Habitat. Masongaleni, 1911, 2 3. 


246 LT.-COL. J. M. FAWCETT ON 


291. GAVARA VELUTINA Walker. 


Habitat. Kedai, 1912, 16th March, 13 ; 10th December, 1¢. 

I identified these specimens in the B. M. as above; there are 
however, two forms of Limacodide which have passed as “ velu- 
dina,” and are given by Hampson in Faun. Brit. Ind., Moths, i. 
pp. 3882 & 398, which are very different-looking insects from 
the present form. : 


292. ALTHA LACIDES Druce. 
Habitat. Masongaleni, 1911, 25th December, 1 ¢ ; 2nd April, 
LO 


Family ARBELIDA. 


293. ARBELODES TETRASTICTA Hampson. 

flabitat. Kedai, 12th January, 1912, 1 ¢. 

This specimen only differs from the specimen in the B. M., 
which is also from British Hast Africa, in the body being on 
what shorter, and the pale markings of the wings being larger. 


Family Coss1pD&. 


*228. Duomitus. pInDARUS Fawcett, P. Z. S. 1916, p. 733. 
GEL he. 20:) 

I am now able to give a figure of this species, which was 
omitted from my last paper fron want of space in the plate. 

Comparison of fignre 27 with that of D, stewiptera Hampson, 
PZ. IS. 9OiG: plate u. fig. 31, will show how much it differs - 
from that species both in colour and size, 


Family THYRIDID4&. 


294, RHODONEURA BRYAXIS, sp.n. (PI. I. fig. 9.) 

Description.— 9 . Head, thorax, and abdomen ochreous suffused 
with rufous. Fore wing reddish ochreous profusely striated 
with rufous. Antemedial line indistinct. Postmedial line bifid 
on the costa, originating from two costal spots, and bent in 
obliquely on veins 2 and 3. Subterminal line very prominent, 
from costa before apex, angled outwards to outer margin at 
veins 4 and 2. Hind wing with a medial double line, a post- 
medial line, and a Sabecrminal line angled to outer margin at 
veins 4 and 2. 

S. Smaller and paler in coloration, being pale ochreous. 
The postmedial line on the fore wing and the medial line on the 
hind wing double and filled in with rufous ; otherwise as in @. 

Habitat. Kedai, 1911, 26th November, 1 ¢ ; 15th December, 
loo 1D 3th January, 1 ei ; 12th March, 1 ©; Sth Decem- 
ber, 1 ¢. Expanse, 6, 287mm 230 mm: 

This form differs from all the specimens of Rhodoneura in the 
B. M. Collection in its prominent subterminal line angled to 
outer margin at veins 4 and 2. 


MOTHS FROM B. E. AFRICA. : 247 


295. DysoDIA INTERMEDIA Walker. 
Habitat. Kedai, 5th January, 1912, 1 ¢. 


The; above example agrees with B. K. African specimens in 
the B. M. Collection which were collected at Taveta by Rogers in 
October and at the end of December, and named flavillula by 
Warren, and which Sir George Hampson evidently regards as 
forms of Walker’s South African species. 


Family PyrawuipDa, 


Subfami'y CRAMBIN&. 


296. SURATTHA RUFISTRIGALIS, sp. n. (PI. I. fig. 3.) 


Description.— 3. Head, body, and legs ferruginous. Fore 
wing brownish ochreous, ferruginous on costa. Highly dentate 
medial and postmedial ferruginous lines from costa to imner 
margin, angled outwardly at vein 5. A ferruginous terminal 
line with dark wedge-shaped spots between the veins. Median 
nervure and inner margin defined by bright ferruginous bands ; 
the interspaces between the veins defined with fuscous irroration. 
Hind wing ochreous, with ferruginous irroration in the subapical 
area: cilia ochreous. 

Habitat. Kedai, 18th November, 1911, 13. Expanse 24 mm. 

The figure is slightly Jarger than natural size. 

This form is nearest to S. africalis Hampson, but has no dark 
medial area; the postmedial line is more dentate, and the 
coloration is brighter ferruginous. 


297. ANCYLOLOMIA SIMPLELLA De Jaan. 
Habitat. Meru, B. EB. Africa, taken by Lady Colvile, 1 9. 


298, ANCYLOLOMIA PECTINIFERA Hampson. 


Habitat. Kedai, 9th January, 1913,1 9. 
Also recorded from Somaliland. 


299. ANCYLOLOMIA CHRYSOLINEALIS, sp.n. (PI. I. fig. 2.) 


Descripiion.— $. Head, thorax, abdomen, and fore wing very 
pale ochreous. Fore wing with eight golden stripes in the inter- 
spaces between the veins in the distal area beyond the cell, of 
which those in interspaces | and 7 are shorter than the remainder. 
Each golden stripe is defined beneath by a yellow fascia with 
streaks of black scales on it. A minutely dentate silvery sub- 
marginal line on which are some dusky points. Cilia silvery. 
Hind wing pure white; cilia white. Underside as on upperside, 
with silvery suffusion. 

Habitat. Masongaleni, 26th December, 1911, 1 ¢. Expanse 
34 mm. 

This form is nearest to A. chrysographella Kollar, from: India, 


248 LT.-COL. J. M. FAWCETT ON 


but is distinguished by having the golden stripes of the fore 
wing confined to the distal area, and it is considerably larger than 
chrysographella. 


300. ANCYLOLOMIA GRACILIS, sp. n. (PI. I. fig. 1.) 


Description.— 3. Head, thorax, and abdomen pale ochreous 
brown. Fore wing bright ochreous, unmarked; the interior 
marginal area below vein 1 thickly irrorated with fuscous scales. 
A series of black points at the end of the veins on the outer 
margin. Hind wing silvery grey, with ochreous suffusion on the 
subapical area. Cilia ochreous. Underside as on upperside, with 
silvery suffusion. 

Habitat. Meru, B. E. Africa, taken by Lady Colvile, 1 ¢. 
Expanse 28 mm. 

There is a series of unnamed specimens in the B, M. Collection, 
which appear to be very near this species. 


Subfamily ANERASTIANZ. 


301. EMATHEUDES LENTISTRIGALIS Hampson. 
Habitat. Masongaleni, 6th June, 1912, 1 9. 


New Forms from other Regions. 


Family LYMANTRIADA, 


~ 


302. ANTHELA NIGRISTIGMA, sp.n. (PI. I. fig. 28.) 


Description.-— 3. Vertex of head and collar ferruginous ; 
thorax pale red-brown ; abdomen ochreous. Wings pale ochreous 
with golden suffusion in certain lights. Base of fore wing and 
internal area of hind wing clothed with long pale red-brown 
hair. Fore wing with a round black spot in centre of cell, and a 
larger quadrate black spot on discocellulars. A somewhat in- 
distinct line of submarginal black points cn the veins in the 
subterminal area. Hind wing with a quadrate discocellular 
black spot, and a submarginal series of black points on the veins. 

Underside concolorous with upperside, with an additional black 
spot in the cell of the hind wing which does not appear on the 
upperside, and all the black spots have whitish centres. Cilia 


ochreous. 
Habitat. Townsville, Queensland, Australia, August 1913,1 ¢. 


Expanse 36 mm. 

The above specimen was sent to me by Mr. Feather in a small 
collection from Australia. In the B. M. Collection there is a 
series of unnamed specimens which may, or may not, be referable 
to this species. In some of them the cellular spots are a good 
deal larger and the submarginal series “more distinct. This 
species is near to ocellata Walker, which has ‘the submarginal 
spots black and prominent. 


MOTHS FROM B. E. AFRICA, 249 


Family NocruipD4. 
Subfamily CATocALIN#, 

303. Nycrrpao acuta, sp.n. (PI. I. fig. 26.) 

Deser iption.— 3d. Head, thorax, and abdomen fuscous brown. 
Fore wing with the basal two-thirds very dark red-brown 
the subterminal third pale red-brown with pinkish suffusion, 
profusely striated with red-brown. Apex of fore wing acute. 
Antemedial line obsolescent, and hardly distinguishable. An 
inverted comma-shaped discoidal whorl with bilobate head defined 
by black, and its head by silvery blue with a red-brown patch 
beyond it; its centre reddish brown with an ochreous line on 
inner side of tail. Medial line black, arising below the costa, 
excurved round the discoidal whorl, and angled inwards below it 
on vein 2, and then erect to inner margin. This line is defined 
outwardly by a pale round ring beyond the whorl. Postmedial 
line curved, starting from a large triangular black patch below 
the costa and continued as an indistinct fuscous line to below 
vein 2, where it forms the outer edge of the basal dark area 
to inner margin. ‘This line is defined outwardiy by a narrow 
ochreous line from costa to below vein 7, and thence to inner 
margin by an indistinct line of the ground-colour. Subterminal 
line obsolete. Hind wing with the basal two-thirds dark red- 
brown, the terminal third pale red-brown with pinkish suffusion ; 
the postmedial line represented by double fine lines of the ground- 
colour; subterminal line represented by an indistinct, waved, 

subterminal shade. Cilia pale red-brown. 

Underside of both wings fuscous brown; fore wing with a 
white postmedial patch from below costa to vein 6, where it is 
broadest. A subterminal series of white oblong spots, incurved 
in interspace 4, excurved in interspace 3. 

9. Similar to gd, but the comma-shaped discoidal whorl is 
somewhat larger, and there is a pale ochreous postmedial patch 
from costa to just above vein 6, diffused outwardly. 

This form has, I think, been overlooked by later writers; I 
say this because I cannot find amongst the described forms in 
C. L. P. vol. xii. (published in 1913) a description and figure 
that absolutely tallies with my specimens. ‘The nearest to it 
is Vyctipao gemmans Guén. It differs from all the forms of 
Nyctipao which I have seen in having the apex of the fore wing 
acute. The form crepuscularis Linn. is nearest to it in this respect, 
but its outer margin is highly crenulate, whereas in this species 
it is evenly rounded. From gemmans in particular it ‘differs in 
the absence of the white postmedial line, which is represented by 
a short ochreous striga below the costa. 

Habitat. Hills E. of Toungho, Burma, May 1896, 13,1 @. 
Expanse 74 mm. 

The above specimens were taken by some native Lepcha col- 
lectors sent to the Burma hill-country to collect Lepidoptera by 
Mr. F. Moller, Mr. J. Apcar, and myself, 


Proc. Zoot. Soc.—1917, No. XVIII. 18 


250 ON MOTHS FROM B, E. AFRICA. 


EXPLANATION OF THE PLATE. 


Fig. Fig. 
1. Ancylolomia gracilis, 3. 17. Cortyta minyas, 6. 
2. Ancylolomia chrysolinealis, 8. 18. Cortyta griseacea, 3. 
3. Surattha rufistrigalis, g. 19. Rhodesana mintha, 6. 
4. Thosea perseis, g. 20. Leocyma candace, 6. 
5. Thosea chloris, 6. 21. Pteronycta cervicornis, g. 
6. Thosea undosa, &. 22. Parastichtis orylus, &. 
7. Euproctis rubricosta, &. 23. Plecoptera polymnia, ¢ . 
8. Lacipa albula, g. 24. Brevipecten icarus, & 
9. Rhodoneura bryaxis, &.' 25. Parastichtis lysis, 6. 
10. Syntomis polyxo, 8. 26. Nyctipao acuta, 6. 
11. Bithra aganice, 2. 27. Duomitus pindarus, 6. 
12. Cladocerotis ceneus, 8. 23. Anthela nigristigma, 6. 
13. Cynisca thisbe, 9. 29. Hylemera lepta, 3. 
14. Giaura arethusa, 8. 30. Heterocrita metis, 2. 
15. Odontestra avitta, @. 31. Clenusa rectilinea, g. 


16. Heteronygmia aurelia, o. 


DEFORMITY OF OS PENIS IN A SEAL. 251. 


16. Deformity of Os penis in a Seal (Phoca caspica. Nilsson). 
By Seretus ALPHERAKY *, 


[ Received October 5, 1917: Read November 20, 1917. ] 
(Text-figure 1.) 


My friend Mr. T. N. Arnold, our well-known ichthyologist, 
has kindly lent me for description an abnormal os penis of an 
adult Phoca caspica he himself shot a few years ago on the 
Caspian Sea. 

This abnormality is evidently due to an accidental fracture at 
about the first third part of the total length of the bone. ‘That 
the. bone has here been broken in some way, and has again 
united with the portions in a distorted position, can clearly be 
seen from the accompanying drawing, made by my son Dmitry 
Alpnéraky. The drawing shows the bone from above, and as it 
is very exact in form and size, I think it gives a better idea 
of the malformation than could be given by any amount of 
description. 


Text-figure 1. 


The fractured part is so strongly and solidly united with the 
main part of the bone, that the fracture must have taken place a 
long time prior to the animal’s death. 

It is dificult to find a satisfactory explanation of the cause of 
such a fracture, and Mr. Arnold thinks that it may have been 
the sequel of some violent movement of one of the animals during 
the act of copulation. 

As far as Mr. Arnold remembers, the penis outwardly had no 
signs, or scars, to show that anything was amiss inside, and 
the deformity was discovered only during the cleaning of the 
bone. This was not an easy thing to do, as the apical part of the 
bone hal the cavernous integument round it strongly adherent. 
This apical part of the bone has now a very rough, porous, 
uneven surface, showing that the erectile tissue had been deeply 
embedded. 


* Communicated by Mr. O. Tuomas, F.R.S, 


oh 


CONCEALMENT BY THE APPARENT DISRUPTION OF SURFACE. 253 


17. Some Observations upon Concealment by the apparent 
Disruption of Surface in a Plane at Right Angles to 
the Surface. By J. C. Morrram, M.B. (Lond.) *. 


[Received October 22, 1917: Read November 6, 1917. ] 


(Text-figures 1-4.) 


INDEX., 
MorPHOLOGY: Page 
FAMALY SIS OF PALLETS iiss sessacnstsronssirrncncaiseraecsecess 920M 
ETHOLOGY : 
Concealinge power of patterns. (1s. ..cvises drecsncvscescnsnses «200 
2 


The forms of animals are often rendered inconspicuous by the 
methods which the artist usually employs in the representation 
of three dimensions on a flat surface. “ 

Attempt is here made to show, first, what use the artist 
makes of Simultaneous Contrast for the apparent disruption of 
the surface of his paper or canvas: and subsequently to ascertain 
whether similar arrangements occur in the patterns of animals. 

The Artist's Use of Simultaneous Contrast.—When a dark 
object is viewed against a light background or vice versa, it will 
be seen that where the dark and light areas join, the dark tone 
appears darker than the rest of the dark area, and the light, 
lighter than the remainder of the light area. 

The same appearance will be seen on the artist’s paper if 
he copies exactly the tones of the original. If, however, he 
exaggerates the contrast by surrounding dark objects with dark 
lines and light objects with light, then the objects will appear to 
stand out from the background, and different portions of his 
paper to occupy different planes. For this purpose, foreground 
figures are often surrounded with dark lines, clouds are edged 
with light tone, and trees against the sky are outlined with black, 
whilst the neighbouring sky is greatly heightened in tone. 

It is not uncommon to find a series of receding hills outlined 
with dark lines of gradually decreasing intensity: it is remark- 
able how realistically distance is thus represented, 

The introduction of dark tone between coloured areas results 
in disruption of surface for another reason. On viewing the 
junction of two coloured areas, there will be seen for a short 
distance a flowing-over of each colour into its neighbour. This 
is well seen by drawing a narrow white line across red and blue 
surfaces: where the white lies on the red, it appears reddish, 
where on the blue, ight blue. If the white line be surrounded 
on both sides with black lines, then this flowing-over is prevented 
and the white line retains its purity. By thus preventing this 


* Communicated by the SECRETARY. 


254 DR. J. C. MOTTRAM ON CONCEALMENT BY 


blending, coloured areas in juxtaposition have the appearance of 
standing out, one from the other, and occupying different planes. 

These appearances may be explained in another manner. ‘The 
mind largely recognises objects by their outline, and to some 
extent judges their distance by its sharpness. If in representing 
objects on a flat surface, they be edged with white or black lines, 
then the sharp outline thus given will make them appear nearer 
the eye than their surroundings, and the flat surface appear to be 
disrupted in a plane at right angles to the surface. 

Surface Disruption in Animal Pattern.—Concealment in the 
animals consists chiefly in the hiding of outline, solidity, and 
surface. Outline is largely concealed by ‘“ Disruptive Coloration,” 
solidity by ‘‘ Counter Shading.” 

‘Disruptive Coloration” conceals outline in the following 
manner: when an animal covered with a number of large 
patches of different tone and colour comes to rest on a back- 
ground which harmonises with any one of the components, then 
its characteristic outline will be broken, as shown in text-fig. 1. 


Text-figure ie 


Disruptive Coloration: A, a pattern of three components which largely interrupt 
the margin. B, the appearance when seen against a background harmo- 
nising with the spotted component, C and D when seen against a striped 
and a plane background. 


The disruption is in the same plane as the wings of the insect, 
and is dependent upon a pattern breaking out over a large portion 
of the margin. 

Disruption of surface consists in an arrangement of pattern 
and coloration, which causes some portion of the surface to 
appear close to the eye, and others far away. In this case the 
disruption is in a plane at right angles to the plane of the object. 
As has already been shown, this can be produced by the use of 
Simultaneous Contrast and, thus brought about, may be termed 
Surface Disruption -by Contrast. ‘Outline Disruption” and 
“Surface Disruption by Contrast” are often combined as shown 
in text-fig. 2A (compare text-fig. 1). Text-fig. 2B illustrates 
an insect showing only Surface Disruption. 

An examination of RBiitish Lepidoptera is now made in order 
to estimate to what extent this method is employed for purposes 


THE APPARENT DISRUPTION OF SURFACE. 955 


of concealment. It may be said with confidence, that the whole 
of the Geometers in respect of the dorsal surface of their wings, of 
the Noctuas with regard to the dorsal surface of their anterior 
wings, and of the Rhopalocera in respect of the ventral surface of 
their posterior wings, present concealing pattern and coloration, 
Here, therefore, ‘‘Surface Disruption by Contrast” may be 
looked for. 


Text-figure 2. 


A B 


A, a pattern similar to that shown in text-fig. 1, but with the addition of Surface 
Disruption produced by light and dark lines between the components ; 
in C the surface is further disrupted by the representation of solidity. 
B shows “Surface Disruption by Contrast ’’ not associated with disruption 
of outline. This is rarely found: Thyatira batis is a good example. 
D shows the combination of Surface Disruption by Contrast with Surface 
Disruption by Solidity. Large ocelli generally appear to be of this nature. 


Text-figure 3. 


5 


Shows in diagram seven types of pattern commonly found in Geometers and other 
Lepidoptera, in which Surface Disruption by Contrast is combined with 
Outline Disruption. Many variations occur of type no. 6; in some cases the 
bands are wavy or zigzag, more especially in the fore wings of Noctuide. 


An examination of the Geometridee (Newman) shows that 
surface disruption occurs, associated with well-marked types of 
outline disruption. ‘These are illustrated diagrammatically in 
text-fig. 3, and are seen to consist of well-defined bars of tone 
interrupting the margin and which are separated from the rest of 


256 DR. J. C. MOTTRAM ON CONCEALMENT BY 


the insect’s wing by distinct black or white lines (only black lines 
are shown in the diagrams). 

The distribution of these types is given in the Table on p. 257, 
where it can be seen that “Surface Disruption by Contrast” occurs 
in the Geometers in 69 out of 276 insects. In the Noctuas the 
patterns occur 34 times in 311 insects, in the Cuspidates 11 in 33, 
and in the Rhopalocera 10 in 60. 

It follows that this method is often used for purposes of con- 
cealment amongst the Lepidoptera ; numerous examples are also 
to be found in the patterns of Vertebrates. 

Surface disruption can also be produced by the representation 
of solid objects on a flat surface: this, indeéd, is the method 
chiefly employed by the artist. This method is very often found 
in the pattern of animals generally combined with “Outline 
Disruption ” and “Surface Disruption by Contrast.” Examples 


Text-figure 4. 


The Oleander Hawk-Moth (Cherocampa nerii). From ‘The Natural History of 
British Butterflies and Moths,’ by Edward Newman, F.L.S., F.Z.S. See text 
below : reference is made only to the fore wings; the hind wings are more or 
less concealed by the fore wings in the position of rest. 


ave shown in text-fig. 2, C and D; in C all three methods are 
combined, in D surface disruption by solidity with contrast. No 
better example of snch combinations of methods can be found 
than Cherocampa nerw illustrated in text-fig. 4; 1u this insect 
the components of the pattern break out at the margin, are shaded 
to represent excrescences and depressions, and are separated from 
each other by well-defined white and black lines. 

Opportunity is here taken to call attention to the fact that 
many small details of pattern are of value in concealment; it 
would seem dangerous therefore to conclude on negative evidence, 
that small differences such as often distinguish species can have 
no value in the struggle for existence. 


THE APPARENT DISRUPTION OF SURFACE. 


TABLE.* 
pees Geometers. | Noctuas. | Cuspidates. Rhopalocera. 
(eee 3 i | 3 
Die Cec, 1 me 2 4 
3 ae ae ee 4, 2 1 ar 
AR tetas 10 8 3 3 
OPE, Pasar 25 a 4 
nee 21 15 
ieee 5 2 
| eee as itt! Prats! | 
Total 69 34 | 11 | 10 
SO 311 33 60 


: | 
species. J 


\ 


a 
| 


| 
| 


* Compiled from Newman, ‘The Natural History of British Butterflies and 


oo 


P.Z.S. 1917. TALBOT, & [OICEYeRlaae 


Horace Knight, del. MENPES PRESS, WATFORD. 


LEPIDOPTEROUS ABERRATIONS, 
NEW SOUTH AMERICAN ARCTIIDE. 


ON NEW SOUTH-AMERICAN RHOPALOCERA. 259 


18. New South-American Rhopalocera. By J. J. Joicry, 
F.L.S., F.Z.8., f.H.S., and G. Tatsot, F.1.S8, 


[Received June 20, 1917: Read November 20, 1917. | 


(Plate 1.*) 
INDEX. 

SYSTEMATIC : Page 
Papilio erlaces chinchipensis, subsp. Nn. ....6..+...... 259 
Papilio iphidamas ayabacensis, subsp. n. eee ee 3), 
Papilio harmodius tabaconas, subsp. n. ... ........... 260 
Leodonta zenobia monticola, subsp. n. .................. 260 
Catasticta philone nimbata, subsp. n. .................. 260 
Catasticta flisa tamboonsis, SUDSPe Ms. ( hcteeqar cates 200 
Catasticta lathyi,sp.n. ...... etic 2 Ok 
Catasticta nimbice peruviana, subsp. Mee hago Ok 
Vila eueidiformis, sp. n. ....... Hess ll 
Vila eueidiformis ab. radiata, new ‘aberration .. ...... 262 
Polygrapha cyanea G. &S., Qo... ccccce cece tees ee ees 268 

eee 

“ila eueidiformis and lewis CONIG CS a vieahce akon on 263 


The types of the forms herein described are in the collection 
of Joicey. 
PAPILIONIDAE, 


PAPILIO ERLACES CHINCHIPENSIS, subsp. n. 


3. Upperside like typical erlaces Gray. Underside with spots 
as in lacydes Hew. 

2. Resembles lacydes but with no post-discocellular spots. 

This race connects erlaces erlaces with lacydes through 
canthias R. & J. 

Hab. North Serer tae River, Sept., li did, 62°; 
Charape, 4000 feet, 29 9; Tabaconas, 6000 feet, 2¢ ¢. These 
places are on the eastern slopes of the Andes. Collected by 
A, t,, & H Pratt. 


PAPILIO IPHIDAMAS AYABACENSIS, Subsp. n. 


Nearest to the race calogyna Roths. & Jord., from West Ecuador 
and Colombia. 

3. Upperside with reduced green patch on the fore wing. 
Hind wing with three red spots, all separate. 

Underside of hind wing with the spots smaller and more 
widely separate; ground-colour not so black and browner than in 
ealogynda. 

2 similar to calogyna 2, but band of hind wing much whiter 
and with an extra spot in cellule 6, similar to a specimen in the 
Tring Museum from Zaruma, S.W. Ecuador, which has the band 
of the hind wing half white. 

Hab. North Peru-—Ayabaca Mtns., 6 od, 2292; River 
Tabaconas, 6000 feet, 1g. Collected by A. E., & F. Pratt. 


* For explanation of the Plate see p. 276. 


260 MESSRS. J. J. JOICEY AND G. TALBOT ON 


PAPILIO HARMODIUS TABACONAS, subsp, n. 


Near the race weniades Hew., from Ecuador. 

3 like harmodius Doubl., but with a narrower white patch on 
the fore wing, straighter on its proximal edge. 

2 with no constant difference. 

A og in the Tring Museum from Zamora, -S.W. Ecuador, 
occurring as an aberration with typical weniades. 

The 2 presents a strong resemblance to the Q of P. erlaces 
chinchipensis found in the same. district. 

Hab, North Peru—River Tabaconas, 6000 feet, 12 ¢ ¢6, 19 
(types); Charape, 4000 feet, 27 g ¢; Ayabaca Mtns., 1 9 ; 
Rentema Falls, Upper Maranon, 1000 feet, 1g. Collected by 
A. E., & F. Pratt. . | 


PTERT Da. 


LEODONTA ZENOBIA MONTICOLA, subsp. n. 


3g. A larger race with deeper yellow markings. 

Upperside of fore wing with the two post-discocellular spots 
-farther from the cell, there being a wider space between the 
Jower spot and the one in 3; the subterminal spot in 3 is 
larger. 

Underside with a larger cell-patch, which extends basad beyond 
vein 2; patch below vein 2 slightly more extended basad. Hind 
wing with no constant difference. 

Length of fore wing 80-34 mm.; in zenobia zenobia it averages 
29 mm. 

Hab. Uruhuasi, §8.E. Peru, 7000 feet, April to May, 1910. 
H.& C. Watkins. 5 3 ¢. 


CATASTICTA PHILONE NIMBATA, Subsp. n. 


3. Upperside very similar to philone Feld. from Colombia, 
but basal markings less well-defined. 

- Underside of fore wing with the three anterior spots of the 
Aieal band smaller. Mind wing with the discal white band 
much narrower in the cell and below it; the yellow spots deeper 
in colour. 

This race is not well differentiated. 

Hab. Peru, 1 @. 

This and the following forms of Catasticta were received from 
Monsieur E. Le Moult of Paris. 


CATASTICTA FLISA TAMBOENSIS, subsp. n. 


3. Upperside more brown than black. Hind wing with the 
band narrowed in the cell and postdiscal spots minute. 

Underside of fore wing with cell-spot larger; spots of post- 
discal row larger and nearer the margin, distally concave, form- 
ing lunules. Hind wing with precostal area white, discal band 
widened costally ; outer subterminal area more washed with 


NEW SOUTH-AMERICAN RHOPALOCERA. 261 


white; marginal spots yellow, large, and triangular, their apices 
almost touching the postdiscal spots. 

Hab. Peru, 1 ¢. 

This form is probably the same as the specimen recorded as 
flisa H.-Sch. by Weymer & Maasen, Lepidoptera, p. 73 (1890), 
from Tambo Almirante. We do not know another record of this 
species from Peru. 


CaTASTICTA LATHYI, sp. n. 

Allied to philais Feld., from Colombia. 

g. Colour above nearly as in manco Doubl. and pattern 
similar, but easily distinguished by the brown margin of the 
hind wing and different underside. Differs from philais above 
in colour, philais being brown; the pattern is the same, but the 
dark margin of the hind wing is broader. 

Underside of fore wing with reducel cell-streak; marginal 
spots smaller but more produced proximally, forming streaks ; 
the spots of the postdiscal series are smaller. These markings, 
which are brown in philats, are here grey-white. Hind wing 
with markings very much as in philais, but the yellow intra- 
neural distal streaks longer. 

Length of fore wing 26 mm. 

Hab, Loja, Ecuador, 1 g. Also in the Tring Museum from 
the following places: Loja; Zamora, Ecuador, 3000-4000 feet ; 
Cushi, Huanuco, Cent. Peru, 1820 m. 


CATASTICTA NIMBICE PERUVIANA, subsp. n. 


3. Distinguished by the more rounded wings and _ paler 
markings. ‘Lhe discal spot in 4 on the fore wing is much 
smaller than the spot above it; spots in 7 and 8 absent ; post- 
discal spot in 3 shifted inwards; no marginal dots. Hind wing 
with the pale band widened and its outer edge nearer the post- 
discal spots; spot in 3 shifted inwards; spots in 2 and 4 farther 
from the band than the others. 

Underside of fore wing with the band, cell-spot, and lower five 
postdiscal spots pale straw-colour. Hind wing with the discal 
band reduced distally, the spot at base of 2 being smaller, and 
widened proximally; postdiscal and marginal spots larger. ‘The 
markings which are orange-yellow in numbice are bright yellow 
In perwviand. 


Hab. Peru, 1d, 


NYMPHALIDA - 


A remarkable new species of the Nymphaline genus Vila. 


VILA EUEIDIFORMIS, sp. n. (PI. I. figs. 1, 1a.) 

3. Upperside black-brown. Fore wing with pale yellow mark- 
ings. A basal wedge-shaped streak in lower part of cell; a 
wedge-shaped spot in end of cell, placed transversely, its pointed 


2O2, MESSRS. J. J. JOICEY AND G. TALROT ON 


end posteriorly ; a large, somewhat quadrate spot beyond end of 
cell divided by vein 5, the lower part in 4 larger than the upper, 
the inner edge slightly curved ; a small spot above it in the angle 
of 6; an elongate oval spot below it at base of cellule 3. Hind 
wing paler in the costal area. A black patch of androconia in 
base of cellule 7, 9 mm. long. Red basal streaks in la, 6, c, and 
in the cell. In a second specimen, from Bolivia, they are a little 
reduced, but in a third, from the Upper Amazon, the red streaks 
are greatly enlarged and more extended, the one in the cell filling 
its base and whole upper part, forked at the end and forming two 
short streaks in 4 and 5; there are also two additional short 
streaks in 2 and 3, 

Underside with paler ground-colour. Fore wing with markings 
as above, and a white dot at base of costa. The black patch of 
androconia, common to all members of the genus, is longer than 
in other species and does not touch the median vein; it is placed 
obliquely in the submedian area, whereas in other species it 1s 
parallel to vein 2; the inner margin is shining grey; a white 
subterminal line from tornus to vein 2 and represented above 
this vein by a few scales. In the species with increased red on 
hind wing above, the basal fore-wing stripe is only represented 
by a few scales. Hind wing with brick-red stripes between the 
veins; base of costa and precostal area brick-red, joined to a 
stripe bordering vein 8 in 7; stripes in lc, 2-6, nail-headed 
distally, the one in 4 joined to a broader cell-stripe, stripe in la 
filling the cellule; a subterminal seriés of white spots between 
the veins. In the specimen with increased red on hind wing 
above, the stripes are broader below. 

Antenne black-brown, club and inner surface yellow-brown ; 
palpi black, first and second segments white at sides; head, 
thorax, and abdomen black, collar with white dots; abdomen 
below dark brown, legs dark brown, coxe with white tufts. 

Length of fore wing 35 mm. 

Hab, Jurua River, Brazil (type and paratype); Bolivia (1d). 
This latter locality is doubtful. 

We propose the name radiata for the aberration with increased 
red rays on the hind wing above (PI. I. fig. 1a). . 

This species agrees in structure with other species of the genus, 
possessing the characteristic inflated basal part of the costal vein 
of the fore wing, and also the curved lower discocellular touching 
the cell before vein 3. In pattern, its relationship with other 
species may perhaps be traced in the fore-wing cell-streak, the 
white mark at base of costa on fore wing below, the white sub- 
terminal mark near tornus, the red subcostal stripe on hind wing 
below, and the series of white spots which may represent the 
subterminal line. The colour of the palpi and antenne also 
shows affinity with other species. 

The chief feature of this species lies in its remarkable mimetic 
resemblance to the Heliconine genus Hueides as represented by 
E. eanides Stich. and its form aides Stich., which inhabit the same 


NEW SOUTH-AMERICAN RHOPALOCERA. 263 


region. The form eanides is mimicked by the specimen with 
increased red rays on the hind wing above. The hind-wing 
underside, with its rayed pattern and marginal row of white 
spots, shows the greatest resemblance. It also bears a strong 
likeness to Pericopis mimica Feld., a day-flying Hypsid moth, 
which occurs from Colombia to Peru. In a lesser degree, the 
species also resembles two forms of the Nymphaline genus Hresia, 
represented by the darker specimens of 4. perilla Hew. and 
cornelia Stgr. The diversity-of co-existent mimetic forms, seen 
in the species of Hresia, is analogous to the wide departure from 
the normal form now seen to occur in the genus Vila. 

The typical forms of Vila possess a somewhat Ithomiine-like 
pattern, but more nearly resemble some white-banded forms of 
Eresia, the likeness being marked on the underside of the hind 
wing. Ithomiines would probably serve as models for Vila on 
the wing, and in such a white-banded association would be 
included Adelpha epione Godt. Another Adelpha, coryneta Hew., 
resembles Vila azeca D. & H. on the hind wing below, as also 
does Hresia clio L. 

We have, therefore, a mimetic association of Vila eweidiformis 
with Hueides, Hresia, and Pericopis, and another association of 
Vila azeca with Ithomiine species and the Nymphalines Adelpha 
and Hresia. It would appear probable that the resemblance of 
Vila to Ithomiines and Adelpha would be most marked when on 
the wing, whilst presumably the resemblance on the hind wiug 
below between Vila azeca, Hresia clio, and Adelpha coryneta 
would be sufficiently marked when at rest. 

No doubt the white-banded Ithomiines, probably of the genus 
Leucothyris, have served as models, as they are more abundant 
than the Nymphalines. Adelpha is fairly common, and so are the 
white Hresia, but Vila is comparatively rare. 


The Female of PoLYGRAPHA CYANEA G. & 8. 
(PI, 1. fig. 2.) 


The male of Polygrapha cyanea Godm. & Salv. is not rare in 
collections, but no description of the female seems to have been 
published. We describe it from the only specimen which we 
believe to exist in collections. This specimen, which is much _ 
damaged, was taken by Mr. W. F. H. Rosenberg’s collector, 
Mr. G. Palmer, at Alpayacu, Rio Pastaza, E. Ecuador, 3000 feet, 
and was found drowned in a native “ Chicha” pot outside a 
house. The collection containing the specimen was acquired by 
the late Mr. Herbert Druce. 

Upperside with dark brown ground-colour. Fore wing with 
yellow-brown oblique band from vein 4 to the inner margin near 
tornus; the band is much broader in cellules 2 and 3 and its 
inner edge is incurved in cellule 2 and in 10, the outer edge 
is evenly curved outwardly ; a pale brownish streak at base of 
cellule 6, and beyond this the costal area whitish brown, not 


é 


264 ON NEW SOUTH-AMERICAN RHOPALOCERA. 


extending to the base nor distally beyond the streak in 6; a small 
triangular brown spot in 5 in the distal area. Hind wing with a 
yellow-brown costa] edge and apical suffusion ; a-series of yellow- 
brown subterminal spots, well-defined proximally, but distally 
suffused ; outer margin narrowly edged with yellow-brown. 
Underside marked as in the 3, but the specimen being rubbed 
we cannot distinguish any differences in colour. 

Head and antenne missing. ‘Thorax and abdomen black-brown. 

Length of fore wing 41 mm. 

The completely different pattern presented by the female of 
this insect suggests some sort of mimetic resemblance. It pre- 
sents a great similarity to the pattern of many species of the 
Brassolid genus Opsiphanes, four species of which, with brown 
bands on the fore wing, are recorded from Ecuador and Peru. 
Mr. W. J. Kaye informs us that Opsiphanes fly round the houses 
at dusk, and this observation, coupled with the fact that the 
female Polygrapha was found in a water-pot near a house, points 
to the possibility of the two insects flying in company. 


> 


ON NEW SOUTH-AMERICAN ARCTIIDA. 265 


19. New South-American Arctiide. By J. J. Joicey, 
ELS: 1:Z.S,, FHS... and G. Tarsor, F.E.S. 


[Received June 20, 1917: Read November 20, 1917. | 


(Plate I.*) 


INDEx. 
SYSTEMATIC: Page 
RIC AUUSIAVELNOFAD, SY. /Is bers0scesdevserenesias ss sesgsesds ate, 200 
Neonerita metapheenica, sp. nN. ......... cee cece eee eee ees 265 
Areéomolis nigripuncta, SP. Me... .cccce cece cece teens eee 266 
PARMLOMOLIS AGNIVENA, SPs Ne .aseie..-bep-2cncvececseos seems 200 
Automolis ochreogaster, SP. De ....0.0.c.ccbeeveceeeeesene ess 267 
Automolis ochreomarginata, Sp. ...... ce. ce cece 267 
Glaucostola maroniensis, Sp. Ne 2... ..c cece cee eee. 268 
UMICLCSeIAVESCENS"SP)sNe 5x2 rede. tcads xe oreoreradeicien 9208 
OphOTUS OCHRACEA, SPeMe res cteneie: cemnes tesco vex 209 
Neritos purpureotincta, Sp. Ve... .....0cc ccc ceccee eee 269 


The types of the species herein described are in the collection 
of Joicey. Our thanks are due to Sir G. F. Hampson for kind 
help given. 


1. IDALUS FLAVITHORAX, sp. n. (PI. I. fig. 4.) 


Nearest flavoplaga Schaus. 
6. Both sides of fore wing pale purplish brown ; veins white. 
A large pale-yellow distal costal patch, extending to below vein 4 
and filling end of cell, its outer edge parallel to margin and as 
long as its costal edge. Inner margin with a yellow stripe on its 
outer half, the edge of the stripe curved above the submedian. 
Hind wing pale yellowish white, costa and margins more deeply 
tinted. 

Head, thorax, and abdomen ochreous-yellow, three end seg- 
ments of abdomen above black fringed with grey, below yellow 
excepting end-segment. Antenne white above, brown below. 
Palpi greyish-white. Legs white, coxe of fore-legs yellow. 
Length of fore wing 15 mm. 

Hab. 8. Brazil, Rio Grande do Sul. A single specimen. 


2, NEONERITA METAPHENICA, sp.n. (PI. I. fig. 5.) 


Nearest to yahuase J.& T., Ann. & Mag. N. H. ser. 8, vol. xviii. 
p. 54, pl. xiv. fig. 3 (1916) (Peruvian Amazon). 


3. Differs from yahuase in its smaller size, broader band on 
the fore wing, and hind wing washed with crimson except over 
the apical area. Antenne, thorax, and abdomen above crimson, 
the thorax being much duller with a greyish admixture. Length 
of fore wing 9 mm. 

Hab. French Guiana, St. Jean du Maroni, 1 ¢. 


* For explanation of the Plate see p. 276. 
Proc. Zoou. Soc.—1917, No. XIX. 19 


266 MESSRS. J. J. JOICEY AND G. TALBOT: ON 


3. ARHOMOLIS NIGRIPUNCTA, sp. n. (PI. I. fig. 6.) 


Near sanguinea Hamps.*, but more strongly irrorated with 
yellow, the discal line strongly curved, and the three subapical 
spots placed in a curve. 

3. Upperside of fore wing crimson, strongly irrorated with 
yellow. Costal edge greyish white for proximal two-thirds, 
Markings yellowish grey tinged with crimson. Three ill-defined 
basal spots edged with blackish outwardly, the upper one in the 
cell; a discal band of spots, its Inner edge strongly curved and 
defined by blackish dots, its outer edge irregular, though the 
upper part defined by blackish dots forms a curve to below 
vein 3; below this the band is much narrower and reaches the 
tornus ; anteriorly it is limited by the subcostal and is composed 
of a double spot in the end of the cell, three outside the cell and 
five below it; three subapical spots forming a curve, and below 
the third two smaller and darker spots parallel to the margin ; 
the termen marked by black dots at the ends of the veins. 
Hind wing with distal area salmon-pink, the discal and basal 
area being hyaline. 

Underside of fore wing with basal area to vein 2 grey tinged 
with crimson, costal edge grey, distal area crimson strongly tinged 
with yellow, markings of upperside faintly showing. Hind wing 
paler than above. 

Antenne grey-brown except basal two-thirds of shaft, which is 
crimson above; palpi crimson inside and grey-brown on outer 
side; frons grey-brown; head and thorax crimson irrorated with 
yellow ; abdomen above salmon-pink, basal two segments and 
anal tuft grey-white; thorax and abdomen below grey-white ; 
legs grey-white marked with crimson, anterior coxze crimson on 
inside, Length of fore wing 16 mm. 

Hab. French Guiana, St. Laurent, Maroni River, July—Sept. 
J ui raueh tole 

Whilst Sir George Hampson regards this as an Arcomolis, it 
may be useful to point out that it differs from his original 
diagnosis of that genus. The fore wing has vein 10 originating 
before 7, and the hind wing has 3 and 5 on a short stalk, and 6 
and 7 from the cell. The fore wing has a patch of androconia 
below the cell in the median area. The hind wing has the costa 
lobed, and bears a small patch of androconia on the upperside 
above the cell; on the underside of the inner margin is a fold 
containing a brush of androconial hairs which are club-shaped. 
The antenne are serrate and fasciculate. 


4, AUTOMOLIS IGNIVENA, sp. n. (PI. I. fig. 7.) 


Near ditissima Walk. from 8. Brazil, but larger and with a 


difterently shaped apical patch. 
2. Upperside of fore wing fuscous-brown irrorated with 


* Areomolis sanguinea Hampson, A. M. N. H. ser. 7, vol. xv. p. 442 (1905) 
Cayenne). 


NEW SOUTH-AMERICAN ARCTIIDA. © 267 


blackish, darker on the inner margin, basal area paler; apical 
patch pale yellow, extending to vein 3, outwardly defined by pale 
orange, which also colours the veins traversing it; the part filling 
cellule 4 is half the length of the areas in 3 and 5; a discal zigzag 
crimson line from the costa at vein 6 to the middle of inner 
margin, outwardly curved to vein 3, then directed basad to sub- 
median, thence distad to margin on the black area, being here 
much thickened and enclosing a yellow spot; anteriorly the teeth 
formed in cellules 3-5 are filled in with pale orange; some 
reddish suffusion at the base; a short crimson basal streak below 
cell; costa pale orange; veins crimson. Hind wing dark brown, 
also fringes; apex produced and a short anal lobe. 

Underside of fore wing dark brown, apical patch as above ; 
basal two-thirds of cell and a stripe below it pale yellow. Hind 
wing with a pale-yellow subbasal spot below costa. 

Antenne brown, white anteriorly; palpi with first segment 
white, the rest grey on outside, black on inside; frons grey-white; 
head pale orange; thorax yellowish white marked with orange, 
two black dots at base; abdomen crimson above, white below; 
pectus white; legs white, fore-legs marked with pale orange. 
Length of fore wing 18 mm. 

Hab, North Peru, Charape River, Tabaconas, 4000 feet: A. E. 
Prawb 1912: 12. 


5. AUTOMOLIS OCHREOGASTER, sp.n. (PI. I. fig. 8.) 

¢. Upperside silky-white; underside similar with the veins 
brownish. ¢ 

Head, tegule, patagia, and base of abdomen silky-white ; 
antennee with shaft white above, brown below, serrate and fasci- 
culate; frons with a tinge of ochreous; an ochreous line at side 
of eye; abdomen pale ochreous; legs grey-white; palpi white, 
ochreous on inside. 

Hind wing with 3 and 5 stalked, 6 and 7 stalked, 8 from 
middle of cell. 

Length of fore wing 17 mm. — 

flab. Sapucay, Paraguay, July 1902, 1°. 

Similar to albescens Roths., but differs in neuration of hind 
wing. 


6, AUTOMOLIS OCHREOMARGINATA, sp. n. (PI. I. fig. 9.) 


Nearest sicilia Druce, from Mexico and Panama. 

d. Upperside.—Vore wing orange-yellow, the apex thinly 
scaled with black. A narrow terminal buff margin widening 
posteriorly. Hind wing paler yellow than fore wing, darker 
along outer and inner margins. . 

Underside as above.—Vore wing with apex and costal edge 
narrowly black to within a quarter from base. Hind wing 
deeper yellow on costal area. . 

13% 


268 MESSRS, J. J. JOICEY AND G. TALBOT ON 


Head and thorax orange-yellow; frons metallic blue, anteriorly 
ochreous; vertex with a metallic-blue spot: palpi black, ochreous 
on the outside; antenne black, serrate and fasciculate ; ; legs 
ochreous striped with black; abdomen black above, first and 
second segments orange-yellow, segments 4-6 with a metallic-blue 
dorsal spot, a lateral and sublateral series of metallic-blue spots 
reaching terminal segment, ventral surface ochreous with narrow 
black bands. Length of fore wing 16 mm. 

Hab. French Gian St. Jean du Maroni, 1 ¢. 


7. GLAUCOSTOLA MARONIENSIS, sp.n. (PI. I. fig. 10.) 


3. Upperside.—fore wing blackish brown, greyish between 
the veins; two large irregularly-rounded white spots, one sub- 
basal, the other subapical; inner margin at the base narrowly 
orange. Hind wing pale orange with black outer margin, which 
is widest at angle of the wing and narrows posteriorly. 

Underside.—- ore wing with basal area white and fovea bearing 
a tuft of white hair. Hind wing as above. 

Head orange, a black spot on vertex, frons black; palpi black, 
paler on outside; antenne black, serrate and fasciculate ; tegule 
orange with mesial black spot; thorax grey-black; abdomen 
orange, paler below, segments 5 and 6 black dorsally and later- 
ally, a lateral row of black spots; legs grey-black and ochreous. 
Length of fore wing 13 mm. 

Hab. French Guiana, St. Jean du Maroni, 1 3. 

Differs from typical Glaucostola in antenne being serrate aud 
fasciculate, and in fore wing having vein 3 from before angle of 
cell. Near metaxantha Schs. 


~ 


8. MELESE FLAVESCENS, sp. n. (PI. I. fig. 11.) 


This distinct species belongs to Sect. 11. of Hampson’s diagnosis, 
and the antenne are serrate and fasciculate. 

3. Upperside of fore wing fuscous-brown irrorated with 
yellow. A yellow dot at the base, a yellow spot below the celi on 
the submedian, another boyoud it touching submedian and vein 2, 
one at the base of cellule 2, a rounded yellow spot beyond the 
cell in 5. Hind wing hyaline, costal and inner margins pale 
yellow, veins yellowish, fringes yellowish white. 

Underside of fore wing paler than above, grey-white in the » 
basal area. 

Antenne brown; palpi and lower part of frons pale orange- 
yellow, upper part of frons fuscous, vertex yellow ; tegule fuscous, 
rest of thorax yellow mixed with fuscous; abdomen yellowish 
brown, fringed at base with yellowish-white hair; pectus and 
fore-coxe pale orange-yellow, femora and tibize fuscous, tarsi 
grey-white. Length of fore wing 17 mm. 

Hab. Argentine, September, 1 3. 


bo 
(or) 
eo) 


NEW SOUTH-AMERICAN ARCTIID®, 


9, OPHARUS OCHRACEA, sp. n. (PI. I. fig. 12.) 


This distinct species has no near ally. 

©. Fore wing pale ochreous-brown with white markings. A 
small patch at the base with two black dots; a median band, its 
upper part directed distad, its lower part below cell narrower and 
directed basad to Inner margin; a broader discal band broken at 
cellule 2, its upper part formed of an oblong patch at right angles 
to costa, its lower part below vein 2 narrower and parallel with 
median band; a narrow postdiscal band of spots, its upper part 
to vein 4 at right angles to costa and composed of four spots 
joined together, the fourth much smaller, its lower part curved. 
basad to inner margin and composed of four separate spots 
smaller than the upper first three; a subterminal series of 
irregular spots, the first in 7 the second in 6, and nearer the 
margin, the third and fourfh joined in 4 and 5,and below the 
first, the fifth in 3, the sixth and seventh joined in 2 and 1, the 
last being curved and close to tornus. Mind wing semi-hyaline 
with a pale ochreous subterminal band, narrowing posteriorly and 
not reaching inner margin. 

Antenne rufous, basal segment white with a black dot; palpi 
pale ochreous, black at sides; frons pale ochreous, anteriorly with 
a black spot ; vertex white with a black spot; tegule white, bear- 
ing two black spots; patagia white with black spots and fringed 
with ochreous-brown ; abdomen pale orange above, grey-white 
below, with a dorsal and lateral series of black spots; pectus 
grey-white ; legs ochreous-brown, fore-coxe bearing a black spot 
on inside. Length of fore wing 25-27 mm. 

Hab. North Peru, Charape River, Tabaconas, 4000 feet: A. E. 
Pratt. £912. 2 O° 2. 


10. NeRITos PURPUREOTINCTA, sp.n. (PI. I. fig. 13.) 


This distinct species belongs to Sect. 111. of Hampson’s diagnosis, 
and the antenne are serrate and fasciculate. 

3. Upperside of fore wing pale purple, with a darker brownish 
basal area; a crimson dot at extreme base, another at base of 
eell, another, encircled by black, in upper basal part of cell ; 
black line strongly curved outwards, from origin of vein 2 to 
inner margin and bordering a rounded crimson inner spot; 
heyond it a second curved blackish line below vein 2; some 
blackish sealing in outer part of cell; a crimson basal str eak on 
inner margin; a broad postdiscal semihyaline band of pale 
amber-yellow, its inner edge straight and oblique, margined with 
crimson, the distal edge curved inward, also margined with 
crimson, cutting off a rounded apical patch; veins traversing 
apical patch crimson near margin and blackish behind ; costal 
edge crimson except on the band; fringe of outer margin pale 
yellow. Hind wing pale orange-yellow, costa pinkish, 

Underside of fore wing similar to upperside, basal area darker, 


* 
270 ON NEW SOUTH-AMERICAN ARCTIID. 


paler in the median space; crimson edging to the band better 
defined ; apical area darker, more invaded by crimson. Hind 
wing with costa pale crimson. 

Antenne reddish brown, crimson in basal part; palpi crimson, 
second segment marked with brown, first segment white in outer 
and basal part; head and thorax crimson and with a blackish- 
brown patch; abdomen orange-yellow above with a white basal 
dorsal spot; pectus and abdomen below white; legs white, fore- 
and mid-tibie and tarsi pale yellow. Length of fore wing 
13 mm. 


Hab. French Guiana, St. Jean du Maroni, 1 ¢. 


& 
ON NEW BUTTERFLIES FROM AFRICA AND THE EAST. 271 


20. New Butterflies from Africa and the Hast By J. J. 
Jorn | tes.e BP AS.. FLES.,.and G. Tapor, 


[Received June 20, 1917: Read November 20, 1917. ] 


INDEX. 

SYSTEMATIC: Page 
Papilio illyris hamatus, subspe Ny ......s0cececesccreres 201 
Charaxes maudei, sp. n. San eda DEA, 
Charaves smaragdalis or senate. saber Tih see eeee oe ee 
Papilio polydorus ulawaensis, subsp. n. or aberr.n.... 272 


The types of the forms herein described are in the collection 
of Joicey. 


PAPILIO ILLYRIS HAMATUS, Subsp. n. 


So. Upperside with broader bands. Sore wing with band 
straighter on its outer edge. ae wing with a bar at the end 
of cell, almost cutting off a cell-s 

Underside with reduced red on ae hind wing; the red line 
fringing the black spot in 7 is very narrow, and the black spot is 
reduced to a straight bar; white spot beyond cell enlarged and 
entering the cell. 

Hab. German East Africa (British “Cecupaoh )s. ex coll. 
Suffert, 1 ¢. 


Ciranaxns MAUDEI, sp. n. 


Allied to atphares Cram. and citheron Feld., and perhaps 
linking these two species together. 

8 “The upperside resembles, citheron and the underside more 
like wiphares. Much larger than any females we have seen of 
these two species. 

Upperside of fore wing with a broad white discal band, broken 
where 1c, 2, and 3 cross it, the spot in 3 larger than the others, 
in le smaller than the spot below it; 3 subapical spots as in 
citheron, followed by 6 spots in 5, 4, 3, 2, le and 1b, forming a 
subterminal series of fulvous spots, the two upper ones being 
white proximally, the two lower ones placed close to the lower 
median white patches. Mind wing with a broad white discal 
band formed as in citheron, its distal edge with diffuse purplish 
scaling ; a postdiscal series of 5 fulvous spots in cellules 2-6, 
the anterior one touching the discal band; a subterminal series 
of small violet spots; marginal lunules thicker than in the two 
allied species, the anterior 3 fulvous, the posterior 3 dull green. 

Underside with greener ground-colour than in the. allied 
species. More wing with discal band continuous, its proximal 
edge indented so that the black line is strongly curved in the 


* 


272 ON NEW BUTTERFLIES FROM AFRICA AND THE EAST, 


lower median space, and in 2 and 3; black post-discocellular line 
much thicker than in the allied species; subterminal fulvous 
spots much brighter than in citheron, some faint whitish scaling 
distally of the upper 5 spots, which is seen completely developed 
in xiphares; black subterminal spots in 2 and below it as in 
citheron, and not so large as in xiphares. Hind wing with discal 
white scaling from costa to vein 4 more developed than in 
citheron, but much less so than in wiphares; other markings as 
in citheron. 

Length of fore wing 57 mm. 

Hab. German East Africa (British occupation),—Lindi, 1 92, 
ex coll. Suffert. 


CHARAXES SMARAGDALIS ORIENTALIS, subsp. n. 


d. Differs from the type-form in reduced blue on both wings. 
The patch below vein 2 on the fore wing is reduced distally and 
is indistinctly divided by a curved line between vein 2 and the 
submedian, a line which sometimes appears in the typical form. 
The blue band of the hind wing does not extend basad beyond 
the point of origin of veins 3 and 4; ‘it is also reduced distally 
and the spot in 6 is only about two-thirds the typical size; the 
submarginal spots have only the faintest trace of a white dot. 

Underside more yellowish, the yellow subterminal lunules 
thicker and without any white scaling. 

Hab. Kast Africa—Kericho, July 1903 (Jackson), 1 5. In 
Tring Museum 1 ¢ from Rau, Nandi, 12. 2.98 (Dr. Ansorge). 


PAPILIO POLYDORUS ULAWAENSIS, subsp. or aberr. n. 


Near the race polydemon Math. The hind wing with discal 
patches obsolete above and much reduced below. 
Hab. Ulawa Island, North Solomons, 1 2? , ex coll. Grose-Smith. 


- 

+ 
ber 
+ 

. 

7 
= ra 
. 
. 


=m) 


PZ.5. 1917, TALBOT & |@l@zieuiaaNe 


Horace Knight, del. MENPES PRESS, WATFORD. 


GYNANDROMORPH OF PAPILIO LYCOPHRON. HBN. 


A GYNANDROMORPH OF PAPILIO LYCOPHRON, 273 


21. A Gynandromorph of Papilio lycophron Hbn. By 
Jao sorony, f.iis., HZ.5.; f.B.S., and G, Tatsor, 


[Received June 20, 1917: Read November 20, 1917. } 


(Plate II.*) 


This remarkable specimen is a well-marked example of gynan- 
dromorphism, the right hind wing above being normal. It 
belongs to the race phanias Roths. & Jord. 

Both fore wings are asymmetrical and each shows a large spot 
in the end of the cell. The right fore wing has the band inter- 
rupted between the submedian fold and vein 3, two small spots 
being left below vein 2; the outer part of the spot in cellule 3 
remains and the inner portion of it is nebulous, the spot in 4 is 
indented distally, the spot in 5 is only represented by a streak 
above vein 5, the spot in 6 is reduced distally, also the spot in 7. 
The left fore wing shows some traces of the band in the median 
area, and on the inner margin the spot in 4 is reduced as on the 
other wing, the spot in 5 is slightly reduced anteriorly, that in 6 
is represented by a curved proximal line and a distal dot, the one 
in 7 reduced toa proximal dot. The right hind wing is normal 
except for some orange scaling at the lower angle of the yellow 
band. The left hind wing has the band invaded by streaks of 
brown ground-colour, and notably a broad stripe filling the lower 
part of the cell fromthe base to the end; there is some orange 
scaling at the lower angle of the band. The distal area is 
damaged, but there is some green scaling in cellules 3-5; the 
apical spot is much larger than on the other wing. 

The underside of the right fore wing is normal. The left fore 
wing is darkened over the inner and median area and lower half 
of the cell. The right hind wing has the red discal spots larger 
than usual. The left hind wing is more strongly darkened than 
above, leaving a yellow streak in the cell, in 7, and in 6, with 
other minor traces of the band. The red spots are a little larger 
than on the other wing, especially the apical spot. 

The abdomen bears some scattered dark-brown spots. 

This specimen was obtained by Mr.’ A. E. Pratt at the 
Rentema Falls on the Upper Amazon, North Peru, 


* For explanation of the Plate see p. 276. 


is! 


aye 
2 Ts 
: 
2 . 
. 
* 
- 
me * 
7 
S 
rasp 
= © 
> 


se 


Cap 


Pare 


ine) 


ON THREE ABERRATIONS OF LEPIDOPTERA, 275 


22, Three Aberrations of Lepidoptera. By J. J. Jotcry, 
Hit, beZioss, Le tise, and G., LAnBon,, WH.8. 


[Received June 20, 1917; Read November 20, 1917.] 
The specimens described herein are in the collection of Joicey. 


A Melanie Aberration of CATAGRAMMA CYNosuRA Hew. 
(Pl: I. fig.:3.) 


Upperside of fore wing with distal red band much obscured by 
dark scaling and showing faintly; apical spot absent. ind wing 
with red basal patch reduced and limited by vein 6. 

Underside of fore wing with blue subterminal line thickened 
and forming a series of 6 proximally pointed spots; subapical 
yellow streak represented by a thin and shorter line. Yellow 
diseal band narrower and reduced proximally. Hind wing much 
blackened ; yellow costal patch reduced, yellow band encircling 
the disc only represented by a short streak before vein 2, discal 
transverse streak absent, distal series of blue spots nearer the cell 
and smaller. 

The specimen bears the locality ‘‘ Peru.” 


PHILOSAMIA CYNTHIA Drury. 
Aberration. 


This remarkable aberration is a 2 specimen bred in Trinidad, 
B. W. I:, and received from there by Mr. Ernest Swinhoe. 

The ground-colour of the wings is much darker and in the 
distal area is nearly black, excepting the grey marginal border. 
The white bands are very broad. On the fore wing the post- 
discal band is united as the basal bands at vein 4 and the 
submedian, the space between these being white; the distal 
widening of the lower basal band leaves only a small patch of 
ground-colour on the inner margin; similarly the widening 
of the upper basal band and the proximal widening of the post- 
diseal band reduces the discal lunule to a small ovate spot which 
is obsolete on the left wing. 

The bands of the hind wing are united anteriorly so that the 
space between costa and origin of vein 5 is white; the discal 
lunule is very small on the left wing and minute on the right 
wing. The distal normally pink scaling on both wings is replaced 
by grey. 

Abdomen wholly white; head and tegule white. 


PAPILIO NEOPHILUS PARIANUS R. & J. 
Dichromatic Aberration. 
This is an example of erythrism in a ¢. specimen which hag 
the patches on the fore wing salmon-pink instead of green; the 


276 ON THREE ABERRATIONS OF LEPIDOPTERA, 


anterior spot is translucent, and the scales on the underside in 
the area of this spot are also pink. The scales have suffered no 
displacement; those forming the anterior spot are longer and 
narrower than the more ovate ones of the lower patches, and are 
-mostly tridentate, like those of the underside, this being normal. 
Hab. Las Quiguas, Esteban Valley, N. Venezuela, Nov.-March. 


EXPLANATION OF PLATES I. & II, 


PuateE I. 


Fig. 1. Vila eueidiformis. 
aa f. radiata. 

. Polugr apha cyaneaG. & 8S. Q. 
. Catagramma cynosura Hew. Aberration. 
Idalus flavithorax. 
Neonerita metapheenica. 
Areomolis nigripuncta. 
Automolis ignivena. 

a5 ochreogaster. 

=. ochr comarginata, 
10. Glaucostola maroniensis. 
11. Melese flavescens. 
12. Opharus ochracea. 
13. Neritos purpureotincta. 


Die Ses A 


PLATE IT. 


Fig. 1. Gynandromorph of Papilio lycophron Hbn. Upperside. 
2. a 5, Hi a Underside. 


ON ANT-LIKE SPIDERS FROM MALAYA, 277 


23. Ant-like Spiders from Malaya, collected by the 
Annandale-Robinson Expedition, 1901-2, By H. D. 
Bapcock, M.A.*. 


| Received July 7, 1917: Read October 23, 1917. | 


(Text-figures 1-12.) 


INDEX. 
SysTEMATIC. Page Page 
New names :— Descriptions of species previously 
described :-— 

Myrmarachne cornuta ............... 291 aaa 
a COP UORMIUS. acces 296 | Strigoplus albostriatus K. Sin. ... 280 
, CUNEUUA. mucun sede ener: 300 | Amyciea forticeps O. P. Camb. ... 283 
is TAMOSA  ......0.00..... 803 | Hpidius longipalpis Thor......... sac 280 
a albicrurata  .......5 306 | Myrmarachne maxillosa C. L. 
se VACeralis oases SLO ICOCISe ret ec eee enone ase: 
Fs biseratensis ......... 812 | Agorius gracilipes Thor. ............ 318 
Bt gedongensis............ 316 


The Spiders here described were collected by Dr. Nelson 
Annandale and Mr. H. C. Robinson in the Malay States in the 


years 1901-2, and are now under the charge of the British 
Museum, South Kensington. 


The specimens examined were :— 


THOMISID&. 
IDET OPUS akaen Gna p ia cease 1 
AMIN CURD, aicchiawihi tee ayuicins 1 
DEUS ete ctovtarc aunistaar gra es 1 
SALTICIDA. 
Myrmarachne ..........4. DO 
AG OTUU Sin. 038s werastooe net sat 1 


Many of these were accompanied by ants which have been 
numbered for reference, but which have not been identified. 

The species of Thomiside and of <Agorius are known. Of 
Myrmarachne, with the exception of J. mawxillosé, of which there 
are 13 specimens, all seem to be new species. Material in the 
British Museum available for comparison is, however, scanty, and 
the descriptions and especially the figures dealing with known 
species are so scattered and often inadequate, that certainty on 
this point is at present impossible. 

Myrmarachne maxillosa female was very common, and it is 
curious that no male was found. In general shape M. biseratensis 
is very much like it, but it differs in teeth and tibial spines from 


all previous descriptions of the male of that species. It may 
ultimately prove to be a variety. 


* Communicated by the SECRETARY. 


208 MR. H. D. BADCOCK ON 


M. turriformis is a very distinct species of the Plataleoides 
group. 

M. cornuta is distinguished from all other species fibno 
described by the excess of length of the second over the first 
joint of the pedicel. 

The distinction between M. albicrurata and M. lateralis is not 
very satisfactory, and one specimen occurs which suggests that 
these are only varieties of a single species. 

The males If. ramosa and M. gedongensis are well marked, 
but there is no information to suggest to which (if any) of the 
females they belong. 

The chief diagnostic differences between the various species of 
Myrmarachne are given in the key preceding the descriptions. 
The group is a difficult one. In general shape the prosoma falls 
into three classes, viz., somewhat squat with a narrow and shallow 
thoracic groove and thorax falling rapidly behind it (mawillosa, 
biseratensis), caput towering high above thorax (turriformis), 
and caput and thorax separated by a wide groove of moderate 
depth with caput not much higher than thorax (all the rest). 
The prosoma is further characterized by the presence or absence 
of a wedge of white colour below the thoracic groove, but this is 
liable to be almost obliterated. 

Constriction of the opisthosoma or the absence of it seems a 
constant character, though its depth and the extent of light 
coloration about it are variable. The shape of the unguis and 
armature of the falces in the male are always characteristic. . 

The shape of the sternum, though in cases “appearing very 
definite, is certainly variable to a considerable extent and except 
in extreme cases is a doubtful guide. The epigyne appears to 
differ in the figures, but it 1s probably more due to differences in 
condition than specific differences. ‘The male organs all consist 
of a fine coiled spine of about 13 turns with slight, if any, 
specific differences ; but distinction may be based on the relative 
lengths and shapes of the last three palpal joints and the terminal 
process of the tibia, none of which, however, is particularly 
striking. 

In both sexes the number of spines below the patella and tibia 
afford marked distinctions and are probably as good a first guide 
as anything. They are, however, liable to be rubbed off and are 
not absolutely constant. 

In every case the practice adopted has been to start wih a 
detailed description. This has been followed in the case of 
Myrmarachne by a short description or diagnosis. Finally a note 
is given stating the place where and circumstances under which 
specimens were taken, any particulars concerning them and any 
measurements made. In the case of these last it must be 
remembered that the specimens, having been in spirit for years, 
were very brittle and difficult to measure with accuracy, and that . 
the figures given must only therefore be taken as approximate. 


ANT-LIKE SPIDERS FROM MALAYA. 279 


In the following descriptions I have used Lankester’s terms 
“prosoma” and ‘ opisthosoma ” for the main divisions of the 
body, while retaining “ caput” and “thorax,” employed by Cam- 
bridge, as convenient terms for the two parts of the former. 

The ‘thoracic groove” is the depression between caput and 
thorax. ‘Cephalic wedge” the wedge of white hairs or colour 
at the side of the prosoma reaching downwards from the groove. 
“Width ” or “height ” means greatest width or height. 

The eyes are referred to as 1, 2, 3, 4 in the order—central of 
front row, lateral of ditto, central of back row, lateral of ditto; the 
small eye of the Salticide being considered as 3 in this series. 

The intervals between the eyes are expressed as 1.1, 1.2, de. 
When the shape-of a quadrilateral of eyes is mentioned it is con- 
sidered as the quadrilateral circumscribing the eyes in question. 
The letters f, b, 0, u referring to an eye indicate that it is 
directed forward, backward, outward, upward. 

The basal end of the groove of the falx is the end nearest the 
roct of the unguis. 

The joints of the legs are referred to as c¢, tr, f, p, ti, m, and 
ta, with a subscript letter to denote the particular leg where 
necessary; thus tr, means ‘trochanter of third leg. oo ele 
refer to joints of pedicel. 

In describing the prosoma the order adopted has invariably 
‘been: (a) as seen from above, (>) from the side, (c) from in front. 
In the case of the falces and opisthosoma it has been (a) from 
above, (6) from the side, (c) from below, though in none of these 
has it always been thought necessary to describe from all these 
positions. 

The upper margin of the groove in the falces is always that 
furthest from the maxille, even though this in the case of 
Amyciea and some female Salticids may more correctly be 
described as ‘“ outer.” 


Key to Myrmarachne mentioned in this paper. 


1. a. Second joint of pedicel much greater than first. Opistho- 
soma constricted . eases haa cornuta, 
b. Second joint of ret ae in or more sothen ntehi 1es8 


than first .. Wes ee 2 
2. a. Opisthosoma not poerricrede shee eet rahe eek ec Seeder 3 
_ 6. Opisthosoma aes ee AUR ak ere ee ok seta ce 5 
Dre OCC OL SN ONG ston baw Zits soa ticasactdce oBe as (4eeaa hoomaedonc eins, foveie 4 
b. Pedicel aioree Maton cep ete aaa pane canton Hesse, CULT LOTICIS: 
4. a. Spines on Ist fabs B4o pairs. Sera a ere anata esa MINCE IL LOGO = 
6. Spines on Ist tibia 5 pairs .. Pobdsescendota? eee aiass ess csnene | DISCLALENSIS O « 
5. a. Ist patella no spines, Ist aie! pairs.. Sis Seeetncesue PALOTCHULALO 2s 
b. 1st patella 1 spine ........ er ee 6 
c. 1st patella 1 pair of spines, ae aw bt to 1) ATES oat ii 
Sara statis, A VSpimegpAMS 2:20...) avaes.sccbecheasie ise see< slic sascee’ LAMOSA Os 
ByalststibtaowtOuG pails eat nct arnt Meher eslanesteaease lateralis Q. 
7. a. White wedge on side of prosoma . ME eet elmeata: 2. 
Do NOs Waniber Wiel Omer ean heey dec art hi. etnencsiicheiicceel these” GE@ONGENSIS<S. 


280 MR. H. D. BADCOCK ON 


THOMISID4. 
Srrtcopius E. Simon, 1885. 


@. STRIGOPLUS ALBosTRIATUS E. Sim. (Text-fig. 1.) 


S. albostriatus K. Sim. Bull. Soc. Zool. de France, 1885, p. 144. 

Peltorhynchus rostratus Thorell, K. Sv. Vet.-Akad. Handl. xxiv. 
pe2 (1891): 

Prosoma.—(a) Clypeus nearly straight or very slightly recurved, 
with fringe of stiff red hairs projecting forward slightly; notch 
on sides behind clypeus then swells out to § and then inwards, 
ending in a truncation slightly narrower than width of clypeus. 
Thorax and caput cannot be separated except by colour. Clypeus 
and a triangle from outside eyes 4 to centre of cephalothorax is 


Text-figure 1. 


\ 


“ity 


‘ue N 
AY 
“bay tlife om er ren 


NGAI la Wy 


aN | Wg 


d 


Strigoplus albostriatus E. Sim., ¢. 


a. Prosoma from above. 6. Do. profile. e¢. Do. from in front. d. Sternum &c. 
e. Opisthosoma from above. f. Do. from below. 


light yellow (caput), remainder darker yellow-brown (thorax), 
Caput has a still lighter fine line proceeding forward from apex 
of triangle and stopping just beyond line of 3.3 at a transverse 
line of stiff hairs similar to those on clypeus. ‘This line is 
slightly expanded into a dot immediately after its start. A 
similar light line crosses this at right angles between 4.4, the 
two lines together forming a faint but distinct cross. Eyes 2 and 
4 are on tubercles with a distinct blue tinge which is striking. 
Line centre to eye 4 is marked by two light dots similar to that 
on central line, and these dots form approximately the line of 
demarcation between the light and dark portions (caput and 


ANT-LIKE SPIDERS FROM MALAYA. 28] 


thorax) of prosoma. There are also less distinct dots or streaks 
radiating from centre and marking other thoracic divisions. 
Dark portion is bordered by a thin white marking at hind 
corners, and there is a very thin darkening at outer edge. 
Cephalothorax is practically free from hairs except the bristle 
fringes and a very slight thin pubescence at edges. 

(b) Clypeus slopes very gradually backwards and upwards with 
convex section, and on this view is equal in length to eye-cliff, 
its rise being very slight. Portion of face containing eyes 2 now 
rises vertically or rather slightly overhanging, and from here to 
centre of cephalothorax (which is well behind eyes 4) top is flat 
with slight rise; thence 45° slope to rear. White marking is 
prominent at rear end and can be traced round to front as a fine 
white line. Hyes 2 and 4 are also prominent on side view. 

(c) Clypeus slightly procurved and about = eye 1; top convex, 
sides straight. 

Cephalothorax has somewhat the appearance of a death’s head. 

Eyes.—\|st row distinctly procurved. 

1.1 nearly =2x1.2, on slight tubercles of bluish tinge, 
Onued: 

2=2x 1, largest of the 4, on blue tubercle connected by valley 
with tubercle of 4, the whole markedly blue, o. u. f. 

3=4 x1, smallest of all, u. 

4 very slightly <2, blue most marked and tubercle most pro- 
minent, o. b. u. 

2.3 very slightly <1.1; 3.3=4.4 approximately. 

Total length of 2nd row greater than Ist. Trapezium 1.3.3.1 
is narrower behind than before and approximately as ai as 
long. 

Falces nearly horizontal, projecting forward and very pro- 
minent from above, broad at base and rounded, rapidly reducing 
to apex and curving slightly apart on inner side so that there is 
a distinct space between. Short tooth-like spines from about 
3 to apex, one on top near commencement being distinctly 
stronger than the rest. Lower side free from hairs. Groove 
indistinct and toothless. Outer margin has a thin scopula of 
moderate hairs. 

Maxille.—Long, nearly = falces, broadest at base, slightly con- 
eave without and within round labium and tapering to a blunt 
point, greatest slope being on outer edge. Short spines all over 
lower side. Serrula on outer edge after concavity. Scopula 
strongest at angle but also slight on inner side. 

Labiwm shaped like a Florence flask, length = maxilla = 
3 breadth ; blunt-pointed. 

Palpi—One missing, other broken after patella.  Light- 
coloured. stout, cylindrical. 

Legs.—Mostly detached and damaged. 

lst. One missing, other broken after femur. Light especially 

on top of f, which is marked with ‘somewhat indistinct 
longitudinal series of white spots. 


Proc, Zoou. Soc.—1917, No. XX. 920 


‘ 


282 MR. H. D. BADCOCK ON 


2nd. Two detached legs are probably 2nd. These have :— 


f dark with distinct large white spot at centre and one at 
apical end on each side, and a fainter one at basal end 
on one (% both) sides. p dark. 

ti=f=3 p, a little lighter and with white annulation about 
centre; basal } darker and < apical. 

m < ti, deve at Hae) then a white spot followed by hight 
end. 

ta light, <m, and distinctly lighter for basal half. 

Claws Zs pectinated with about. 5 teeth. 

3rd. Only one remains. Much lighter, no distinct rodent 
Slight hairs thickest on m & ta. 
4th. Both missing. 


There are three detached legs resembling 3 and three resem- 
bling 2; presumably, therefore, 3 and 4 are much alike and 2 
and 1, though the detached one is not much like the attached 1. 
In-any case none has any spines, though all have considerable 
hairs on ti, m, ta. All joints are cylindrical except f, and f,, 
which are slightly bow-shaped though not angular. 

Sternwm.—Broad, nearly an equilater al triangle, with fore 
side straight, sides slightly convex, and hind end a blunt point. 

OprstHosoma.— Light and dark brown. 

In front a slightly recurved narrow transverse white line with 
short central white line projecting back and rather indistinet 
returns at ends. . 

About middle a white line, whole width of abdomen, waved 
doubly at centre. 

About same interval another straight line broken on each side 
of centre. 

4th line wavy and shorter. 

5th shortest, just above spinners. 

The darker brown is more or less bounded in front by the 
white line, light after 3rd. 

Shape an amphora with spinners for neck. Sides have some 
more or less irregular white splashes or spots which bound lower 
surface ; a pair on each side of lung-sacs and a shortest pair at 
base of spinners are very marked. 

Below, brown. Lung-sacs small, light, fairly wide. Epigastric 
fold marked, pr ocurved. 

Spinners. —1st Gacelishanod ; 3rd longest and thinner; all 
hunched. 

Note.—Description from a single ? somewhat damaged, from 
Sungkei, found in company with ant no. 2. The prosoma and 
opisthosoma were separated and the legs were not ‘measurable, 

Prosoma 1-2 long x 1:2 wide x 1:1 mm. high. 

Opisthosoma 270 x 1°8 x 1:4 mm. 

ae prosoma is widest and highest at 2 3, the Ppleclere a widest 
at 3, highest at 3. 

Total length estimated at 3 mm. 


ANT-LIKE SPIDERS FROM MALAYA. 283 


Amycr@A KE. Sim., 1885. 
@. Amycr#a ForticEPS O. P. Cambridge, 1873. (Text-fig. 2.) 

Amycla forticeps O. P. Cambr. P. Z. 8. 1873, p. 122. 

Amyciea forticeps Thorcll, Spid. Burm. p. 282. 

Prosoma.—(a@) From above truncated but slightly recurved 
in front, the same but slightly procurved behind, sides slightly 
convex, widest a little behind centre. 

(6) Nearly as high as long, very convex, very steep in front to 
eye 2, then shghtly and convexly rising to highest point over 
eye 4, then with a more gradual and very slightly convex slope 
to hind end. 

(c) In front sides convex, subparallel, front edge slightly pro- 
curved. Clypeus=eye-space. 

Colour reddish yellow, with a few black hairs on front edge and 
between eyes. Very faint traces of fovea immediately behind 
eyes and striz therefrom. 


Text-figure 2. 


e 


Amyciea forticeps O. P. Camb., 2. 
a. Profile. 6. Prosoma from in front. c. Sternum &c. 


Hyes.—¥irst row straight and subequally spaced, 1.1 being 
possibly very slightly <1.2. 2 large on bluish tubercle = 4x 1. 
‘2nd row strongly recurved so that 3.3 is well in front of 4.4. 

Quadrilateral 1.3.3.1 is very much wider behind than before. 
3 considerably <1. 

Quadrilateral 2.2.4.4 slightly wider than long, a rectangle. 
4 large on blue tubercles but slightly < 2. 

Falces not longer than clypeus, slightly stouter at base than 
apex, which is cut square. On side view rather more tapering. 
Unguis small. A weak scopula below unguis, and inner sides of 
falces with short, dark, stiff hairs in a regular row, longest at 
unguis. : 


20* 


284 MR. H. D. BADCOCK ON 


Maxille.—Narrow at base, at first parallel then curving in- 
wards to rounded points facing each other in front of labium. 
On inside curves closely round labium. Serrula from well on 
outer side round end to point, very weak. Scopula very weak 
just inside point. 

Labiwm.—Length = maxilla. Breadth <4 length. Sides . 
slightly constricted immediately after base and then convex to a 
round point. 

Palpi.—Spring from base of maxilla. All joints cylindrical. 
f much longest, >c, ; p, ti, ta increasing in length, but all short ; 
‘ta has a single simple claw. 

Legs.—2.1.4.3, with 2 and 1 practically equal and marked 
differences between the others. All are cylindrical except coxe, 
which are rather rectangular on section. Proportions of joints 
much the same in all, f, ti, m, ta being in decreasing order 
with m about 4 ti or 2 ta. Femurs of 1, 2, and 4 have a weak 
short spine above near apex, otherwise there are no spines. 
Tarsi have two claws, pectinated, with more teeth on one than 
the other ; teeth close together and extending threughout. 

Sternum.—Shield-shaped, cut straight and broad in front, and 
with a short point behind. Nearly as broad aslong. ¢;,1 whole 
width of sternum, ¢c4,4 close but not in contact. 

The colour of the prosoma including appendages is a dull 
orange-yellow. 

Pepicet.—Moderate, distinctly visible from above, slightly 
tapering forward, single-jointed, covered at top with a long 
narrow lorum. Nearly white. 

OpristHosoma.—Length nearly = 2 width or height. Some- 
what flask-shaped from above, distinctly widest at %. Light grey 
or mouse-colour, with two longitudinal sets of white spots, rather 
irregular in disposition, proceding from front to widest place and 
spreading outward. These terminate each in a large black spot. 

Below plain. Lung-sacs very dark and wide apart. LEpi- 
gastric fold distinct. 

Epigyne.—Rosette-shaped. Not noticeable. 

Spinners.—All short; 1st and 3rd pair making a rectangle 
considerably breader than long; 2nd pair between 3rd, so that 
3.2.2.3 is a straight line. All spring from sockets in a 
common projection, all 2nd joints very small. 

Ist stout barrel-shaped. 2nd shortest, cylindrical, thinnest. 
3rd cylindrical, intermediate. 

Anus.-—Tubercle rather broad and thin, nearly as high as 3rd 
spinners. 

Note.—Description is from a single 9 dropped from a tree in 
the jungle at Biserat. 


The following are the measurements :— 

PROSOMUG Re Hin Siete: 2°0 x 1:5 wide x 1°6 mm. high. 
Opisthosoma ...... 27 < LSeeee x aled Pe 
Palpus see. ne. rian ae 


ANT-LIKE SPIDERS FROM MALAYA. 285 


Legs.—1 ...... c+tr 0°6, f 2:2, p+ti 2:2, m+ta 22 =7:2 
yA ts 0°6, SRE DMs tls 2-2, al Oe ua dee 
Dilip tees cc OSG sia Leo, ptr +m+ta2°4 =4°5 
ov ney poe OF Om eel IE F 37 =5°4 


Ant accompanying, none. 
Eripius Thorell. 


3g. Eprpius tonerpaueis Thor. (Text-fig. 3.) 
E. longipalpis Thor. St. Rag. Mal. i. 1877, p. 152. 


runcated or very slightly convex in 
front, swells out slightly and concavely at sides to thorax, which 
is much wider and rounded at sides and slightly concave at rear. 
Caput is cut off above by strong striz, and there is a deep fovea 
and thoracic striz. Caput has a central longitudinal indentation 
from fovea to 2nd line of eyes, Colour dirty brown, lighter at 
indentations and rear of thorax. 

(b) Clypeus vertical, gradual slope up from Ist eyes to mid- 
thorax, then more abrupt slope to end. 

(c) Clypeus nearly = face, very slightly procurved. Caput 
convex above and at sides, thorax convex at sides and flattened 
above. 


Text-figure 3. 


Epidius longipalpis Thor., 3. 


a. Prosoma from above. 6. Do. from in front. c¢. Sternum &c. 
d. Tarsus. e. Palpus. 


Eyes.— 1st row recurved, 1.1 =1.2, 1 = 42. 2nd row re- 
curved as viewed from above but ace s0 than “Tat row, slightly 
procurved as viewed from in front, considerably longer than Ist. 
3.3 very slightly >3.4, 4 very slightly > 3. 

Trapezium 1.3.3.1 longer than broad and nearly twice as 
broad behind as before. 

Falces.—Vertical or very slightly sloped forwards, parallel 


286 MR. H. D. BADCOCK ON 


sides with very oblique long truncation, groove distinct. Upper 
margin, one small tooth at angle with one smaller above it, and one 
or more very small below. Lower margin, one tooth half way to 
angle and two at angle, one of which is fairly large; these two 
may, however, be really one with two cusps. <A strong and long 
scopula of light hairs on outer margin. 

Maxilla.—Moderate, length =14 breadth, subparallel, being 
shghtly wider in front than at base. Outer apex rounded, inner 
obliquely truncated with scopula. Serrula from well before 
outer rounding to truncation. 

Labium.—Length > maxilla, broader than long, strongly 
barrel-shape. 

Palpi.mRemarkably long = about % leg 4. Tr simple, much 
broader than long, breadth = nearly least width maxilla; f length 
= prosoma, cylindrical, slightly S curved; p very short, <+ ti; 
p+ti>f, ti<f and shghter, S curved; four steut bristles at 
lower apex pointing directly forward and reaching beyond base 
of fovea; a short sharp apophysis pointing directly forward at 
lower outer apex. Ta a little >p, very short, subelliptic, with 
blunt point and fovea occupying about half of it. Organs have 
largish but flat bulb ending in blunt-pointed extension forward. 
Style springs from centre and points forward over extension of 
bulb, ending in a sharp black spine-lke point. 

Legs.—Only Ist and 38rd on one side remain, Ist being much 
stronger and about twice as long as 2nd. 
lst. ec length = nearly 2 breadth. 

tr, a few short dark hairs at ridge, fore section distinctly < aft. 

f, 2 or 3 spines at top in longitudinal row, a very few 
smaller at side. 

p. =} ti, hairless. 

ti slightly curved, slenderer than f; 3 or 4 spine pairs and 
a few spines along upper sides. 

m nearly =,ti, two spine pairs below but stronger and a few 
at sides. 

ta = 3 m, hairless, 2 claws with 5 teeth and fasciculus. 

3rd. General proportions as Ist, but much shorter and f 

stronger. Spines on back of f and p, others as Ist. 

Sternum.—Cordiform, as broad as long, very smooth, front 
corners rounded, rear point rounded. c.; = full width of 
sternum. c4y,4= width of c,. 

PrpiceL.—Short and stout, with a darker narrow lorum above. 

OpistHosoma.—Very much damaged, but appears to be almond- 
shaped from above, with two or more pairs of spots in a longi- 
tudinal series at the fore part above. On a side view very high 
in front, rising vertically or with a slight overhang from pedicel 
and falling with a straight slope to spinners. 

Spinners compact, short. Ist stout at base, tapering, 2- 
jointed. 2nd slender, longer, cylindrical. 3rd in shape and 
proportions between Ist and 2nd. 

Anal tubercle.—Wide and noticeable. 


=" 


ANT-LIKE SPIDERS FROM MALAYA. 287 


Note.—Description is from a much damaged ¢ specimen from 
Biserat, Jalor, in company with the Salticid spider M/yrmarachne 
biseratensis ¢ @ and three ants of species 2 and 17. 

The following measurements were taken :— 


WE OSOMUG! Gee vic. es +s 1-7 long x 1:7 mm, wide. 

Opisthosoma ...... 2°0 or longer, damaged. 

Palpi::..<.. f 1:8, p very short, p+ti2°0 =abt. 4:0. 

Legs.—1...... e+tr+f 3°6, p+ti 3°9,m+ta40 =11'5 
om ae 55 Oi ay AO ge PLO, ae Org) 


SALTICIDA. 
MyRMARACHNE Macleay, 1839. 


Q. MyrMaracune MAxILLOsA C, L. Koch. (Text-fig. 4.) 
Toxeus maxillosus C. L. Koch, Die Ar. xii. p. 19, tab. cecelxxvi. 
fig. 1090. 
Synemosina procera Thor. Ann. Mus. Genov. x. p. 538 (1877). 
Toxeus procerus id. ibid. xxv. p. 346 (1887), xxxi. p. 220 (1892). 
Salticus modestus id. Ann. Mag. Nat. Hist. ser. 6, ix. p. 235 
(1892), Q jun. 
Toxeus maxillosus id. Spid. Burm. 1895, @. 


Text-figure 4. 


Myrmarachne mavillosa C. L. Koch, 2. 


a. Profile. 6. From below. 


Prosoma.—Length = patella + tibia of 4th leg, slightly longer 
than opisthosoma. Greatest breadth and greatest height each 
nearly 4 length. Caput = or very slightly < thorax. Front wall 
straight or very slightly recurved and stepped back at eye 2. Side 
walls straight or very slightly convex. Back wall somewhat pro- 
curved as it slopes down to the groove, where the cephalothorax 


288 MR. H. D. BADCOCK ON 


is narrowed above but not at base. Thorax swells out again 
behind groove towards width of caput, being strongly rounded, 
and terminates in a truncated or even slightly hollowed end of 
about half greatest width of thorax. 

Clypeus very low, top of caput flat and sloping very slightly 
upward to eye 4, over which it is convex. Groove is very shallow, 
hardly reaching as low as bottom of eye 4. Thorax rises very 
slightly behind it, and then falls with an even but slightly convex 
slope to rear end, where it rises concavely into a rim which can 
be traced along sides to caput. 

Colour dark mahogany, with black eye-frame broken in front 
of eye 4 and terminating just below the level of eyes 2 and 4. 
Scanty short grey hairs all over (Text-fig. 4, a.) 

Hyes.—1st row straight by summits. 1.1 in contact, 1.2 
close but not in contact. 2=41 and slightly behind it. 4= or 
very slightly > 2.3 < 4 2, very slightly nearer 2 than 4 and on 
the upper tangent to 2 and 4. Hye square slightly broader than 
long, with eyes 4 projecting prominently at hind corners and 
4.4 very slightly > 2.2 and > distance between itself and rim 
of cephalothorax. 

Falces.— Very nearly as long as caput. Length > 2 breadth. 
Project horizontally forward. Sides subparallel, with inner edge 
straight to about § and then rounded off to base of margins, 
outer edge very slightly convex. On side view straight at lower 
edge, upper edge strongly convex making falx very much thicker 
at + to 4 than elsewhere. Unguis ==4 falx, sickle-shaped with 
base much thicker than tip and curve with a tendency to being 
straight in the middle with angle inside near base. Groove is 
marked. Upper margin has typically 8 teeth, of which the first 
three are close together at base of unguis, increasing rapidly in 
size from 1 very small to 3 large, 4-6 are large teeth, 7, 8 rapidly 
decreasing till 8 is very small. 4-8 evenly spaced. 1 and 2 and 
8 are sometimes missing. Lower margin has 8 to 10 or more 
very small teeth close together and reaching from base of unguis 
to a little beyond angle, but always terminating before the upper 
ones. They or at any rate the middle ones are alternately large 
and small. There is a strongish scopula of long grey hairs on 
upper margin, largest at angle, and of short dark scanty hairs on 
lower margin. 


Colour dark mahogany as cephalothorax. The falces are 
glabrous and somewhat rough. 

Maaille.—Length = coxa + troch. of 1st = falx nearly. Outer 
edges parallel from base to apex of labium, then still remain 
straight but incline slightly outwards. Outer corner is well 
rounded and end square, a serrula following the edge from about 
middle of outer side to end of straight portion. Inner corner very 
slightly rounded and truncation straight and nearly parallel axis 
runs back more than half way to labium. Hence slopes slightly 
outwards and follows labium back to base, being hollowed out to 
enclose it. . Truncation has a strong scopula of black hairs which | 


ANT-LIKE SPIDERS FROM MALAYA. 289 


is extended towards labium and also outwards to inner angle at 
termination of serrula, where hairs are grey and as long as width 
of maxilla. 

Width at end = lJabium. Colour light mahogany except for 
outer edge, which is as dark as cephalothorax. 

Labium.—Iin length % maxilla, terminating half way up it. 
Breadth 4+ length. Tapers very slightly from base to apex, which 
is square. Sides straight. Apex has a scopula of dark hairs. 
Colour as dark as outer portion of maxilla except for apex, which 
is ight. 

Palpi.Spring from near base of maxilla. Trochanter length 
=width maxilla. Dark brown except for light yellow on inner 
basal side. Femur > half length of caput, flattened, narrow 
at both ends where its width > 4 length. Patella < 4 femur, 
wedge-shaped, broadening at apex from narrow base. Tibia 
broadens from base to apex, where it is twice width of patella 
apex. ‘Tarsus slightly longer than patella, with rounded point. 
Tibia and tarsus together form a lanceolate plate, glabrous below, 
nearly twice femur in length, with strong fringes of hairs on 
both sides and round point. All joints except patella have short 
grey hairs on back. General colour dark brown-green, except 
as mentioned for trochanter. Total length of palpus where 
extended = 2 falx, the basal end of tibia being about level with 
apex of falx. 

Legs.—Order 4.1.5.2, with 4 and 1 practically equal and 
likewise 3 and 2. In the lst leg patella + tibia and in the 4th 
metatarsus + tarsus are notably long compared with the other 
legs. 

Ist. Coxa length = 23 breadth, very slightly tapering, light yellow 
except for brown patch at lower outer vertex. 

Trochanter = ¢ coxa, narrower, light yellow except for con- 
tinuation of coxal brown patch strengthened. 

Femur strongly clubbed at base, < patella + tibia > tibia, 
dark except below at apical end. 

Patella > 3 tibia, slightly curved downwards. No spines. 
Dark at sides, hght above and below. 

Tibia. Has normally 7 spines on the inner and 6 on the outer 
side below, but there are frequently 6 pairs. The spines 
are evenly spaced from base to apex, but the largest are 
in the middle and they get rapidly shorter, so that the 
apices of the last 3 or 4 are all about level near the end of 
the joint. Dark at sides, light above and below. 

Metatarsus > patella, with two pairs of strong spines longer 
and stronger than those of tibia, the last of which reach to 
beyond the middle of tarsus. Dark. 

Tarsus < 3 metatarsus, with claw-tuft and 2 claws. 

2nd.c equally broad and long, swollen near base, < 3c,. Dark. 
brown with a touch of light at apex. 

tr short = 3c, cylindrical, brown with light patch below. 

f slightly clubbed at base, brown with light below at apex. 


290 MR. H. D. BADCOCK ON 
p =¢# ti, curved downwards, no spines, Light above and 
below, dark sides. 
3 pairs of feeble spines evenly spaced, with a 4th pair 
extremely short and feeble at extreme apex often missing. 
Light above and below, dark sides. 
m> ti. 2 spine pairs stronger than on ti, but much weaker 
chan on m,, basal is lar hea apical only just reaches ta. 
Light oowe fail below, dark at sides. 
ta< 3m. Light. 
ards Ca nae larger and move cylindrical than 2nd Cc, buowall 
tr short, cylindrical, brown. 
f brown, slightly clubbed at base. 
p short, curved downward, brown except for light patch 
above at basal end. 
ti brown except at tip where light above. 
m. feel) = = ti, light above and ‘below brown sides. 
ta > 3m, light. 
4th. c > ec, but not more than § ¢,, cylindrical, brown. 
tr larger and a little slenderer than ¢, cylindrical, light. 
f long and slightly clubbed at base, pravent 


Peep 
ti brown throughout, slender. 
m very long, nearly = ti, slender, brown except for very 


small hghter patch at upper apical end. 
ta = ¢ m, light above and below, dark at sides. 


In Eee appearance the posterior legs are much darker than 
the anterior, and the light (almost white) Ist ¢ and tr and 4th tr 
are very noticeable. 

Sternum.—Lanceolate, short-pointed between C1. 1, which are 
; their width apart. Widest from C1.2 to C2. 3, thence tapering 
to a long point in front of ¢4,4, salvich are in contact. Sides 
undulate to accommodate coxe but do not run into points 
between them. Broad for the genus, being > width of middle 
cox. Gaps c),2 and c2,3 about equal and not materially greater 
than ¢3,4. Colour mahogany, lighter than cox 2,3, 4. (Text- 
fig. 4, 6.) 

PrpicEL.—Short but visible from above. First joint covered 
by brown plate overlapping second. Second joint longer and 
covéred by longitudinal narrow plate. Whole length of pedicel 
not greater than its w dth. It is received into a short projecting 
socket in abdomen. | 

OpisrHosoma.—EHee-shaped from above, widest about 3 back. 
From side overhangs in front and about same level for first 3. 
Above, general colour a dark green-brown. Below, general colour 
the same but coarser, and at “sides distinctly composed of darker 
lines separated by light spaces. Lung-sacs very wide apart and 
dark red-brown, space between around epigyne lighter red-brown. 
A smoother and darker longitudinal band from epigastric fold to 
spinners, slightly wider at spinners. 

Kpigastric fold procurved. 


ANT-LIKE SPIDERS FROM MALAYA. 291 


Epigyne.—A pair of white oval spots with a darkish mahogany 
bridge between expanded at both ends. The whole ou a ground 
of the same colour. (Text-fig. 4, 6.) 

Spinners.— 1st stout, subconical, light green-brown. 2nd very 
thin, white, slightly longer than Ist. 3rd a little stouter and 
darker than 2nd, much thinner than Ist, slightly longer than Ist. 

All have 2nd joint very much longer than Ist. 

Anus on a tubercle of two joints. 


Suort Descriprion. 

2. Prosoma = opisthosoma. Caput = thorax. Groove very 
slight, with thorax not rising appreciably behind. 

Falces project horizontally forward, nearly as long as caput, 
giving the appearance of ¢. Teeth, upper 8 strong at centre, 
subevenly spaced ; lower 8 small, alternate sized, close. 

Legs 4.1.3.2, with 4 and 1, 3 and 2 nearly equal pairs. c, 
and tr, and tr, are very prominently white. Ti, has 6 spine pairs 
or more often 6 and 7 spines, m, 2 pairs., p, none, p, none, ti, 
3 pairs, occasionally a 4th, m, 2 pairs stronger than on ti,. 

Sternum rather broad. Gap c2.3 not > others. 

Pedicel short but visible from above. 

Opisthosoma squat, much thicker than usual in the genus. 

Note.—Description based on 13 9 from Biserat, K. Mahek, 
c. Patani. It was found practically everywhere and is much the 
commonest. ' 

2. The ¢ is possibly JZ. biseratensis. 

Length varied from 3°5 (imm.) to 7°0 mm., the normal length 
being 5 to 7. 


The accompanying ants were numbers | and 2. 
Legs of a specimen, 2 :— 
| aes e+ti 1:2, £2°5, p+ti2°9, m+ta 16 


==8°2 
7 apa c--f 2°0, p-ta 2:9 =4°9 
Barrer c-f 2:0, p-ta 3°2 = D78 
es e+til4, £2°7, p+ti2°9, m+ta2°3  =9'3 
Legs of another specimen, length 6:0, 9 :— 
Tee ae c-f 3°6, p+ti3:2, m+tal4 =8-2 
Dianne Ree os tne oie p-ta 2°8 =U) 
teh ate ee See 3 —)".) 
LS ee » 26, p+ti2-7, m+ta26 =89 


MYRMARACHNE CORNUTA, Sp. n. (Text-fig. 5.) 


2. Prosoma.—(a) Length = opisthosoma. Greatest breadth 
very slightly >‘greatest height and each nearly 2 length. Caput 
= thorax. Front wall straight, slightly stepped back to eyes 2. 
Side walls slightly convex. Rear wall procurved where it slopes 
back to groove where p.s. is narrowed above but not at base, 
though white wedge makes it look as if strongly constricted. 
Thorax swells out again behind groove to width of caput and 
then narrows with straight or almost concave edges to base, 


292, MR. H. D. BADCOCK ON 


which is cut nearly straight and about 4 greatest width of 
thorax. 

(6) Clypeus very low. Caput slightly convex. Groove shallow 
and broad. Thorax rises behind it to nearly height of caput and 
then falls with convex and concave slopes to rim, which is slight 
but can be traced round to caput. 

Colour mahogany moderate to dark, with black eye-frame 
broken in front of eye 4 and terminating just below the line of 
eyes 2and 4. In dark species this frame is hard to distinguish. 
Thorax a shade lighter than caput. Between the two on each 
side a wedge of white hairs broadest below. 


Text-figure 5. 


Myrmarachne cornuta, sp. n. 


a. 6: Profile. 6. ¢: Right falx from below, unguis omitted; ¢. Do. from above. 
d. $: Right palpus from below. e. 9: Epigyne. 


Eyes.—1st row procurved by summits. 1.1 subcontact, 
1,2>1.1 but very close. 2>41 and slightly behind it. 4=2. 
3< 72 and very slightly nearer 2 than 4 and on the upper tan- 
gent to 2.4. Hye square broader than long and 4.4 distinctly 
> 2.2and much > 4-rim. Hye 4 projects over edge of prosoma 
but not at all prominently so, and the prosoma is not convex over 
it in either view. 

Falces.— Length < 4 caput =14 breadth, nearly vertical with 
slight forward slope. Outer edge convex, inner more markedly 
sloping inwards from about ? to root of unguis. On side view 
strongly convex above with almost an angle at 4 so that basal 
portion is horizontal and apical nearly vertical; straight or 
very slightly convex below. Unguis regular curve and regular 
diminution in thickness from base to apex. Teeth on upper 
margin 5 or 6, of which 3 above angle evenly spaced and of 


ANT-LIKE SPIDERS FROM MALAYA. 293 


moderate size, others below angle and smaller. Lower teeth 7 
or 8, close, about same size from end to end, but alternate ones 
possibly smaller. Long thickish light scopula largest at angle on 
outer margin. A few long dark hairs along lower margin. 

Colour mahogany lighter than cephalothorax. Unguis at base 
darker, at apex lighter than paturon. 

Maville.—Length > ¢,, nearly = falx. Narrow at hase, outer 
side curving strongly and concavely outwards beyond labium to 
bold rounded corner, truncation shortish and at about 45° with 
axis of body. Inner side slopes outwards from truncation, being 
strongly hollowed in front of and at side of labium. Dark serrula 
from outer corner to angle. Light scopula along truncation. 
Width at base =+ labium, at end > labium. 

Colour mahogany, markedly lighter at inner end. 

Labiuwm.—Length > 3 maxilla. Breadth at base > 3 length. 
Sides parallel at base, slightly inclined towards each other at 
apex, which is cut off square and has thin scopula of dark hairs. 
Colour darker than darkest part of maxilla except at apex 
where light. 

Palpi.Spring from close to base of maxilla. Trochanter < 
width of maxilla. Femur = + caput, flattened, reduced at ends, 
greatest width = + length. Remainder as I. macillosa, 

Colour = caput with end joints darkest. 

Legs.—Impossible to measure lengths. 

Ist. Proportions as mawillosa except patella =4 tibia. Patella 
has 1 spine below towards outer side. Tibia 4 spine pairs, 
subequally spaced and nearly of equal size but with 
basal slightly the greater. Apex of last spine just reaches 
end of joint. Metatarsus 2 spine pairs, basal strongest 
and somewhat stronger than tibial, apex slightly beyond 
end of joint. 

Colour very light throughout except small dark splash ° 
on outer side of trochanter and larger dark splash towards 
outer basal end of femur and slight darkening on outer 
side of metatarsus. In a light specimen all these dark 
markings are absent or merely shades. 

2nd. e¢ nearly as broad as long, markedly swollen on inner side 
about centre. 

tr = 4c, barrel-shaped. 

f slightly thicker at base than apex, > ti < p+ti. 

p = 3 ti, ti 2 spine pairs. m=p, 2 spine pairs. 

Colour light throughout except strong dark longitudinal 
marking on outer sides of c and tr. 

3rd. Proportions as far as patella = mazillosa. Dark brown 
c to f, light beyond. 

Ath. c > ¢,, cylindrical or very slightly wider at base. 

tr = 3¢, slightly barrel-shaped, both these joints are brown 
with longitudinal light centre line below. 

f brown, slightly thicker at base. 

Remainder missing. 


294 MR. H. D. BADCOCK ON 


The white or very light yellow anterior legs are very noticeable, 
as also the proportions of c, and tr,. 

Sternum.—Sharply but shortly pointed between c,,1 which are 
in subcontact, short-pointed at sides between ¢;.2 which are close, 
curves cut strongly between c2,3 which are wider, narrows again 
before c, and then proceeds with straight parallel sides to ¢,, in 
front of which it ends in a moderately long point. ¢4.4 in 
contact. At widest sternum about =c,. Colour dark mahogany. 

PrpiceL.—Length > caput. 2nd joint considerably longer than 
Ist (as. seen from above j, = 2j,). Ist plate broader than 2nd 
and procurved behind. Pedicel received into very short projecting 
socket of opisthosoma. 

OpistTHOosoMA.—Subcylindrical with constriction at +, widest 
at 3, pointed at spinners. Dark green-brown above, with white 
marking at constriction starting at top of sides and widening 
below. In front of epigastric fold and behind the white marking 
colour is as above. Epigastric fold straight ; lung-sacs wide apart 
—and dark. 

Epigyne.—An equilateral triangle with base along epigastric 
fold and vertex forward. Edges red, inside white, with central 
darker bridge and darkest spots at its edges. Not very distinct 
but more characteristic than others of the genus. (‘Text-fig. 5, e.) 

Spinners. — Proportions as maillosa. All a light brown 
colour. 

3. The chief differences from @ are:— 

Prosoma.—Cephalothorax = # abdomen. Groove sharper at 
bottom and thorax does not rise so much behind it, so that 
it is considerably below caput and general shape not unlike 
maxillosa. Rim as well as wedge has grey hairs. 

Falces = cephalothorax. From above inner edge straight at 
first, apical half concave, outer edge convex. Width = length. 
End cut off square, with strong concavity and very strong tooth- 
like projection forward and inward from inner angle, with blunt 
end. From side falces project straight forwards, with upper and 
lower edges almost parallel, granulated. 

Groove not well defined. Upper margin has 6 teeth. Ist 
smallish close to base of unguis, remainder subequally spaced 
over basal half of falx, 2-5 being very large, 6 small = 1. 

Lower margin has 9 teeth, mere dots subequally spaced. 

Unguis darkest and strongest at base, S curved with slight 
swelling of tip-curve. Length = paturon. (Text-fig. 5, 6, c.) 

Macille.—Rather straighter outer edge and stronger serrula. 

Palpi.—Trochanter = width maxilla. Femur = 4 caput, less 
flattened than in 2. Patella < trochanter, curved downwards ; 
from above, sides almost parallel, apex distinctly wider than 
base. Tibia length = patella, much wider at apex. A very 
small blunt process on outer apical corner visible from below. 
Some longish hairs on inner side. Tarsus longer and broader than 
tibia, oblong, with short hairs on inner side. Fovea reaches to 


ANT-LIKE SPIDERS FROM MALAYA. 295 


about 3. Organs have a black spine springing from near inner 
apical end, encircling fovea outwards and backwards and termi- 
nating in a fine point pointing outwards at about middle of apex 
‘after completing somewhat more than a circle. (Text-fig. 5, d.) 

Legs.—4 considerably > 1; 3 missing but up tof = 2. Gene- 
rally as Q except that in 2 anterior pairs there are strong black 
lines along sides of joints which are much strongest in femurs. 
and the white centre lines of c, and tr, occupy the whole of the 
lower side, and ti, has 3 spine pairs instead of 2. 

OpistHosoma.— Abdomen longer and narrower. Width = 1 
length. White marking at sonstnicuen only at sides. Above 
and below central brown coriaceous lines running nearly the 


whole length of the body. 


SuHortr DEscrRIPrion. 


Cephalothorax = abdomen in 2 = ?abdomen in 3. Caput= 
thorax, moderately high, with broad but shallow groove and 
thorax lower. Strong white-haired wedge at sides of groove, and 
3 white hairs along rim. 

@ falx: upper teeth 5 or 6, subevenly spaced, 3 above angle, 
moderate, remainder small; lower 8 or 9, close, small. 

J falx = cephalothorax, subparallel edges, inner concave near 
apex and terminating in strong bow. Unguis, § curve, slight 
swelling near point, =paturon. Upper teeth 1 at base of unguis, 
5 evenly spaced over basal half, of which 2-5 very strong. 
‘Lower 9 dots evenly spaced with dark ridge at base. 

Sternum narrow, rounded between ¢2,3, short points in front 
and between ¢}. 2, longer point behind, ¢c;,; about + their width 


apart. 
Legs.— First 2 pairs white, black side-lined in ¢. Posterior 
dark with c¢, and tr, dark helow, > .c,. tt, = = 3¢,. Order 4.1 


(2. Bh P, 1 spine ‘below towards outer side, ti, 4 spine pairs, 
m, 2 spine pairs, ti, 9 2 spine pairs (¢ 3), m, 2 spine pairs. 

Pedicel: 2nd joint notably longer than Ist. 

Abdomen cylindrical, narrow, gently constricted at 4, with 
white oblique lines down side, in 2 extending below. has 
brown longitudinal centre above and below. 

Kpigyne an equilateral triangle. 

3 palpus: very small process at outer apical angle of tibia. 
Organs are encircled by a black spine pointing outwards at centre 
of apex. 

General colour of prosoma dark mahogany with even darker 
eye-frame, of opisthosoma dark grey-brown. 

Mote The description is based on 1 ¢ and 2 9, one of the 
latter being much lighter in colour. The ¢ and this 2 were 
both found at Bukit Besar, alt. 2500 ft., on tree-trunk 35 ft. from 
ground on 6 Sept. 1901; the other 2 at Bukit Besar. 

Ants accompanying, nos. 12 and 16. 


296 MR. H. D. BADCOCK ON 


The following measurements of the ¢ were taken :— 
Length 8-0 mm.,__p.s. 3:1x2°0x2-0, 0.8. 4:0 x 1:0. 

ihegs ae e+f 2:2, p+ti2d, m+tal2 = 5-5 

7 Aes AU). miele » hth is 

ened Sa aes missing 

ers e-etr 15.0 a ls (Pies woe Soke, Oe ee meee 
No measurements of 2 legs were obtainable; total lengths of 

the two 2 5:8 and 5:0 mm. 


MYRMARACHNE TURRIFORMIS, sp.n. (Text-fig. 6.) 

2. Prosoma.—(a) Length considerably > p+ ti of 4th = 
p+ti+m-+ta of 3rd = 24 width = 23 height. Caput < thorax. 

Front wall of caput straight or very slightly recurved, and 
stepped back at eye. Side walls straight. Back wall procurved 


Text-figure 6. 


Myrmarachne turriformis, sp. i. 


a. 6: Profile. 6. g: Sternum. ec. ¢: Left palpus from below. 
d. 9: Epigyne. 


as it slopes down to groove, which is narrowed above but prac- 
tically full width at base. Thorax straight behind groove with 
slightest undulations to # where it narrows with straight edges to 
end, which is cut nearly straight and 4+ to 3 width of thorax. 

(6) Front wall straight. Caput slightly convex and distinctly 
sloping upwards from eye 2 to 4, well rounded and steeply sloping 
to a broad groove, top of which is generally only about 4 height 
of caput. Thorax rises slightly behind groove to 4 where it is 


ANT-LIKE SPID#RS FROM MALAYA, 297 


much lower than caput, and then with convex and concave 
slope to hind rim, which is weak but can be traced well towards 
caput. 

Colour mahogany, slightly darkest on caput, which has black 
eye-frame sub-broken in front of eye 4 and terminating just 
below the line of eyes 2-4. Slight indication of grey- haired 
cephalic wedge. 

Clypeus very low. 

Eyes.—\st row straight or very pape recurved by summits. 
1.1 subcontact, 1.2 considerably > Poe. 2 land ‘behind, 1b. 
2.3 very slightly < 3.4, Eye square much broader than long 
and 2.2 distinctly > 4.4 > 4-1im. 4 projects over edge of 
prosoma but not at all prominently, and the prosoma is not 
convex over it on ES view. 

Falces —Length < 3 caput = 2 breadth. Hdges parallel, trun- 
cation somewhat short and square. Groove marked: Unguis 
hight, with regular curve and oe diminution from base to 
apex, not very strong, length = paturon. Upper teeth 5 to 
angle, not quite in contact, with i or 2 dots wider spaced below. 
Lower teeth 6, as strong as upper, close, largest in centre, but no 
great difference in size. 

Maaxille.—Length = c, = falx. Narrow at base, outer edge 
curving strongly and concavely outwards beyond labium to bold 
rounded corner. Hnd square, truncation shortish, steep with 
strong hght scopula, longest at apex. Inner edge slopes outwards 
from truncation, being strongly hollowed in front of and at side 
of Jabium. Dark serrula from about middle of outer edge to 
angle. Colour light. 

Labiwm.—Length > 3 maxilla, parallel sides, cut off square 
at end, a little darker than maxilla except at end. A slight dark 
scopula at truncation. 

Palpi.—Spring from near base of maxilla. Trochanter length 
= 4 maxilla. Femur < # caput, flattened, breadth in centre 

=1 length. Patella narrow at base, broad at end. Tibia 
== 1h ae at base = width patella, at apex very much wider. 
Tarsus = 2 tibia. The tibio-tarsus plate a little > femur, with 
inner ge rather short. Colour = labium, with hairy parts 
of tibio- tars sus br wre ereen, 

Legs.—4.1.3.2, abl marked steps between each. 

Ist. c white, length < 2 breadth, very slightly tapering. 
tr white, cylindrical, length = breadth < least width of ec, 
faint. 
f light yellow, lightest at apex, dark-lined cn outer side, 
length > 2(e+tr), clubbed at base. 
p= sti, light, thin oe line each side, 1 spine pair below. 
tif; coloured as p, D spine pairs evenly spaced and about 
equal strength. 
m light, dark-lined on side, 2 spine pairs in basal half, at 
least as strong as ti spines, last reaching well on to ta. 
ta = 2 ti, light, m+ta = $(p+ti). 
Proc. Zoou. Soc.—1917, No. X XI. wil 


DW) 


ivy) 


298 MR. H. D. BADCOCK ON 


2nd.c white, length = $c, > breadth, considerably stouter at 
base. 
tr as tr,. 
f helt: slightly clubbed at base, > 2 (e+ tr). 
p = ti, light, faint dark side-lines. 
ti < f, light with dark side-lines, 3 spine pairs evenly spaced 
nearly as strong as on ti,. 
m slightly > ta, light, 2 spine pairs nearly as strong as on ti. 
ti light, ti + ta slightly < ti. 
3rd. c brown, cylindrical, > c,, length = 2 breadth. 
tr = 3 ¢, brown, length slightly > breadth, slightly wider at 
apex than base. 
f lighter brown, = f,, very slightly clubbed at base. 
p brown, light at base, < } ti. 
ti light witn dark lines at sides, p + ti > f. 
m & ta light, m+ta < p+ti. 
Ath. ¢ light with dark above and sides, length = 2 breadth, 
>c,, very slightly wider at base than apex. 
tr cylindrical, light with dark above and sides, nearly as long 
as c and much > any other tr. 
f = 2(c+tr), brown, very slightly wider at base than apex. 
p light with apex dark above, = 4 ti. 
ti light brown, p + ti > f. 
m hghter, nearly as long as ti, light. 
ta = +m, light. 

In general appearance the anterior legs are lighter than the 
posterior. 

Sternum.—Truncated between c;.; and stoutly and bluntly 
pointed between ¢),9, spreads out into an advanced point between 
€2.3, Which are rather widely separated, a long point in front of 
c4.4, Which are in subcontact. Greatest width > width of Cs, 
c:,; = nearly width of ¢,. Colour mahogany, darker than any 
other part of prosoma. (Text-fig. 6, 0.) 

Prepicet.—Slightly < caput; lst joint brown, slenderer and 
very much longer than 2nd, hight with dark sides and front, very 
short, inserted into socket on opisthosoma. 

OPpIstHosoMA = or very slightly > prosoma. Breadth and 
height nearly 3 length. Pear-shaped, widest at 4. White belt 
at about 3, distinctly in front of widest part. Lung-sacs brown, 
wide apart, lighter brown rest in front of epigastric fold, which is 
slightly procurved. General colour brownish green. 

Epigyne.—No distinct formation, but in some specimens a light 
circle with faint brown bridge ending basally in a dark bifurcate ; 
general appearance is sometimes something like the ace of clubs. 
(Text-fig. 6, d.) 

Spinners.—I|st pair stoutest. Basal joint short, thick, light 
with dark longitudinal markings; 2nd joint larger, slightly 
barrel-shaped, thinner. 2nd pair white, thinner. 3rd _ pair 
darker than 2nd, 2nd joint much longer than Ist. 

Anus.—Iwo segments, rather prominent. 


ANT-LIKE SPIDERS FROM MALAYA. 299 


3. The chief differences from 9 are :— 

Prosoma.—Caput very much higher and back wall nearly 
vertical, so that caput is about three times as high as groove and 
twice as high as thorax, standing up like a square tower (text- 
fig. 6, a). 

Hyes.—4-rim = 4.4 or very nearly. 

Falces.—Longer than prosoma, after an initial sharp con- 
striction at top they curve outwards to 2 and then in; end cut 
square but somewhat pointed forward at inner corner in con- 
tinuation of straight inner edge. On side view guise edge is 
convex, widest at. centre, lower straight. Width = + length. 
Nearly horizontal. Unguis, length slightly < paturon, S curve 
with branches much flattened but sharp curve at tip, considerably 
thicker and darker at base than tip. Groove not defined. Upper 
margin contains 7 teeth, of which the first 2 are close to base of 
margin, the 4 at least are strong and subevenly spaced, the last a 
little weaker. Lower margin 6 or 7 teeth, very small, and much 
further apart at apical than basal end, but teeth a little variable. 

Palpi.—Tibia only a little larger than patella, with moderately 
strong pointed black-tipped process on upper outer apex. Tarsus, 
has fovea large and extending nearly to the end, with a fine spine 
encircling rather more than once. 

Legs.—The contrasts of colouring are not nearly so distinct 
‘as In 2, 

OpistHosomMA.— White belt missing. Upper and centre of 
lower sides coriaceous. 


SHort DESCRIPLION. 


Prosoma = opisthosoma. Caput = thorax. Caput very high 
with nearly vertical walls, tower-like in ¢, not so marked in °. 
Occasionally faint white-hsired cephalic wedge. 

2 falx. Upper teeth 5 to angle with 1 or 2 dots beyond; 
Jower teeth 6, as strong as upper, close. 

$ falx. Longer than prosoma, width + length, outer edge 
convexly curved on upper and side aspect. Upper teeth 6 or 7, of 
which first two comparatively close to root of unguis. Lower 6 
dots wider spaced at root of unguis. Unguis slightly < paturon, 
flat S curve with rather sharp curve at point. 

Orpalp. Tarsus =, 2 tibia. 

$ palp. Tibia has moderate, black-pointed, slightly curved 
process at outer upper apex. Organs surrounded by circle and a 
bit of thin black spine. 

Legs 4.1.3.2, with marked difference between each 2; anterior 
pairs lighter than posterior, bnt not always in 6, dark side-lines 
in some joints. p, 1 spine pair; ti, 5 spine pairs, evenly spaced, 
not very strong; a 2 spine pairs; ti, 3 spine pairs nearly as 
strong as on ti,; m, 2 spine pairs nearly as strong as on ti,. 

Pedicel nearly as long as caput, 2nd joint very short as 


compared with Ist. 
Pil 


300 MR. H. D. BADCOCK ON 


Opisthosoma. Fusiform, not constricted. White belt at about. 

2 in 93 missingsim, oC . 

General colour of prosoma lightish mahogany with black eye- 
frame, of opisthosoma dark greeny brown. 

Note. —The description is based on 12 2 and 8 Q, all from 
Bukit Besar and Talum. Nos. 93 2 and 94, 95 ¢ were all in 


same tube, and this fact as well as the general resemblance is. 


evidence of identity of species. 
The followi ing measurements were taken :— 


250mm. ps. 2°3x1:2X1-2, 0.8. 24% 14% 1-2, 

iesss le e+, et+tr+f 17, p+til5, m-+tal:l ee hs 
De eace iy leOe pt+titm+ta 2°5 =oo 
Oise obe 1-4, p+til-0, m-+ta missing 
Ai ios e+tr 0: Os fal 8, oi alee or allel ='5-7 
Gg OFZ mms p.s. 2°4 x 1:4 x 2:0} 2°4., 

Legs. 1... e+tr+f2:2, p+ti 2°4, meta 18 
25 ie ': 136; ee ale8, 5 heeled 
San 2k, 5, 188, 5 mellem 
2 eS e+tr Tl, f 2-0, ees oe yy bee 

The lengths of the specimens varied from 5:2 to 3°7 mm., the 


3 being on the average slightly longer than @. 
Ants ‘accompanying, nos. 4, 14, 15, 1% 


Q. MYRMARACHNE CUNEATA, sp. n. (Text-fig. 7.) 


Prosoma. — (a) Length = p,+tr,+m,+ta, nearly, slightly 
> opisthosoma. Caput = thorax. Front wall slightly projecting 
in centre and stepped back to eyes 2. Side walls straight, very 
slightly approaching backwards. Back wall strongly projecting 
at centre into bridge. Bridge narrowed at top, only very slightly 
at base, though it. is liable 1 to appear so owing to white wedge. 
Thorax edges « curve very slightly outwards to 4, then inward to 
rear, which is cut off convexly straight. 

(b) Front wall vertical, top of caput straight and level, groove 
shallow and broad, bottom not below eye 4. Thorax rises behind 
to above half way up eye 4 at 4and then slopes convexly back- 
~ wards to rim, the final concavity before rim being negligible. 
Rim rather marked and traceable round to caput. 

Colour medium mahogany, lightest at sides and much darkest. 
at top of caput. Eye-frame black, rather broader than usual and 
very distinctly broken between eyes 3 and 4, A very strong 
white wedge on either side of bridge, smooth and not due to 
hairs. 

Eyes.—\st vow straight by summits, 1.1 and 1.2 subcontact, 
2 slightly behind 1. 2=431. Eye square slightly broader than 
long and sides‘slightly approaching backwards. 2.3 very slightly 
< 3.4,.4= or very slightly > 2. 4 projects but not markedly 
so, and 4-rim very much < 4.4. 

Faices.—Project nearly horizontally forward by lower edge, 


‘ 


ANT-LIKE SPIDERS FROM MALAYA, 301 


visible from above, length > 3 caput, upper edge strongly convex 
being subangular at 3, length = 2 breadth. From above edges 
subparallel, outer being slightly convex. Upper teeth 6, 1 basal 
close to root of unguis, small and liable to be missed; 2.3.4.5 
largest with increasing spaces, 6 beyond angle very small. Lower 
teeth 6, strong, close, from base of unguis to level with 5 of 
upper. Thickish light upper scopula longest at angle; thin dark 
lower. Unguis rather strong. Colour hghter than cephalo- 
thorax, glabrous. . 
Text-figure 7. 


Myrmarachne cuneata, sp.n., 2. 


a. Profile. 6. Epigyne. 


Maxille.—Length =c,+tr, =falx nearly. Outer edges parallel 
to end of labium, then inclined straight outwards to rounded 
outer corner. Serrula at end only, which is slightly rouffded. 
‘Truncation long, very oblique, with very strong black scopula. 
Below scopula inner edge hollowed in front and at side of labium. 
Colour light. 
 Labtum.—Length > 2 breadth, parallel sides, slightly con- 
stricted at middle; strong dark scopula. 

Palpi.—Spring from near base of maxilla. Tr = width 
maxilla; f = 4 caput, parallel sides, rounded ends, flattened, 
length < 3 breadth; p = 4 f, broadening to apex from narrow 
base. Ti < 2p, broadens from base to apex, which is 2 width p 
apex. Ta very slightly > ti, rounded end; the two together 
making a plate =2f, hairy on both edges. Total length of 
palpus when extended =2 falx. 

Legs.—Order 4.1.3.2, with all differences marked. 

Ist. c white, length = 24 width, parallel or very slightly 
tapering. ; 
tr white, with sometimes dark patch on outer side ; breadth 
— length = 7¢, 


302 MR. H. D. BADCOCK ON 


f le, strong black side-lines, clubbed at base, = ti. 
p= L ti, white with dark side- lines, 1 spine pair. 
ti white with dark side-lines, 7 spine pairs, or 7 and 6, last 
3 or 4 pairs all terminating about level at end of joint. 
m light, > p, 2 spine pairs, of which last reaches to middle 
of ta. 
ta light = 3m. 
2nd. ¢ brown, strongly swollen at centre, length = breadth. 
tr broader than long, brown except for patch of white on 
fore side. 
Other joints, proportions, and colours as Ist, but are slightly 
greater in proportion, and ta only slightly < m. 
ti has 3 spine pairs, -m 2 pairs. 
ord. c a little > c,, brown. 
tr short, cylindrical, brown. 
f > ti < p+ti, brown, very slightly clubbed at base. 
p = 7 ti, curved downward, light except above apical end. 
ti brown, slender, slightly lighter at end. 
m nearly = ti, light, darker shading at basal sides. 
ta = 4m, light. 
4th. ec < c, brown except for light below. 
tr much longest, white with dark basal side-markings, 
cylindrical, > c. : 
f brown, slightly clubbed. 
p white except for brown above at base and apex, =j ti. 
ti brown, lightest at apex. 
m brown, nearly = ti. 
ta white, < 3m. 


The light ¢, and tr, and tr, are very noticeable, as also the dark 
side-lines on the light anterior legs. 

eee .—Blunt-pointed behind ¢;.;, which are about } their 
width apart, and between c2.3. Wider and pointed close behind 
c,, attempt at a point at c,, and then tapers to long point in 
fr ont of c4,4, which are in contact. At widest point =c,. Colour 
dark mahogany. 

PrepiceLt.—Short = 4 caput, rather stout. Joints about equal, 
first very slightly longer. 

OpistHosoma.—Middle thickest, constricted at +. Brownish 
green with white transverse band at constriction, somewhat 
lighter below, with longitudinal central portion bordered by faint 
white lines from epigyne to spinners. Lung-sacs dark, wide 
apart. Epigastric fold procurved. 

- Kpigyne.—A lighter semicircle in front of epigastric fold, in 
the centre of which is a dark marking somewhat houtrglass-shape 
(text-fig. 7, 6). 

Spinners. —1lst brown, thickness slightly eveater at base than 
apex. 2nd longer, thinnest, lighter, cylindrical. 3rd as light as 
2nd, slightly thicker, subeylindrical. 


ANT-LIKE SPIDERS FROM MALAYA. 303 


SHort DESCRIPTION. 


Prosoma very slight, longer than opisthosoma. Cephalic groove 
broad and shallow, caput not very much higher than thorax. <A 
strong white wedge not due to hairs at sides of groove. 

Falces nearly horizontal by lower edge. Upper teeth 6, with 
4 middles much largest. Lower teeth 6, strong, close. Ta of 
palp = or very slightly > ti. 

Legs 4.1.3.2, with all differences marked. Ist has white c 
and tr, and 4th white tr larger than c. The anterior legs are 
light with strong dark side-lines f to ti. p, has 1 spine pair, ti, 
7 pairs or 6.7, m, 2 pairs. Ti, has 3 spine pairs, m, 2. 

Pedicel shortish, with j, very slightly > j.,. 

Opisthosoma has thin white belt at 7 and is widest at % back. 

Notes.—Description based on 4 @ found at Bukit Besar 
3500 ft., Talum 4000 ft., and Gedong. One of the specimens only 
(from Bukit Besar) is accompanied by its mimicked ant of species 
no. 2. The type-specimen is rather lighter than any of the 
others. Differences from specimen to specimen are very slight. 
In one tr, has a distinct dark exterior marking not visible in the 
others. The sternum point is rather unusually far from the base 
of the labium. The cephalic wedge is always very noticeable and 
serves as a recognition mark, 

This species is in many ways very like Wyrmarachne macillosa, 
bunt it may be distinguished without difficulty by general shape of 
prosoma and opisthosoma, which are less squat, teeth of lower 
margin, white cephalic wedge, larger pedicel, and shape of 
sternum, which is pointed instead of undulated between coxze and 
constriction of opisthosoma. 

Total lengths 7:3, 7°6, 7:3, 6-2 mm. 

Legs of specimen 6°2 mm. long :— 


tise e+tr 1:0, £1:9, p+ti2°3, m+tal-l] =6:3 

Ditch e+tr+f 1:8, Repel ee me Oa ail 

Basen: e ra eae ye ek pO 

1 eee e+tr 11, f 2-1, Os Ty eo) ely ae 
Ants accompanying, no. 2. 


3S. MyrMARACHNE RAMOSA, sp. n. (Text-fig. 8.) 


Prosoma.—(a@) Length = p,+ti,+m,-+ta, nearly, slightly > 
opisthosoma. Caput=thorax. Front wall recurved, not stepped 
back to eye 2. Sides straight or very slightly convex, parallel. 
Back wall straight with well rounded corners. Thorax joined 
across groove by distinct paralled-sided bridge which appears 
from above as a sharp constriction, though groove at base is 
nearly as wide as caput. Thorax swells out at sides to about 3, 
and then narrows with slightly convex and concave slopes to end 
which is cut square. 

(6) Front wall of caput nearly vertical but with very slight for- 
ward batter, top straight sloping very slightly up to eye 4, over 


304 MR. H. D. BADCOCK ON 


which it is rounded. Back wall slopes straight down to narrow 
groove, height = 4 caput. Thorax slopes slightly up to z and 
then convexly and concavely to end where is rim, which can be 
traced round to caput. 

Colour mahogany, darkest on caput. Black eye-frame widely 
broken between 3 and 4. Faint light-haired wedges at sides of 
groove. f 

Hyes.—I|st row slightly procurved by summits. 2=21. 1.1 
an appreciable gap, 1.2 slightly wider. Eye square markedly. 
wider than long and very slightly. wider at 4.4 than 2.2. 2.3 
distinctly <3.4, 3 below upper tangent to 2.4, 4>2 and lighter 
in colour, projects over edges. 4-rim < 4.4. 


Text-figure 8. 


S > SS! 
ce SS 
— 
Die 


Myrmarachne ramosa, sp. n., 3. 


a. Profile. 6. Sternum &c. c. Right palpus from below. 


Falces.—-Longer than caput < prosoma. Length = 3 width. 
Inner edge straight with black ridge, outer convex with black . 
ridge. End hollowed, with onter angle sharp, inner chamfered. 
On side view much deepest just beyond base, decreases consider- 
ably to apex, upper edge being straight and lower slightly hollow, 
Unguis long and tip reaching beyond end of maxille, flat S curve, 
with short branch or knob at 3. Groove non-existent and upper 
and lower teeth liable to be mixed up. Upper teeth 9, of which 
that nearest to unguis is strongest, the next 5 are spread over the 
paturon with gap in centre largest, last teeth closer, smaller, and 
overlapping maxille. Lower teeth about 10, small, spread over 


ANT-LIKE SPIDERS FROM MALAYA. 305 


length of paturon but closer at apical end. The two rows of 
teeth are notably crooked (text-fig. 8, 6). 

Maxille.—Length =c,+tr,. Outer edges parallel to end of 
labium, then inclined outwards to well rounded corner and 
slightly convex with truncation 45°; strong dark scopula. Below 
scopula inner edge hollowed in front and at sides of labium. 
Serrula. Colour light mahogany. 

Labium.—Length > 2 breadth, parallel sides, broadly and very 
slightly constricted near apex, end slightly hollowed, dark scopula. 
Dark mahogany, light at apex. 

Palpi.—Spring from near base of maxilla. Tr > width 
maxilla. f = 4 caput, parallel sides, somewhat reduced at ends 
and slightly flattened. p < tr < $f, broadening to apex from 
narrow base. Ti only a little > p above, = nearly 2p below, 
slightly broadened from base to apex; short, small, blunt, light 
straight process at outer lower apex, largish hairs on inner edge. 
Ta a little > ti, blunt, suboblong. Fovea occupies little more 
than basal half. Organs have a fine black spine springing from 
outer basal rim of fovea passing backwards and inwards with 
double spiral, and finally pointing forward rather on inner apical 
rim (text-fig. 8, c). 

Legs.—4.1.3.2, with difference between 3 and 2 only slight. 
Ist. ¢ length > 2 breadth, rather markedly tapering, brown. 

tr narrower, slightly tapering, < 3c, brown. 
f clubbed at base, = ti, brown, darkest above. 
p = 3 ti, slight, curved, light, shaded at sides. 
ti 4 spine pairs, with sometimes a fifth very small, light, 
dark at sides. 
m= 13 p, 2 spine pairs near base and apex, basal much 
largest. 
ta < 3m, dark. 
2nd.c¢ much shorter, NON at base, lighter. 
tr cylindrical, < 4c, light with shaded sides. 
fi OnLy 2 little thicker at base than apex, lighter, but 
side-lines darker especially fore. 
-p=nearly 3 ti, strong especially at apex, slightly curved, 
light. 
ti light, 3 spine pairs. 
ls Ps 2 spine pairs, Rep: 
ta < $m, light. 
SLGeAC Or ' as 2nd, but dark all over. pp, ti, m, ta slightly 
lighter than these. p = 3 ti nearly, m = ti, ta < 4m. 
4th. c =c,, cylindrical, very slight tapering. 
tr = c, cylindrical, both ¢ and tr lighter than 3rd and dark 
at sides. 
f brown, nearly = p + ti. 
p > ¢ ti, light, with dark apex above and at sides. 
ti, bro a ti nearly, slenderer, and back lighter at apex. 
a < im, ‘light. 


306 MR. H. D. BADCOCK ON 


On the whole the legs strike one as dark, none of the coxe or 
trochanters being really hght. 

Sternum.——Blunt-pointed between c;.1, which are about 4 width 
apart, blunt-pointed between c,.2, which are not in contact, 
swells out with straight edges between c2,3, which are wide, and 
tapers to moderately long point in front of 4.4, which are in 
contact. Widest =c,. Dark mahogany as coxe. 

PEDICEL nearly equals caput. j, brown, longer than j,, below 
=2j,.  j, light with black fore end. 

OPISTHOSOMA = prosoma. Thin and round with broad con- 
striction at ¢. Brown above and below in front of epigastric fold, 
which is slightly recurved. General colour very dark. 

Spinners.— 1st light, stoutish, barrel-shaped. 2nd lige 
slenderer, cylindrical. 3rd longest, darker, slender, oy nieneal 

Anal tubercle. —Prominent, 2-jointed. 


SHort DESCRIPTION. 


Prosoma = or slightly < opisthosoma. Cephalic groove rather 
narrow, with thorax about half height of caput and wedge of 
white hairs not very strong. 

3 falx longer than caput < prosoma, flattened above. Unguis 
as long as paturon, flat S curved, with very short knob-like 
branch. Upper teeth 9, lower 10, but lines liable to be confused. 

3 palp = or nearly = falx. Tibia has short blunt process at 
outer lower apex. Organs have a spine twisted spirally twice 
and terminating pone straight forward at inner apex. 

Legs 4.1.3.2 with difference 3.2 slight, no very marked 
colour contrasts. Tr, 4 spine pairs, m, 2 pairs, ti, 3 pairs, m, 
2 pairs. 

Pedicel nearly = caput, with lst joint markedly greater than 
2nd, especially below. 

Opisthosoma widely constricted at 4, with longitudinal brown 
coriaceous marks above and below and in front of epigastric fold. 

Note.—Description is based on 4 ¢ from Bukit Besar and 
Biserat, two of the specimens from the latter associated with ' 
ants nos. 9 and 16 respectively. There is not much difference 
between specimens except some variation in intensity of leg 
colouring. 

Measurements :— 

Total lengths 5:0, 5:2, 5:4, 5°8 mm. ex falces. 

Legs of specimen 5:2 mm. long :— 


LS acess e+tr0°8, £1:°2, p+til9, m+tal-2 =51 
ee yO 1Os aise Bes Bais ces » 10 Sao 
Sees you 2a bee tae a iw lice, oo 
Aes Bede le al AY ne ee). a ES a9, 215 


Q. MyRMARACHNE ALBICRURATA, Sp. n. (Text-fig. 9.) 


Prosoma.— Length = opisthosoma = p,+tr,+m,; Greatest 
breadth distinctly > greatest height. Gaput = or slightly < 


ANT-LIKE SPIDERS FROM MALAYA, 307 


thorax. Front very slightly recurved, slightly steppedgback to 
eyes 2. Side walls very slightly convex and tapering outwards 
towards eyes 4, Back wall strongly procurved at groove, which 
is strongly constricted above and much less so below. Thorax 
curved rather strongly outwards from groove to slightly in front 
of centre and then inwards to rear, which is cut nearly square. 
Rim dark. 

Caput somewhat convex and higher behind than before. Slopes 
at moderate angle behind, with groove low and < half height of 
caput and broad. Thorax rises to about 3 and then falls convexo- 
concavely to turned-up but not abrupt rim, which is dark and 
traceable round to caput. <A slight white wedge at thoracic 
groove. The general shape is well rounded at angles. 

Colour light mahogany, with black eye-frame broken in front 
of 4 and with lower boundary not so well defined as usual. 


Text-figure 9. 


Myrmarachne albicrurata, sp.n., 2. M. lateralis, sp. n., 2. 


a. Sternum &c. of WM. albicrurata. b. Do. of M. lateralis. 
ce. Epigyne of IZ. lateralis. 


Hyes.—\st row straight, intervals 1.1 and 1.2 slight and 
about equal. Kye square broader than long, parallel sides. 
3 midway between 2and 4. 4>2. 4.4 distinctly > 4-rim. 

Faices.—Length = § caput, nearly parallel with inner edge, 
very slightly convex, length = 24 breadth. Side view strongly 
convex or rather angular at nearly 4, base almost horizontal. 
Groove marked, both outer and inner scopule lighter and thinner 
than usual. Upper teeth 8 or 9, slightly irregular in size and 
spacing; lower 8 or 9 closer, especially away from root of unguis. 


308 MR. H. D. BADCOCK ON 


Maxille = cornuta, very light in colour; scopula dark, strong. 

Labium = cornuta, very slightly darker than maxille; scopula 
light, strong. 

Palpi.cSpring from near base of maxilla. Length of tro- 
chanter = narrowest width maxilla. Femur < 3 caput, flattened . 
with rounded ends, of which basal is slenderer. Patella about = 
trochanter. Tibia slightly < tarsus. Usual shape, but tarsus 
slightly more pointed and hairs even on inner edge rather scanty. 
Joints to femur white, patella to end much lighter than usual. 

Legs.—4.1.3.2 with 3 nearly = 1. 


Ist. c cylindrical, length = 2 breadth, white. 

tr length < width c, narrower, white with sight brown 
longitudinal marks on inner side. 

f nearly = caput, shghtly widened at base but not clubbed, 
white with dark longitudinal side-marking near basal half 
of ous side. 

p = 3 ti, light with dark side-lines, no spines. 

ti te 4 spine pairs moderate, evenly spaced and subequal, 
but 3rd rather the strongest, white with dark side-lines. 

m = p, 2 spine pairs of which first is as strong or stronger 
than 3rd of ti, white. 

ta very sightly < m, white. 


2nd.c length =} c, = breadth, gibbous, white. 
tr length = 3c = breadth, white. 
f < f,, white. 
p > ti, white. 
ti 2 spine pairs, p+ti = or < f, very slight dark side-line, 
white. ; 
m 2 spine pairs, white. 
ta very slightly < m, white, m+ta = pt ti. 
3rd. ¢ shehtly > ¢,, brown, less gibbous. 
tr > s¢, oe davies at sides. 
{> f , slightly broader near base, brown. 
bo ‘ti, light with brown above distally. 
ti light with dark sides most marked at base, p+ti slightly 
Sk 
m slightly > ti, white. 
ta < m, m+ta distinctly > p+ti. 
Ath. c distinctly > c¢,, with same proportions, light with dark 
sides. 
tr nearly = c, slenderer, white with dark markings at side, 
of which fore much the stronger. 
f long, sightly clubbed, brown. 
eae ti, white, brown above distally. 
ti brown except for slight white continuation of p below. 
p+ti distinctly > f. 
m long, light except for slight darkening at basal sides in 
continuation of ti. 
ta = 3m and very slightly darker. 


AN'tT-LIKE SPIDERS FROM MALAYA. 309 


Sternum.—Short-pointed between c;,1;, which are < 7 width 
apart, and between cj .2, extends to sides between c2_3, which are 
very wide apart, contracted between c3,4, and the point, which is 
not well marked, is met between cy,4, which are not quite in 
contact, by a very fine-pointed projection from the fore Jorum of 
the pedicel. Greatest width =c,. Brown. 

PEDICEL = 4 caput. j, >jJ,, very much so below; above the 
fore lorum is slightly broader and longer than the second, below 
“much stronger and broader and extended forward into a very long 
fine point. 

OpistHosoMA.—Subcylindrical, length = 3 or 4 breadth, broadly 
but not deeply constricted at 3. Dark green, with faint white 
oblique markings at constriction and suggestion of white mark- 
ings below. Lung-sacs brown and wide apart. Epigastric fold 
recurved, very clear cut. 

Epigyne.—Immature. 

Spinners.—1st much thickest, barrel-shaped, dark. 2nd about 
same length, very thin, white. 3rd about same length, very 
shghtly thicker than 2nd, darkest. 

Anal tubercle.—Moderate. 


SHORT DESCRIPTION. 


Prosoma = opisthosoma, Groove rather deep but broad, and 
thorax not rising much behind it. Constriction appears rather 
sharp from above, cephalic wedges slight and with a very few 
white hairs, 

Ta of palp a little > ti. 

Legs 4.1.3.2 with 1 and 3 nearly equal. The anterior legs 
are almost white as are the coxe of 4. tr, as long as c, but 
thinner, white with dark sides. c¢, greatest coxa. p, no spine, 
ti, 4 spine pairs, ti, 2 pairs. The backward point of sternum 
met by a very long and narrow point reaching forward from 
under side of pedicel. Legs 1 about 7 their width apart; legs 4 
not quite in contact. 

Pedicel = { caput. j,> ],- 

Opisthosoma subcylindrical, broad but shallow constriction 
at 3, with faint oblique white markings sometimes evanescent. 

WVote.—Description based on 3 immature 9 from Ban Sai Kau, 
Nandock, and K. Mahek, Jalor, only one of which was in good 
condition. 

Measurements 5:7, 5-6, and 4°5 mm. 

Legs of specimen 4°5 mm. long :— 


Tere, e+tr+fl-l, p+til4, m-+talO =3°5 
ES lia +3 TOF es 4 es: 
2 eee A a 1 i pra clclle 4) Pd .=3:3 
Ae e+tr 1:0, f£0°9, a alanis a Or © S01 


No ants accompanying. : 
It is quite possible that, this species and the following one may 


310 MR. H. D. BADCOCK ON 


be identical; the grounds on which I keep them apart provision- 
ally are 
(1) Shape of sternum (text-fig. 9, a, b).—-In MW. lateralis legs 2 
are as close to one another as legs 1; in IM. albicrurata 
they are normal. 
(2) Spine pairs.—alhicrurata, p, no spines, ti, 4 prs., ti, 2 prs. 
lateralis, p, 1 spine, ti, 5 prs., ti, 3 prs. 
(3) albicrurata light in colour, lateralis dark. 


Of two specimens, however, which were much damaged and 
cannot be certainly identified, I found in the first (collector’s 
number 4) agreement with albicrurata in (1) and (3) and with 
lateralis in (2); while the other (collector's number 92 A) agrees 
with albicrurata in (1) and (2) and with lateralis in (3). 


2. MyRMARACHNE LATERALIS, sp. n. (Text-fig. 9.) 


Prosoma.—Length < opisthosoma = ti,+m,. Caput = thorax. 
Caput hardly stepped back at eyes 2, well rounded at rear corners. 
Bridge rather broad at top and only very slightly constricted 
below. Thorax spreads out behind to widest at 3, well rounded. 

Groove shallow but broad, with rim thin. ‘Thorax considerably 
lower than caput. 

Colour dark, almost black on caput, strong wedge of white hairs. 

Hyes.—1st row procurved, intervals slight but perceptible. 
1=3 times 2. 2.3=3.4, 3 very small and rather above inner 
tangent to 2.4. 4=2,. 4.4> 4-rim. Eyes 2 and 4 rather 
pr ominent. 

Falces.—Length = 3 caput, nearly horizontal, sides subparallel, 
slightly convex outside at distal end. On side view straight below, 
strongly convex above, with angle at 3 to 4. Teeth 7 or "8, largest 
at centre, falling in size both ways. Lower 8 or 9 close, largest 
at centre, smaller than upper. Moderate greyish outer scopula. 
Inner a few dark hairs. 

Mawxille, labiuwm = cornuta. 

Palpi.Spring from base of maxilla. Tr = width of maxilla 
at narrowest; femur flattened, < 3 caput, reduced and rounded at 
ends, greatest width = 3 length. p very slightly larger than tr, 
expands distally. Ti and ta form flat plate, hairy especially on 
inner edge. Ta > f > ti, but differences not great. 

Legs.-—4.1.3.2. 

Ist. ¢ claviform, length < 2 breadth, brawn. 

tr < width of ¢, light with strong dark side-markings. 

f long, claviform, light with dark side-marks very broad at 
tee 

DiSes 1 ti, ight with dark side-marks, | spine below at 2 

biel; light with dark side-marks ; spines 5 pairs, or 5 fore 
and 6 ae. the 3rd considerably strongest. 

in much < ti and darker, 2 spine pairs of which 1st is very 
strong. 

ta = 2m, dark at base, light at tip. 


ip) 
4 


ANT-LIKE SPIDERS FROM MALAYA. By al 


2nd.c white, < ¢,, gibbous. 
tr white, <4. 
f white. 
Other joints marked as Ist but much lighter. 
= iti, ti 3 pairs of spines, m> 4 ti 2 prs., ta > 3 ti. 
3rd. c & tr considerably stronger than in 2nd, dark brown. 
f slightly clubbed, dark brown. 
p brown but very slightly lighter, = } ti, a long hair above 
at distal end. 
ti dark brown, stout, < f. 
m slightly > ti and slenderer, dark at base, lighter toward 
apex. 
ta heht,. > = m. 
Ath. ¢ largest, light centre strip below, otherwise dark. 
tr light with dark sides, at any rate apically, cylindrical, =c. 
f long and dark, very slightly clubbed at base. 
p light except for apex above, = j ti. 
tiecdank. < 4. 
m dark, slightly < ti and thinner. 
a = +1, lighter. 


Sternwm.—Short-pointed between 1.1, which are very close, 
and between 2.2, very strongly contracted between 2.2, which 
are very close, swells out with boss between 2.3, which are well 
separated. ‘Tapers between 3.3 to a point in front of 4.4, which 
ave as far apart as 1.1. Greatest width =c,. Dark brown. 

PEDICEL = caput. j, much > j.. ‘ 

OpistHosomA.—Fusiform, slightly constricted at 3, widest a 
little behind middle. Dark greenish, with thin oblique white 
markings at sides of constriction extending well back below. 
Epigastric fold straight to procurved ; lung-sacs wide and region 
between them hairy. 

Epigyne.—A dark horseshoe enclosing a central fork. (Text- 
tig. 9). ¢:) . 

Spinners.—I|st barrel-shaped, dark. 2nd lighter, longer, 
thinner. 3rd cylindrical, < 2nd, darkest. 

Anal tubercle.—Two-jointed, not very marked. 


SHorRtT DESCRIPTION. 


Prosoma well rounded, with broad shallow groove and caput 
considerably higher than thorax. Cephalic wedge of strong white 
hairs but not very broad. 

Falces nearly horizontal, with teeth upper 7 to 8 and lower 8 
or 9, the latter close and much smaller than the upper. 

Legs 4.1.3.2, with 4 much the longest. p, has a single spine, 
ti, 5 pairs on 5 and 6, m, 2 prs., ti, 3 prs., m, 2 prs. 2nd legs 
white, lst white except coxe, 

Sternum well rounded, both e2.3, ¢1.1 very close, more so than 
Ci 4, €2.8 wide. 

Pedicel = caput. j, much > j,. 


BHO: MR. H. D. BADCOCK ON 


Opisthosoma = prosoma, fusiform, constricted at 7 and with 
oblique narrow white stripes commencing at sides of constriction. 

Note.— Description based on 2 2 from Bukit Besar and 
Sungkei. Lengths 6°2, 6°3 mm. 

The following measurements are from the specimen 6-2 mm. in 
total length :— 


lepers e+ti0°9, £1:2, p+til'8, m+tall =50 
rey oi 0: eS Bee) oa ae jd a Rees 
ee ee poe eee |). le 5 iA 
A ae oy elo Lgataee ella or Pre eh 350 | 20} eel 


No ants accompanying. 


3. MyRMARACHNE BISERATENSIS, Sp. n. (Text-fig. 10.) 

Prosoma.—(a@) Length slightly > opisthosoma = p, + ti, 
+m,+ta,. Greatest height and greatest breadth each = 4 cach: 
Caput = ‘thorax. Fr ont wall very slightly recurved and stepped 
back at eye 2. Side walls strongly convex. Groove narrow and 
slightly recurved. Thorax well rounded at sides and terminated 
with almost square end of % full width. Rim strong. 

(6) Top of caput flat. Chinon narrow and-shallow. Thorax 
rises slightly behind it and then falls with strongly convex slope 
to end with strong concave rim. 


Text-figure 10. 


Myrmarachne biseratensis, sp.n., 3. 


a. Profile. 06. Palpus. c. Sternum &c. 


Dark, especially on caput, but eye-frame traceable, black. 

Eyes.—\st row straight by summits, 1.1 very close, 1 . 2 close. | 
2— 31, 3 slightly nearer 2 than 4 and on upper tangent, 
4 hehe > 2. Hye square broader than eee 4.4 very slightly 
> 2.2 and about = 4rim. 


ANT-LIKE SPIDERS FROM MALAYA. ole 


Falces = cephalothorax, length = 5 breadth, straight inner 
edge, outer slightly convex, broadest at %. End concavely trun- 
cated and facing slightly outwards so as to make inner end rather 
pointed but not projecting. Upper edge at first flat, then rises 
with strong convexity so as: to make it much higher at §. Lower 
edge at first flat, then a convex step downward and flat to end. 
Lighter colour and stippled. 

Unguis not as long as falx, reaching only to end of flat, doubly 
curved and with distinct prominence (not, however, to be called 
a branch) at 4. Groove rather distinct, but not deep. Upper 
teeth 4, 1 at base of unguis strong but very close and liable to 
be missed, perhaps rather a process at end of falx; the other 3 
equally spaced along 2nd basal quarter. Lower teeth 15, evenly 
spaced, stronger than usual and, at any rate, some of them 
alternate in size. 

Masxille.—Shape as maxillosa. Scopula very strong, but grey. 
Outer portion dark, inner equally wide, light. 

Labium.— Proportions and colouring as masillosa.. Slightly 
constricted in middle, parallel sides, and apex concave with dark 
scopula. 

Palpi.Spring from near base of maxilla. Tr > width maxilla, 
dark sides, light. below; f = 2 tr, dark, with grey hairs slightly 
curved ; p < tr, parallel sides. Ti nearly = f, short process at 
fore, outer, lower corner, widens apically. Ta a little > ti, 
oblong. Both ti and ta are hairy, with a strong fringe on inner 
edge. Tita below. Fovea occupies a little more than basal half 
of ta, and is surrounded by a dark hairy edge. A spine springs 
from basal inner edge, coils up inner side round 12 times and 
terminates in a fine point facing forwards. Style long and looped 
backwards from bulb in front. 

| Legs.— 4.1.3.2, with differences more or less equal. 
Ist. c ovate, lighter than falces. 

tr < width e¢, lighter with dark at sides. 

f clubbed, dark, long. 

p curved = 4 ti, no spines, dark. 

ti dark, spines difficult, but apparently there have been 
5 pairs all weak, of which first two are basal and medial 
and last three fairly close at apical end. 

m = # ti, dark, 2 spine pairs much stronger than ti. 

ta = 3m, dark, 

2nd.c¢ a little longer than broad. 

tr short, dark sides. 

f dark, more clubbed than f,. 

p = 3 ti, curved, dark at apex above and sides, no spines. 

ti dark at sides, 2 spine pairs. 

m nearly = 3 ti, light except for dark marks on sides at 
base, 2 spine pairs. 

vas zm, hight. 

The tibial spines of 2nd are very strong compared with 
those of Ist. | 


Proc. Zoou, Soc.—1917, No, XXII. 22 


aod 


314 MR. H. D. BADCOCK ON 


3rd. ¢, tr, f, dark and all a little longer and stronger than 2nd. 
p == ulboth dark, 
m nearly = ti, light. 
ta = 3m, light. 
Ath. ¢ < c,, stout, length = 14 greatest breadth, dark. 
tr > c, much longest tr, light with dark sides. 
f long, clubbed, dark. 
p= 1 ti, dark. 
tr dark, cylindrical. 
m = ti nearly. 
ta = +m, light. 


Sternum.—Truncated between c,.1, which are about 4+ width 
apart, swells out between other c, especially between c2,3, which 
are only a little wider than the others, tapers with undulating 
sides to a sharp but rather short point in front of c4.4, which are 
in sub-contact. | 

PepiceL.—Short but distinctly visible from above. j, as seen 
from above distinctly longer than j,. 1st upper lorum decreases 
backwards, where it is procurved, much broader than 2nd. 
Whole length of pedicel < 2 greatest width. Itis received into 
a short socket projecting from opisthosoma. 

OpistHosomA.—Egeg-shaped, widest about §.° Above general 
colour dark greenish brown, below somewhat light, the front of 
epigastric fold yellowish, corneous, behind a longitudinal lighter 
band to spinners, widest at fore end, whence it reduces suddenly 
and then parallel. 

Spinners. — 1st stout, subconical. 2nd longest, lightest, 
thinnest, cylindrical. 3rd very slightly stouter, shorter, and 
darker than 2nd. All spring from projections of abdomen and 
are 2-jointed, with 2nd joint very short. 

Anus on a strong tubercle. 


SHORT DESCRIPTION. 


Prosoma very slightly > opisthosoma. Caput = thorax, 
Groove narrow and shallow. Thorax well rounded behind and 
only slightly lower than caput. Sides ef caput convex. Falces = 
cephalothorax, subparallel, on side view much thicker towards 
apex. Unguis slightly < paturon and with distinct prominence. 
Teeth, upper 4 all in basal half and are close to base of unguis ; 
lower 15 partly alternate in size. 

Palpi.—Tibia = tarsus. Fovea only occupies basal half of 
tarsus, a spine springing from basal inner edge coils 13 round and 
terminates pointing forward. 

Legs 4.1.3.2, with moderately equal intervals. Spines of 
ti, noticeably weak but probably 5 pairs, m, 2 pairs, tr, 2 pairs, 
very strong for 2nd leg, m, 2 pairs. tr, and tr, are distinctly 
lighter than other joints. 

Sternum rather broad. 2nd and 3rd coxe only a very little 
further apart than the others. 


aud 


ANT-LIKE SPIDERS FROM MALAYA. 315 


Pedicel shortish. 

Opisthosoma, as whole spider, rather squat. 

Note.—The description is based on a single ¢ from Biserat, 
length 6°2 mm. Its general shape reminds one of M. mazil- 
losa 9. | 


The following are measurements of the legs :— 
ila pttr 13, £1°9, p+ti3°0, m+tal‘9 =81 


Riaaee MPC TeOn Gee yee 1) oe eis! 22557 
Be, Pie iG 8) 9:0. 6-7 
At ers ok OG, , 26 =89 


This spider was found in company with ants nos. 2 and 17. 
It is very probably a do of MM. maxillosa, which it strongly 
resembles in general shape and colouring, all its features being 
those of maxillosa exaggerated even to the falces. The only 
distinct difference is in the spine pairs. It is the only ¢ in the . 
~ collection the least like the 9 mawillosa, which is so common, 
and I should not have hesitated to claim it as the ¢ did it not 
differ so strongly as regards teeth and tibial spines from Thorell’s 
3g M. (Toxeus) maxillosa. It was found in company with 
3 Enpidius longipalpis described above. 


3. MyYRMARACHNE GEDONGENSIS, sp. n. (Text-fig. 11.) 


Prosoma.—Length a little > opisthosoma = p,+t, Caput > 
thorax. Front wall slightly projecting in centre and stepped 
back to eyes 2. Side walls straight, very slightly approaching 
backward. Back wall nearly straight. Groove not marked from 
above, but walls of thorax going on nearly straight with slight 
convex and then concave slope to rear, which is cut nearly square © 
and about 4 greatest width and strongly rimmed. 

On side view caput flat but convex near eye 4, back slope 
steep, groove about = caput, the broad thorax rises convexly to 3 
and then falls gradually convexly and concavely to rim, which is 
shallow. 

Kyes.—1st row straight by summits, 1.1 in contact; 2 subcon- 
tact with 1 and a little stepped back, = 31. Eye square broader 
than long, parallel. 3 slightly nearer 2than 4. 4>2. 4.4= 
4-rim. 

Falces.— Length = cephalothorax, breadth = 7 length. From 
above inner edge straight, outer convex, being widest at %, end 
cut square. From side shghtly concave above, below a double 
flat step to 3, then straight to end. Lighter colour and granu- 
lated. Unguis > 2 paturon, bisinuate, with basal curve nearly 
flat, simple. Teeth, upper 6 or 7, of which Ist is very strong at 
root of unguis and pointing forward, 2 or 3 close to it, and 
remaining teeth at other end of groove and wide. Lower 7 to 9, 
subequally spaced and decreasing from base to apex of unguis. 
Groove well marked. 

Maxille.—As turriformis, but outer corner sharp. Length =c.,. 

22% 


316 MR. H. D. BADCOCK ON 


Labium.—As turriformis, but base is very distinct, terminating 
in a short point well in front of sternum. Apex very slightly 
concave and scopula light. 

Palpi.Spring from near base of maxilla. Tr > width maxilla; 
f slightly > tr, < ¢,, rather strongly curved; p slightly < tr, very 
slightly widening for ward. Ti a little longer and much stronger 
than patella, much wider apically and with strong pointed outer, 
lower, apical process. Fringe of long dark hairs on inner edge. 
Ta broader at base, cut off square at APEX, slightly longer than ti, 
hairy. Fovea occupies about %. A spine springs from outer 
base of organs and ‘passes round on inner edge of fovea, term1- 
nating with a slender forward point at about middle of apex. 


Text-figure 11. 


Myrmarachne gedongensis, sp. n., 3. 


a. Profile. 6. Sternum &c. ec. Palpus. 


Legs.—4.1.3.2, with considerable differences. 
Ist. c longest, length = 23 breadth, parallel or very slightly 

tapering, light, slightly dark at base and sides. 

tr long, nearly $c 

f slightly clubbed at base, dark. 

p = 4 ti, 1 pair spines, light. 

ti, spines from 6 and 7 to 8 and 9, strong, and last 4 or 5 all 
terminating at end of joint, light. 

m, 2 spine pairs, of which Ist very long and both terminate 
about middle of ta, darker. 

tari, ame idark:, 


AN'T-LIKE SPIDERS FROM MALAYA. 317 


2nd. ¢ very short and gibbous, dark. 
tr very short, light. 
f ae clubbed than f,, light with daik sides. 
p—ti, no spines. 
ti, 2 ov 4 spine pairs. 
m, 2 spine pairs, all spines are very strong for 2nd leg. 
ta < 4 ti, hght. 

3rd. ¢ & tr ae ger than c, and tr,. 
f Cems ehabbed slightly. 

= iti. 
ti ice 
Pee tl, tar m. 

Ath. ¢ a little See, | but much < ¢, slightly tapering. | 
tr much longest, >c, <c¢,, cylindrical, light with dark sides. 
f long, slightly clubbed, dark. 

= 1 ti, shghtly curved, dark. 
ti cylindrical, dark. 
m shghtly < ti, darkest at end sides. 
ta = 4m. 
Colour contrasts of legs very slight. 


Sternwm.—Short-pointed between ¢;,1, which are about 4% their 
width apart, and between ¢ .2, shaip-pointed between c2.3, aan 
are only very slightly further apart than the others, and thence 
undulating to a not very long point in front of c4,4, which are 
not quite in contact. 

PrpiceL.—Shortish, j, > j,, [st lorum parallel-sided and con- 
cave behind, slightly wider than 2nd. Below lorum is continued 
forward by a narrow projection towards sternum, but not te 
meet it. 

Sat —Ovate, thickest at middle, very slight con- 
striction at 4, which is marked by slight oblique aite. lines at 
sides. General colour brown-green. Below, central longitudinal 
wide lighter marking from epigastric fold to spinners, slightly 
constricted near front. Epigastrie fold procurved, with area in 
front yellow-brown and corneous. 

Spinners.—As biseratensis. 

Anus.—Distinctly protuberant, with two joints. 


SHort DESCRIPTION. 


2 


Prosoma very slightly > opisthosoma = p,+t,. Caput > thorax. 
Groove and thorax moderate. 

Falces = cephalothorax, unguis > 4 paturon. Upper teeth 6 
or 7, of which | very strong at root of unguisand 2 or 3 adjacent, 
remainder distantly spaced at other end. Lower teeth 7 to 9. 

Palpi.—Trochanter rather long, tibial process strong. Ta 
distinctly > ti. Fovea occupies more than half tarsus: organs 
have spine springing from onter base and passing round inner 
edge to peu ale in fine point in front. 

‘Legs 4.1.3.2, considerable intervals. Spines of ti, from 6.7 


318 MR, H. D, BADCOCK ON 


to 8.9, p, 1 pair, tr, 3 or 4 pairs. c, and tr, much larger than 
the others. 

Pedicel shortish, with long and narrow point reaching forward 
between cy,4 towards but not reaching sternum. 

Opisthosoma ovate, with faint attempt at constriction at + and 
slight white lateral markings thereat. 

Note.—Description is based on two males of lengths 5-9 and 
7°0 mm. from Bukit Besar and Gedong. 

The following dimensions are from the 7:0 mm. specimen :— 


Prosoma 3:2. Opisthosoma 3:2. 


hhegs/ lee... e+til5, £2°9, p+ti34, m+tal9 = 97 
ese o: LOh a eonl eb, Mie Os 3p = DO Se 
OBER Opel, sf eas 5) et Se ae 
Ae cio, slid SOL uemeten yor, ip Mees yf onOrere= 2 

One of the specimens was accompanied by ant no. 6. 


Acorius Thor, . 


Thorell, St. Rag. Mal. 1877, p. 216; E. Simon, Hist. Nat. 
Araign. i. p. 537 (1901). 


2. Agorius Gracitipes Thor. (Text-fig. 12.) 


Prosoma. Cephalothorax.—(a) Front truncated with very 
slightly recurved edge beyond front, of which eyes 1.1 project 
slightly and 2.2 less. Sides straight and very slightly approach- 
ing to % thorax, where they commence to approach more rapidly 
with slightly concave edge. Hind end broadly truncated, some- 
what concave. Hind end of caput is slightly indented and 
thoracic grooves indicated. Separation of thorax from caput 
clear, but groove not deep and no constriction at sides. Edge is 
rather dark at end and ends of sides of thorax, but does not 
appear as a distinct rim. General colour light reddish yellow 
except for black eye-frame on front and sides. 

(5) Eyes 1.1 project strongly like the buffers of a waggon, 
2.2 the same slightly above and behind, clypeus is almost 
negligible and underhung, and these together form the front 
wall. Top of caput distinctly convex from eyes 2 to 4, and again 
over 4, being about ¥ as high as long. Dark eye-frame reaches 
only to about bottom of eyes 2.4 and is almost broken just in 
front of 4. Groove slopes at 45° from above 4, and is only about 
as deep as middle of 4. Thorax rises convexly to about 3 and 
falls concavely for last , end being slightly turned up but not 
really rimmed. Thorax as high as caput. 

(c) Clypeus very slight, smooth, procurved. Short grey hairs 
around eyes 1.1, and dark hairs longer on clypeus and above 
eyes. Top of caput flat or very slightly convex. Sides slope 
inwards very slightly and cdnvexly from below eye 2, which 
projects strongly and forms angle. 


ANT-LIKE SPIDERS FROM MALAYA. 319 


Eyes.—Clypeus concave, <4 eye 2. Ist row strongly recurved, 
top of 1 being level with mid 2. 2= 31, all Ist row in sub- 
contact. Hye-quadrangle broader than long and very slightly 
narrower behind. 4 < 2. 3 distinctly nearer 2 than 4 and 
slightly inside the upper 2.4 tangent. 2 are on attached fore ~ 
and aft projections. 4 project strongly outwards. 

Falces.— Almost concealed from above, nearly horizontal, length 
only a little > diameter eye 1. Moderately broad and rounded 
on top and side views. From below slightly tapering, with a 
few dark hairs in centre. Angle well rounded. Groove distinct. 
Ungues slope inwards. Upper teeth 1 large at angle, 1 small 
below. Lower, 1 large about mid-unguis, but slightly less than 
large upper. There appears to be a small tooth at end of 
paturon behind root of unguis. Scopula of long light hairs on 
outer margin. 7 


Text-figure 12. 


L_=___. 


i} 


a ae Ss 
7 wii at 4e NN 
SS 


Agorius gracilipes Thor., 2. 


a. Profile. 6. From above. ec. Sternum &c. d. First leg. e. Epigyne. 


Mazille.—Much shorter than Myrmarachne, < ¢,, rounded and 
expanded at end and slightly spreading. Serrula at outer edge 
and end; dark scopula at truncation, which is short and nearly 
transverse, a few hairs along outer edge. 

Labiwm.—Very short, broader than long. White edge with 
scanty dark scopula. 

Palpi.Spring from about half maxilla. Tr short, < width of 
maxilla, and 2nd division small; f a little > maxilla, slender, 


320 MR. H. D. BADCOCK ON 


cylindrical, straight or very slightly curved, with slight dusky 
shading at sides; p = 4f > tr, slightly wider at apex than base ; 
ti < p but much thicker and more hairy; ta > ti, joint between 
the two somewhat curved. p, ti, and ta all have hairy fringes on 
both sides, increasing from very slight at p to strong on ta; on 
back they are greenish. Ta and ti are slightly flattened, but not | 
nearly to the same extent as Wyrmarachne, and the joint between 
them is quite distinct. 


Legs.—4 = 1-3-2, but measurements not good and either 4 or 1 
may be the longer. 


Ist. c length = 24 width = nearly width of sternum, light, 
cylindrical. 

tr = 14 ¢, cylindrical, a little darker. 

f long, clubbed at base, slender, darker, and very dark at one 
side below. 

p = sternum > f, slenderer, dark below, light at end; this 
extraordinarily long patella is the most characteristic 
feature of the genus. 

ti < p, with angle half way, after which it becomes stouter 
and bends outwards. 5 spine pairs beginning at angle, all 
are strong, lst reaches more than half way to apex, and 
remainder all end level with apex. 

m very short, < tr. 2 spine pairs, lst very strong, springing 
from middle and reaching well beyond 2nd, which springs 
from end and reaches to mid ta. Joint about as stout as 
end of ti. 

ta = 1} m, lighter and slighter. 

2nd, ef << ¢.white: 

tr = 4¢, light, with black blotch at lower apex. 

f.very slightly if at all clubbed at base, the black of tr is 
carried up as a strong black line below or rather to the 
side of whole length of joint. 

p = #f, slightly curved and stouter at apex. 

ti = 2 p, slender. 

m= $ti; ta missing. 

3rd. c a little shorter and stouter than c.. 


tac, 

f very slightly, if at all, clubbed at base, slightly clouded at 
sides. 

p, ti, m as 2, but slightly stouter. 

ta == Ml. 


4th. ¢ very strong, slightly broader at base than apex, longer 
than c,, stouter than c.. 
ire cand > ir, 
Remaining joints as 3. 
All tarsi have claw-tufts and two pectinate claws. 


Sternum.—Shield-shaped, with expanded ears at front. c¢.1 are 
wide apart, springing from opposite the ears; ¢1.2 is broad, 
nearly = breadth c,, ¢,c, less, c,c, least, cy.4 in contact, sternum 


ANT-LIKE SPIDERS FROM MALAYA. SA 


terminating in a short point in front of them. The legs appear 
to spring from a separate chitinous plate lying between dorsal 
shield and sternum, and produced into a long socket to take ¢.. 

Prpicet.—Very short; it is just possible to see its point of 
entrance below dorsal concavity at top, but it cannot be seen at 
all from below. 

Opistnosoma.— Abdomen = 13 cephalothorax, subeylindrical, 
width = height = < 4 length. Rounded in front and behind, 
being slightly wider behind, shght constriction half way. On 
side view constriction at 3. Gr ound-colour greenish brown above 
formed by a light ground dark-lined, two small white blotches at 
constriction in a transverse line, followed by a large central white 
blotch. From the small blotches white lines slope downwards 
and backwards across the sides, getting wider as they proceed and 
connecting with white ground-colour which prevails below behind 
epigastric fold to spinners. Spiracular plates chitinous, yellow- 
brown, and so wide apart as to be hardly visible from below and 
noticeable from the side. Between them a strip of same colour, 
but I think not chitinous. Epigastric fold is strongly procurved, 
but not well marked. 

Hpigyne.—A small white semicircular pit edged with dark red 
and with a pair of dark markings forming ares of a semicircle 
outside. Not very marked, but clearer than in MWyrmarachne. 

Spinners.—As seen from below, spring from a very distinct 
half-socket, the edges of which, however, are not continued round. 
All spring from separate sockets inside the above and are 2- 
jointed, the basal joint being very long, the second very minute. 
Ist. Very slightly tapering, Ageare twice as long as broad, brown- 
yellow. 2nd. Equal length, lightest, slender est. 3rd. Dare as 
Ist at base, light as 2nd at apex, intermediate in stoutness but 
nearest 2nd, longest. 

Note.—The description is from a single Q found at Bukit 
Besar, 2500 ft. altitude, in cavity under the bark of a rotten 
sapling. 

The following are measurements :— 


Total length 7-0 mm. 
Prosoma ...... 2°7 long x 1:6 wide x 1:4 high. 
Omsihosoma,... 4°6 ,,. xX. 2°0' "4, “x 20 


99 


Legs 
lhe e+tr 14, f2°6, p2°8, t12°0, m+tal0 =9-8 
"ae? ae e+tr+f 2-4, ptti 20), em eo 
eet xe e Deo a0, a Pe s(),, Sen 
Aas, e--tr 1d, -f 2°38; eee Os oO 2 a=98 


The legs were damaged and impossible to measure with 
accuracy, so that either lor4 may be the longest. 
Ants accompanying, none. 


s 


ORBITO-l1EMPORAL REGION OF SKULL OF LEMUR, 323 


24, The Structure of the Orbito-temporal Region of the 
Skull of Lemur. By F. Woop Jones, M.B., D.Se., 


F.Z.S. 
[Received and Read November 20, 1917. ] 
(Text-figures 1-5.) 
INDEX. 
STRUCTURE and MoRPHOLOGY ............ pp. 323-329 


The question of the structure of the orbito-temporal region of 
the Lemurs is one that has by now accumulated a very bulky 
literature ; and this fact is, in itself, almost a guarantee that some 
difficulty and uncertainty is attached to interpreting appearances 
which are open to ordinary inspection in the skulls of animals 
that are neither particularly minute nor particularly rare. 
The uncertainty is at once made apparent by a study of this 
literature ; for very different accounts are given of the elements 
which enter into the composition of this limited portion of the 
skull. But another factor may be detected, and this is a rather 
more subtle one than any mere difficulty in recognising ill-marked 
suture-lines. It is obvious that some have wished to see an 
ordering of the bones upon lines indicative of a Primate kinship, 
while some have neither looked for, nor expected, any indications 
of such affinity. 

The observations recorded here were made upon two full-term 
foetuses—the one of Lemur catta, and the other a hybrid between 
L. albifrons $ and L. melanocephala 2 , both born in the Gardens 
of this Society. Beyond the limits of these two representatives 
of the genus Lemur the present inquiry is not extended so far as 
the study of feetal material is concerned. The question that has 
been an outstanding one for now nearly a century is the presence 
or absence in the orbital wall of an “os planum” or “ planum 
orbitale” of the ethmoid bone. If we turn to a recent work 
(‘Morphology and Anthropology,’ by Dr. W. L. H. Duckworth, 
1915), we find the following as a description of this region in 
Lemur :—‘* On the inner orbital wall, the frontal and maxillary 
bones join along a suture for a distance of about 5 mm., and thus 
widely separate the ethmoidal from the lachrymal bone” (p. 73). 
The condition here described depicts a phase only separated 
by degree from that prevailing in the Anthropoidea, and the 
description leads one to suppose that the lachrymal in the fore 
part of the orbit is separated from the os planum of the ethmoid 
in the hind part of the orbit by the meeting of the frontal and 
maxilla from above and below over the intervening interval 
of 5mm. The figure that accompanies this description lacks 
definition in some of its index lines; but apparently it bears out 
this diagnosis of the disposition of the elements. 


324 DR. F. WOOD JONES ON THE 


By far the most critical and exhaustive account of this region 
is that contributed by C. I. Forsyth Major to the ‘ Proceedings’, 
of this Society on Feb. 19th and March 19th, 1901. Here the 
os planum of the ethmoid is described and figured in an extensive 
series of skulls of Lemuroidea. 

In his earlier paper (p. 131) the author's says -— ~“ Amongst the 

Malagasy Lemurs a fairly large os planum” (of the ethmoid) 
“is present in all the species of Microcebus. In the other genera 
the planum becomes fused with the palatal at a very early date. 
A distinct small os planum is often visible in young specimens of 
Hapalolemur, Chirogale, Lepidolemur, and Avahis, and sometimes 
in Lemur.” But in the later paper the statements concerning 
Lemur ave rather more extended, for at p. 258 the following 
statement occurs :—‘* The lateral part” (of the os planum of the 
ethmoid) ‘“‘ has become united with the palatal at a very early 
stage; a remnant of the suture with the latter bone is seen in 
the adult at its antero-medial extremity ; in exceptional cases, 
e. g.,1n Lemur catia and young specimens of Lepidolemur, the 
planum remains completely or almost completely distinct from 
the palatal.” <A figure of the orbito. temporal region of L. catta 
leaves no doubt as to the author’s meauing. The os planum of 
the ethmoid is diagnosed as intervening between the lachrymal 
in front and the palatal behind. 

Here, therefore, we have an admission of the same bony elements 
as are described by Duckworth; but their arrangement 1s depicted 
in a strangely different order, for whereas Forsyth Major makes 
the ethmoid coterminous with the lachrymal anteriorly, Duck- 
worth separates it from this element by a meeting of frontal and 
maxilla over an interval of 5 mm. 

Going somewhat further back in the gearare of the subject, 
we find a series of authors denying the presence of an os planum 
ethmoidale in the orbit of the Lemurs. It was indeed to refute 
the findings of these authors that Forsyth Major published the 
papers from which the above quotations have been made. 

The French zoologists, for the most part, have denied the 
pfesence of this element, and probably in this they have been 
euided or assisted by their intuitive discrimination which sees 
little but superficial resemblance between the Malagasy Lemurs 
and the Anthropoidea. Cuvier declared that the os planum of 
the ethmoid was in the Lemurs altogether enveloped by the 
frontal and the palatine, and therefore did not appear as an 
element in the wall of the orbit. | 

MM. Grandidier and Alphonse Milne Edwards regarded the 
os planum as being overlain by the frontal which ther refore came 
in contact dir ectly with the maxilla. Obviously Flower came to 
the same conclusion, for he says, “* The os planum of the ethmo- 
turbinal does not enter into the inner wall of the orbit, but is 
shut out by the maxilla.” 

In this cursory survey of the literature we have therefore four 
distinct diagnoses of the elements entering into the formation of 


td 


ae 


ORBITO-TEMPORAL REGION OF SKULL OF LEMUR. a2) 


this region in the Lemurs. The first two (those of Duckworth 
and Forsyth Major) admit the presence of the os planum of the 
ethmoid, but differ widely as to the arrangement of the different 
elements. The second two (those of Grandidier & Milne Edwards 
and Flower, and of Cuvier) exclude the presence of an os planum 
of the ethmoid, but differ as to the elements which overlie and 
exclude this bone. The position will be simplified by reducing 
the four diagnoses to diagrams in which cognisance is taken only 
of the elements which axe in dispute (see text-fig. 1). 


Text-figure 1. 


Diagrams to represent the diagnosis of the elements entering into the formation 
of the orbital wall of Lemur. 


F.=frontal. L.=lachrymal. 2H.=ethmoid. Pal:=palatal. M.=maxilla. 


A. According to Duckworth. B. According to Forsyth Major. 
C. According to MM. Grandidier & Milne Edwards, and also Flower. 
D. According to Cuvier. 


It is often impossible to comment upon the work of previous 
authors unless their written statements can be correlated with 
the specimens from which the descriptions were taken. IJndi- 
vidual variation may play a large part in discordant description, 
and for this reason it behoves every author to specify the material 
from which his descriptions are drawn. In only one case (that 
of Dr. Forsyth Major) is this condition fulfilled. In every case 
in which he describes and figures a specimen he notes its number 
in the National Collection preserved in the British Museum. 

It was therefore the diagnosis of Forsyth Major that I first 
attempted to verify. In 1914, by the kind permission of 
Mr. Oldfield Thomas, | examined the actual specimens described 
in the paper of 1901. The conclusion I arrived at was that the 
problem was one of exceeding difficulty when confined, as it was 
in this case, to the skulls of adult Lemurs. But I also gathered 
the impression that the interpretation of presumed suture-lines 
described and figured by the various authors was, in many cases, 
open to grave doubts, and that the so-called * os planum” was not 
in all cases the same element (see figs. 65 & 66, p. 258, op. cit., of 
specimens 97,9.1.18 and 97.4.6.1). ‘A glance at these two figures, 


326 DR. F. WOOD JONES ON THE 


and more especially an inspection of the actual specimens, will 
convince anyone that the bone labelled “pl” in the two figures 
is not the same element in the two specimens. The same criticism 
holds good throughout the paper. From specimen to specimen 
the irregularity of the apparent suture-lines increases, and the 
only conclusion at which it was possible to arrive was that the 
question was incapable of solution by an examination of adult 
specimens; since it was certainly essential to see the simplified 
condition in the feetal or new-born individual. 

It was not until the present year (1917) that two fatal Lemurs, 
preserved by Mr. R. I. Pocock, were placed unreservedly in my 
hands, and I had the opportunity of examining the structure of 
the orbito-temporal region before any of the post-natal, secondary 


Text-figure 2. 


The skull of a foetal specimen of Lemur catta. Actual length 39 mm. 


changes had set in. Of the two specimens that of L. catia is 
the less highly ossified, and the constituent bones of the cranial 
wall are seen with diagrammatic clearness (see text-fig. 2). The 
condition of the orbito-temporal region is shown in text-figure 3. 
The orbital plate of the frontal takes a very large share in the 
formation of the orbital wall; it meets the nasal and lachrymal 
in front, but at this stage it is separated from the frontal process 
of the maxilla by an unossified interval. Behind it meets the 
parietal at the coronary suture, and below that it meets the 
anterior edge of the very small orbito-sphenoid. Along the lower 
border of the orbital plate of the frontal runs an elongated bone 
shaped somewhat like a ploughshare. This bone articulates in ~ 


ORBITO-TEMPORAL REGION OF SKULL OF LEMUR. BUA 


front with the lachrymal, and behind the orbital plate of the 
frontal it is in contact with the orbito-sphenoid and alisphenoid. 
Below, the greater portion of its length is free, but, in front, the 
maxillary, which is very small and only partially ossified, meets 
its lower border. This long bone is the orbital process of the 
palate, and at about its mid-point it is perforated by two foramina, 
presumably the naso-palatine canals. 


Text-figure 3. 


| ee ee 
Se ee) 


The orbito-temporal region in the foetal L. catta. 


T.=lachrymal. F.=frontal. Pal.=palatine. O.S.=orbito-sphenoid. 
; A.S.=Alisphenoid. P.=parietal. /.=maxilla. 


In the other specimen, which is rather more advanced in 
ossification, the condition is similar in all essential points. ‘The 
sphenoid and the maxilla are somewhat further ossified ; but 
the most interesting feature is a slight bending of the axis of 
the orbital plate of the palate at the site of the naso-palatine 
foramina (see text-fig. 4). 

In the adult of Lemur catta the picture has changed very 
remarkably, for the portion of the palate anterior to the naso- 
palatine canals has become inflated with an air-sinus; and it has 
been somewhat bent back over the uninflated plate posterior to 
these canals (see text-fig. 5). 

Reference to Forsyth Major’s figures of the orbit of Lemur 


328 3 DR. F. WOOD JONES ON THE 


catia B.M. 59e (figs, 61 & 62, p. 256) will show that it is this 
anterior inflated portion of the palate which is diagnosed as the 
os planum of the ethmoid. Concerning the inflation of this part 
of the orbital wall Forsyth Major says: “It” (the so-called os 
planum of the ethmoid) “always forms the roof of a pneumatic 
cavity, which often (L. catta, L. macaco, L. varius) is but an 
appendix of the maxillary sinus. The anterior portion of the 
palatal also participates, as a rule, in the formation of this 
pneumatic cavity, by forming its posterior cul-de-sac.” As a 
matter of facet, it is this variability of the posterior extension of 


Text-figure 4. 


Orbito-temporal region in the fetal skull of L. albifrons X L. melanocephala. 
Length 41mm. Lettering as in text-fig. 3. 


the inflated area that produces the variability of the presumed 
suture-line. The bones of the orbital wall are particularly thin 
and transparent in the Lemurs, and in adult examples the septa 
between air-chambers may be far more conspicuous than actual 
suture-lines. This condition is carried to extremes in old sgpe- 
cimens of some types, such as Galago, in which the whole of the 
orbital wall (and of other regions of the skull) has the appearance 
of cracked ice, each crack corresponding to a bounding wall of, a 
small sinus, and the whole creating such a meshwork of lines 


ORBITO-TEMPORAL REGION OF SKULL OF LEMUR. 329 


that the determining of sutures separating different bony elements 
becomes a hopeless task, 

More than this, synostosis of the different elements is effected 
at a comparatively early stage in the Lemurs. 


Text-figure 5. 


( KA 
GES MW \ ik 
Aen } 


Za™\ 
ae Wii ZZ 
| eae 


ti My mi 
: ey 


Orbit of an adult 2 LZ. catta. Lettering as in text-figs. 3 & 4. 


The palate-bone is not indicated by lettering and is apparently subdivided into 
two portions by the inflation of its anterior extremity. 


Before the comparison of different species, or the much more 
important contrasting of the Malagasy and non-Malagasy forms, 
can be undertaken, an examination must be made of foetal and 
young stages, and until that is done the question of the variability 
of this region must remain an open one. But, for the genus 
Lemur, the two foetal examples described here appear to indicate 
clearly that the os planum of the ethmoid is lacking from the 
orbital wall, and that a very common mammalian, but definitely 
non-primate, arrangement of the bony elements is present in this 
region. | 


Proc, Zoou. Soc.,—1917, No. XX ITI. 23 


THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 35 


EXHIBITIONS AND NOTICES. 
October 23rd, 1917. 


Dr. A. SmirH Woopwarb, F.R.S., Vice-President, 
in the Chair. 


The Secretary read the following Report on the Additions 
made to the Society's Menagerie during the months of May, 
June, July, August, and September, 1917 :— 


May. 


The registered additions to the Society’s Menagerie during the 
month of May were 241 in number, Of these 125 were acquired 
by presentation, 46 were received on deposit, 60 by purchase, 
1 in exchange, and 9 were born in the Gardens. 

The number of departures during the same period, by death 
and removals, was 135. 

Amongst the additions special attention may be directed 
ton 

1 Grison (Grison furax), 1 Cordovan Skunk (Conepatus proteus), 
new to the Collection, and 1 Viscacha (Lagostomus trichodactylus), 
from Cordova, in the Argentine, presented by W. A. Smithers, 
C.M.Z.S., on May 4th. 

4 Ostriches (Struthio camelus), from the Northern Territories, 
West Africa, presented by Capt. C. H. Armitage, C.M.G., D.8.0., 
F.L.S., on May 31st. 

2 Burmeister’s Cariamas (Chunga burmeisteri) and 1 Black 
Vulture (Cathartes atratus), from Cordova, Argentina, presented 
by W. A. Smithers, C.M.Z.S., on May 4th. 


JUNE. 


The registered additions to the Society's Menagerie during the 
month of June were 168 in number. Of these 54 were acquired 
by presentation, 54 were received on deposit, 42 in exchange, 
and 18 were born in the Gardens. 

The number of departures during the same period, by death 
and removals, was 100. 

Amongst the additions special attention may be directed 
WO) 

1 Hensel’s Cat (Lelis pardinoides), new to the Collection, from 
Bahia. presented by W. A. Smithers, C.M.Z.S., on June 11th. 

1 Eland (Taurotragus oryx), born in the Menagerie on June 
21st. 

A collection of Reptiles, including 3 Anacondas (#unectes 
murinis), 1 Thick-necked Boa (Hpicrates cenchris), and 1 Cooke's 


aoe THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 


Tree-Boa (Corallus cookii), from Trinidad ; 1 Antillean Boa (Boa 
diviniloqua), from Dominica ; 2 Black Cr ibos (Oxyrhopus clelia), 
3 Rat-tailed Snakes (Lachesis lanceolatus), from Trinidad; and 
1 Terrific Rattlesnake (Crotalus terrificus), from British Guiana, 
deposited on June 21st. 


JULY. 


The registered additions to the Society’s Menagerie during the 
month of July were 72 in number. Of these 27 were acquired 
by presentation, 7 were received on deposit, 11 by purchase, and 
27 were born in the Gardens. 

The number of departures during the same period, by death 
and removals, was 75. 

Amongst the additions special attention may be directed 
t0.:—— 

1 White-bearded Gnu (Connochetes albojubatus), born in the 
Menagerie on July 12th. 

1 Barbary Sheep (Ammotragus lervia), born in the Menagerie 
on July 19th. 

1 Hyacinthine Macaw (Anodorhynchus hyacinthinus), from 
S. America, deposited on July 21st. 

1 Diamond Python (Python spilotes), from Australia, purchased 
on July 2nd. 

AUGUST. 


The registered additions to the Society’s Menagerie during the 
month of August were 80 in number. Of these 22 were acquired 
by presentation, 14 were received on deposit, 4 by purchase, and 
40 were born in the Gardens. 

The number of departures during the same period, by death 
and removals, was 85. 

Amongst the additions special attention may be directed 
O::—— 

1 Ocelot (Lelis pardalis) 3, from South America, and 1 Serval 
(felis serval), from Africa, presented by Sir Claud Alexander, Bt., 
on August 30th. 

1 Brindled Gnu (Gorgon taurinus) 3, from South Africa, and 
1 Anoa (Anoa depressicornis) 3, from Celebes, deposited on 
August 30th. 

1 Three-streaked Skink (Mabuia trivitiata) and 1 Small-scaled 
Zonure (Zonurus polyzonus), the latter new to the Collection, 
from South Africa, presented by Dr. R. Broom. 


SEPTEMBER. 


The registered additions to the Society’s Menagerie during the 
month of September were 83 in number. Of these 62 were 
acquired by presentation, 2 were received on deposit, and 19 
were born in the Gardens. 

The number of departures during the same period, by death 
and removals, was 69. 


EXHIBITS FROM THE CAIRD INSECT HOUSE. 333 


Amongst the additions special attention may be directed 
TOs; 

3 Coypus (Myocastor coypus), from South America, presented 
by Sir Edmund G. Loder, Bt., F.Z.S., on September 20th. 

2 Yucatan Blue Jays (Cissolopha yucatanica), from Yucatan, 
presented by Guy Falkner, F.Z.8., on September 29th, 

1 White-breasted Kingfisher (Halcyon smyrnensis), from Ceylon, ° 
presented by J. R. Lee Booker on September Ist. 
~ 6 Anacondas (Hunectes murinus), born in the Menagerie on 
September 19th. 


Effect of Captivity on the Cranial Form of the Lion. 


Prof. F. Woop Jongs, M.B., D.Sc., F.Z.8., exhibited the skull 
of a Lioness which had lived for 17 years in the Gardens. This 
skull presented marked changes when compared with the crania 
of wild animals shown at the same time for comparison. The 
changes involved the whole contour of the skull and depended, 
for the most part, on loss of function. The parts concerned with 
actual mastication of food were but little changed ; but the whole 
mechanism for seizing and holding living prey had undergone an 
atrophy in the menagerie animal. This atrophy was especially 
marked in the facial contour related to the canine teeth, and 
manifested itself by the production of a somewhat ursine appear- 
ance of this portion of the skull. At the same time the cranial 
capacity had undergone a marked diminution. The brain of this 
specimen was also exhibited. 

The skull of a cub which was recently born in the Gardens 
was also shown; but no skull of a wild-born eub was avail- 
able for comparison. The wild-born cub is a desideratum, the 
examination of which might possibly yield exceedingly valuable 
information. : 


Exhibits from the Caird Insect House. 
(Text-figure 1.) 


Miss EvELyN CunEsMAN made remarks on the feeding habits 
of the Long Water-Scorpion, Ranatra linearis, on the burrows 
made by the Mole-Cricket, Gryllotalpa vulgaris, and on the 
formation of the case by the Caddis-worm Phryganea grandiosa, 
based. on observations made in the Caird Insect House, and 
exhibited examples of the living insects to the Meeting. 


Ranatra linearis 


has the first pair of legs modified to form instruments for seizing 
prey. These are kept scrupulously clean and ready for action. 
The morning toilet is an elaborate ceremony, each limb being 
minutely overhauled with special attention to the joints. 


334 EXHIBITS FROM THE CAIRD INSECT HOUSE. 


This insect is of an extremely lethargic nature, and will sit 
motionless for hours in the weed or on a plant-stem until some- 
thing swims within reach, when the long fore-legs shoot out with 
a rapidity quite at variance with its other movements. 

It occasionally hunts the bottom, probing the mud for prey, 
but makes no effort to pursue. The food consists principally 
of Asellus and Gammarus, small water-beetles, small dragonfly 
larve, etc., and even snails. : 


Text-figure 1. 


77 i 


RS ~<a Se 4" Se 
rw N i ee 
a 3 ‘Ny 
(C) > - = 3 Ker aie 


vik i FESS 
Colts hts 


Method of feeding of the Long Water-Scorpion, Ranatra linearis. X 3. 


(a) Waiting for prey; (6) Seizing prey; (c) Piercing prey with the rostrum. 


Gryllotalpa vulgaris 
is fortunately rare in England. The burrows being formed 
regardless of obstacles do much damage in cultivated ground, 
and make it a serious pest wherever it occurs in great numbers. 
It is carnivorous, feeding mainly on worms. ‘There is no 
previous record of its having been exhibited in the Caird Insect 
House, but this year three specimens were sent in from different 


ON THE SPALTEHOLZ METHOD OF PREPARATION. i) 


sources for identification ; in two instances the senders spoke of 
their being in numbers when found. 


Phryganea grandiosa, 


the largest of our Caddis-flies, when in the larval state forms its 
case from segments of the hollow stems of aquatic plants, then 
protects each joint by adding a frill composed of leaf-lobes from 
the Hornwort. To do this the larva climbs a plant, and forces 
itself into the top whorl, head downwards: then, when the leaf- 
lobes close round it, cuts them off and glues each singly on the 
case, the loose ends pointing backwards. 


Mr. C. Tate Reaan, M.A., F.R.S., F.Z.8., described the present 
state of knowledge regarding the life-history of the Common Kel, 
illustrating his remarks with lantern-slides. 


November 6th, 1917. 


Dr. A. Smira Woopwarb, F.R.S., Vice-President, 
in the Chair. 


The Spalteholz Method of Preparation. 


Mr. R. H. Burne exhibited a Common Frog prepared by the 
method of Dr. Spalteholz, in which the highly elaborate saccus 
endolymphaticus of the internal ear was very clearly shown. 

The method of preparation in question was introduced in 1906 
by Dr. Spalteholz of Leipzig, and was subsequently brought by 
him to very considerable perfection. Its aim is to render the 
tissues of an organ or even of an entire small animal trans- 
parent. 

The specimen is first thoroughly dehydrated and then fully 
permeated by a mixture of oils of refractive index as nearly as 
possible the same as that of the dehydrated tissue. 

It will be noticed that the method is quite similar to that 
employed by the microscopist in clearing sections or small objects 
for examination by transmitted light. Its success when applied 
to gross objects depends on the degree of accuracy with which 
the refractive index of the clearing agent is adjusted to that of 
the tissues to be rendered transparent. With a clearing agent 
accurately compounded the results are often very brilliant and 
of considerable value for research or museum purposes. ‘This is 
especially the case with complex organs or whole animals owing 
to the fact that different tissues have slightly different indices of 
refraction, so that in the finished preparation the whole object is 
not uniformly transparent, but the several parts appear in it as 
filmy shades of various opacity. 


te 
336 THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 


The method is particularly useful in the study of cavities, such 
as the blood-vessels, which when filled with an opaque injection 
can be seen in their true relations down to the minutest detail. 

In the Frog exhibited the saccus endolymphaticus, owing to 
the fact that it is packed with calcium carbonate (otoconia), 
remaius opaque, in the same way as an injection, though the rest 
of the body has been made transparent. Thus it stands out from 
its surroundings sharp and black when viewed by.transmitted 
light, or brilliantly white if seen against a dark background by 
reflected light. 

It is interesting to recall in connection with this comparatively 
new method of preparation that the older Anatomists were in 
the habit of using a method very similar both in principle and 
practise to obtain transparency for the study of the blood-vessels, 
centres of ossification and similar structures. Many of John 
Hunter’s preparations were “dried and preserved in oil of tur- 
pentine,” and even after more than 100 years show results, 
rougher no doubt, but still worthy to be compared with those 
obtained by Dr. Spalteholz. 


Prof. F. Woop Jongs, M.B., D.Sc., F.Z.S., exhibited a series 
of brains prepared in the Prosectorium, and called attention to 
the fact that specimens preserved by the spirit method differed 
not only in size but in surface details from those prepared by 
more perfect formalin methods, and commented on the necessity 
of revising much descriptive work on the anatomy of mammalian 
brains from the better material that was now available. 


——————_——— 


The Secretary, on behalf of Mr. D. Seru-Smiru, F.Z.S., 
Curator of Birds, exhibited a set of lantern-slides prepared from 
photographs of mammals taken by Mr. Seth-Smith in the 
Society’s Gardens. 


November 20th, 1917. 


KE. G. B. Meape-Waxpo, Esq.,: Vice-President, 
in the Chair. 


The SrcreTary read the following Report on the Additions 
made to the Society’s Menagerie during the month of October, 
1917 :-— 

The registered additions to the Society’s Menagerie during 
the month of October were 118 in number. Of these 36 were 
acquired by presentation, 77 were received on deposit, 3 received 
in exchange, and 2 were born in the Gardens. 


ON PARTURITION IN MARSUPIALS. 337 


The number of departures during the same period, by death 
and removals, was 125. 

Amongst the additions special attention may be directed 
toe 

1 Humboldt’s Woolly Monkey (Lagothrix humboldti), from 
Maniaos, presented by Miss Stella Wickham on October Ist. 

3 Coypus (Myocastor coypus), from 8. America, presented by 
Sir Edmund Loder, Bt., V.P.Z.S., on October 12th. 

2 White-necked Cranes (Pseudogeranus leucauchen), bred in 
Northamptonshire, received in exchange on October 5th. 


The Horns of a Castrated Bushbuck and the Antler-growth 
in Pére David's Deer. 


. R. I. Pocock, F.R.S., F.Z.8., Curator of Mammals, ex- 
tee — 

1. The head of a castrated Bushbuck (Tragelaphus) to show 
the effects of the operation on the horns. The horns differed 
from those of a normal Bushbuck in being slender, subcylindrical, 
untwisted, unkeeled, with a backward curvature like that of the 
horns of a Goral (W. emorhedus) or Serow (Capricoriis). The 
Bushbuck had lived several years in the Zoological Gardens at 
Clifton, and the head was lent for exhibition by Mr. Edward 
Gerrard. 

2, Three successive pairs of antlers of a fully adult Pére 
David’s Deer (Hlaphurus), showing progressive increase in length. 
The first pair shed on Feb. 5th, 1916, measured 15 inches; the 
second pair shed ca Oct. 22nd, 1916, 25 inches; and the third 
pair, unshed at the time of the stag’s death, 40 inches. 


Parturition iv Marsupials and Hxaternal Characters of 
New-born Young. 


Prof. J. P. Hitz, D.Sc., F.R.S., F.Z.8., gave an account of the 
new-born Marsupial and its mode of parturition. Lantern-sldes 
of the recently born young belonging to species of the genera 
Perameles, Dasyurus, Didelphys, Trichosurus, and Phascolarctos 
were exhibited, and their chief external features described. 
Attention was directed to the fact that, whilst the new-born young 
in the different genera all exhibit similar adaptive modifications 
(small size, anterior half of body much in advance of posterior in 
degree of development, fore-limbs strongly developed and provided 
with recurved claws, sucking mouth, presence of thick epitrichial 
layer, covering eye. and ear-pinne, &c.), there are recognisable 
structural and developmental differences between them—e. g., the 
new-born Dasyurus is characterised by the presence of a curious 
swelling in the neck-region and by the absence of any indication 


Proc. Zoou, Soc.—1917, No. XXIV. 24 


338 ON PARTURITION IN MARSUPIALS., 


of the digits of the foot. The early initiation of the lip-fusion, 
leading to the establishment of the sucking mouth, was demon- 
strated in a series of embryos of Z'richosurus, and an interesting 
comparison was instituted between the recently hatched young of 
Ornithorhynchus and the new-born Marsupial. 

The remarkable and unique mode of parturition which occurs 
in certain Marsupials was explained and illustrated by a series of 
micro-photographs. It was shown that in Perameles, Dasyurus, 
and 7'richosurus the young reach the exterior, not through one or 
other of the lateral vaginal canals, but by way of a direct median 
passage, constituted in front by the epithelially lined median 
vagina and behind by a cleft-like ‘ pseudo-vaginal passage,” 
actually formed at the time of parturition as a tear in the 
connective tissue situated behind the median, and between the 
lateral vagine. The occurrence of the same mode of parturition 
was demonstrated for Phascolarctos (the fact being recorded here 
for the first time), and lantern-slides were shown indicating that 
the same mode of birth also holds good for the Gamba (Didelphys 
aurita). The bearing of these facts on the patent condition of 
the median vagina in certain Macropods was briefly discussed. 


INDEX. 
1917.—Pages 217-338. 


| New names in clarendon type. 


; eae 
Systematic references in italics. 


(z.8.1.) indicates additions to the Society’s Menagerie.] 


Acidalia sagittilinea, 243. 

Adisura atcinsoni, 236. 

Agorius gracilipes (Fig. 12), 318. 

Altha lacides, 246. 

Ammotragus lervia (z. 8. L.), 832. 

Amyciea forticeps (Fig. 2), 28+. 

Ancylolomia chrysolinealis, 
sp. m. (Pld. fig. 2), 247. 

— gracilis, sp. n. (Pl. I. fig. 1), 
248, 

pectinifera, 247. 

simplella, 247. 

Anoa depressicornis (z. 8. L.), 382. 

Anodorhynchus hyacinthinus (2. 8. L.); 
332. 

Anthela nigristigma, sp. n. (PI. I. 
fig. 28), 248. 

Apluda incinata, 245. 

ARACHNIDA : 
Thomisidee 

Malaya: systematic, 277. 

Areomolis nigripuncta, sp. n. 
(PVT fie. 6), 206. 

Arbelodes tetrasticta, 246. 

Automolis ignivena, sp. n. (PI. I. 
fig. 7), 266. 

echrecgaster, sp. n. 

fie. 8), 267. 

ochreomarginata, sp.n. (PI. I. 

fig. 9), 267. 


and Salticidse from 


(Pik 


Bagostomus trichodactylus (z. 8. L.), 
331. 


Bithra aganice, sp. n. (PI. I. fig. 11), 
235. 

Boa diviniloqua (z. s. u.), 332. 

Brevipecten icarus, sp. n. (Pl. I. 
fiz. 24), 240. 


Casama vilis, 241. 

Catagramma cynosura : 
(Pld. fie, 3). 275. 

Catasticta flisa tambcensis, 
subsp. n., 260, 

— lathyi, sp. n., 261. 

—— nimbice peruviana, subsp. n., 
261. 

—philone nimbata, subsp. n., 
260. 

Cathartes atratus (z. s.L.), 331. 

Charaxes maudei, sp. n., 271. 

smaragdalis orientalis, 
subsp. n., 272, 

Chloridia obsoleta, 236. 

Chunga burmeisteri (z, 8. L.), 301. 

Cissolopha yucatanica (z, s. L.), 3833. 

Cladocerotis ceneus, sp. n. (PI. I. 
fiz. 12), 236. 


aberration 


| s a s s . 
Ceeliccia campioni, sp. n. (Figs. 


3, 4), 224. 

flavostriata, sp. n. (Figs. 1, ~), 
223. 

macrostigma, sp. n. (igs. 5, 6), 
225. . 


membranipes race nemoricola, 280. 


Proc. Zoou. Soc.—1917, No. XXV. 25 


Xi 


Coeliccia nigrohamata, sp. n. (Figs. 
We 48), 228: 

octogesima, 231. 

sp. (ig. 9); 229. 

Conepatus proteus (z. 8. L.), 331. 

Connocheetes albojubatus (2. 8. L.), 332. 

Corallus cookii (z. s. u.), 332. 


Cortyta griseacea, Boos 


minyas, 239. 
Cosuma marginata, 245. 


rugosa, 245. 

Crotalus terrificus (2. 8. L.), 332. 

Clenusa rectilinca, 239. 

Cynisca, gen. n., 240. 

—— thisbe, sp. n. 
240. 


(BY alpetia: 


Duomitus pindarus, 246. 
Dysodia intermedia, 247. 


Ematheudes lentistrigalis, 248. 

Epicrates cenchris (4. 8. 4.), 331. 

Epidius longipalpis (Fig. 3), 285. 
istigmene lemniscata, 239. 

235. 


wh 


linea, 
tenuistrigata, 230. 
Ernonoey. 
Insecta: Lepidoptera, 253; Exhibits 
from the Caird Insect House, 333. 
Eunectes murinus (4. 8. b.), 831, 333. 
Euproctis rubricosta, sp. n. (PI. 1. 
fig. 7), 242. 
Eurranthis pennigeraria, 243. 
Eurythecodes flavedinaria, 244. 


Felis pardalis (2. 8. 1.), 332. 
pardinoides (z. 8. 1.), 352, 
serval (z. 8. L.), 332. 


Gavara velutina, 246. 
GEOGRAPHICAL: 


Aracuyipa: Thomiside and Salti- 
cide from Malaya, 277. 
Insecta: Heterccera from B. Hh. 


Africa, 233; Rhopalocera from 
South America, 259 ; Arctiide from 


South America, 265; New Butter- | 


flies from Africa and the East, 


271. 


INDEX. 


Giautra arethusa, 
fig. 14), 238. 

Glaucostola maroniensis, sp. n. 
(Pl. I. fig. 10), 268. 

Gorgon taurinus (z. s. L.), 832. 

Grison furax (z. 8. L.), 331. 

Gryllotalpa vulgaris : exhibited, 333. 


sp. a, CRIA. 


g 


‘ Halcyon smyrnensis (z. s. L.), 333. 


Hleterocrita metis, sp. n. (Pl. I. 
fig. 30), 244. 

Heterogomphus sumatranus, 232. 

Heteronygmia aurelia, sp. n. 
(Pl. I. fig. 16), 242. 

Fonorana erebaria, 243. 

Hylemera lepta, sp. n. 
fig. 29), 243. 

Fypene jussalis, 241. 


21, UE. 


strigata, 241. : 


Idalus flavithorax, sp. n. (PI. I. 
fiz, 4), 265. 

InsEcra. 

On Bornean Dragonflies: structure, 
systematic, 223; Heterocera from 
B.E. Africa: systematic, 233; Lepi- 
doptera: concealment by apparent 
disruption of surface, 253; Rhopa- 
locera from South America: syste- 
matic, 259; Arctiude from South 
America: systematic, 265; New 
Butterflies from Africa and the 
East: systematic, 271;- a Gynan- 
dromorph of Papilio lycophron : 
systematic, 273; Three Aberrations 
of Lepidoptera: systematic, 275; 
Exhibits from tire Caird Insect 
House, 308. 


Lachesis lanceolatus (z. s. L.), 332. 

Lacipa albuia, sp.n. (Pl. I. fig. 8), 
242, 

Lagothrix humboldti (z. 8. u.), 3387, 

Laphygma exempta, 273. 

Lemur: structure of skull (Figs. 1-5), 
323. 

Leocyma candace, 289. 

Leodonta zenobia menticola, 


subsp. n., 260. 


“INDEX. 


Mabuia trivittata (z. s. L.), 332. 
MaAmMALIA. 

Deformity of Os penis in a Seal: 
structure, 251; Orbito-temporal 
region of skull of Lemur: struc- 
ture, 323; The Spalteholz Method 
of Preparation, 335; Parturition 
in Marsupials and External Cha- 
racters of New-born Young, 337 ; 
Horns of a castrated Bushbuck, and 
the Antler-growth in Pére David's 
Deer, 337; Photographs of Mam- 
mals, 336; Exhibition of preserved 
brains, 336. 

IWelese flavescens, sp. n. 

fig 11), 268. 

Microloxia ruficornis, 2438, 
Morprronocy. 


(Pl. T. 


See STRUCTURE, 

Myocastor coypus (z. 8. L.), 833, 337. 

Myrmarachne albicrurata, sp. n. 
(Fig. 9), 306. 

—_— biseratensig, sp. n. (Fig. 10), 
312. 

—— cornuta, sp. n. (Fig. 5), 291. 

—— cuneata, sp. n. (Fig. 7), 300. 

—— gedongensis, sp. n. (Fig. 11), 
3165. 

—— lateralis, sp. n. (Fig. 9), 310. 

macillosa (Fig. 4), 287. 

—— ramosa, sp. n. (Fig. 8), 303. 

—— turriformis, sp. n. (Fig. 6), 296. 


Neonerita metaphcenica, gp. n. 
(Pl. I. fig. 5), 265. 

Neritos purpureotincta, sp. n. 
CPi fig. .13); 269. 

Nyctipao acuta, sp. n. (PI. I. fig. 26), 
249. 


Odontestra avitta, sp. n. (Pl. I. 
fig. 15), 236. 

Ophacus ochracea, sp. n. 
fig 12), 269. 

Oxyrhopus cleelia (z. 8. L ), 832. 

Ozarba abscissa, 239. 

sinud, 239. 


(Pl. I. 


subterminalis, 238. 
—— yaria, 238. 


xii 


Papilio erlaces chinchipensis, 
subsp, n., 259. 

-harmodius tabaconas, subsp. 
n., 260. 

—— illyris hamatus, subsp. n., 271. 

iphidamas ayabacensis, 

subsp. n., 259. 


—— lycophron: gynandromorph of, 


273. 


- neophilus parianus: aberration, 


. 


bo 


1s 

—— polydorus ulawaensis, subsp. 
or aberr. n., 272. 

Parastichtis lysis, sp. n. 
fig. 25), 238. 

cxylus, sp. n. (Pl. I. fig. 22), 

237... 


ik 


Perigea capensis, 287, 

Philosamia cynthia: aberration, 275. 
Phryganea grandiosa: exhibited, 323. 
PISCES : 

Scyllium canieula: structure, 217 ; 
Life-history of the Common Eel, 
335, 

Plecoptera polynuvia, 239. 
Plecopterodes moderata, 239. 
Plestosaurus, 221. 

Polygrapha cyanea, Q (Pl. I. fig. 

263, 

Porosagrotis caffra, 236. 
Pseudogeranus Jeucauchen (4. 8. L.), 

337. 

Pteronycta cervicornis, sp. n. 

CBD figs 21), 240: 

Python spilotes (z. 8, u.), 332. 


9 
) 


al 


Ranatra linearis: exhibited, 333. 
Repvinia: 
Use of the Names Plesiosauria and 
Sauropterygia: systematic, 221. 
Rhodesana mintha, sp. n. (PI. I. 
fig, 19), 221. 
Rhodoneura bryaxis, sp. n. (Pl. I. 
fig. 9), 246. 


Salticus modestus, 287. 

Seyllium, hermaphroditism in, 218, 
Stenagrion dubium (Fig. 10), 261. 
Sirigoplus albostriatus (Hig. 1), 280. 


X1V 


STRUCTURE, 

Mammatta: Orbito-temporal region 
of skull of Lemur, 323; Effect of 
Captivity on the Cranial form of 
the Lion, 333; Horns of a cas- 
trated Bushbuck, and the Antler- 
growth in Pére David’s Deer, 337. 

Insecta: Prothorax of Ceeliccia sp., 
Q , 229; Lepidoptera (concealment 
by amparent disruption of surface), 

D3. 
Struthio camelus (z. s. L.), 331. 
Surattha rufistrigalis, sp.n. (Pl. I. 

fig. 3), 247. 

Synemosina procera, 287. 
Syntomis polyxo, sp. n. 
fig. 10), 234. 


(Pl. I. 


Taurotragus oryx (z. s. L.), 331. 

Teinobasis superba, 231. 

Teracotona subterminata, 235. 

Thosea chloris, sp. a. (Pl. I. fig. 5), 
245. 

ganale, 244. 

—— perseis, sp. n. 
244. 


(Pl. I. fig. 4), 


INDEX. 


Thosea undosa, sp. n. (Pl. I. fig. 6), 
244, 

Toxeus maxillosus, 287. 

procerus, 287. 

Trimetepia etheraria, 243. 


VARIATION, 
Manmatta: Deformity of Os penis in 
a Seal, 251; Effect of Captivity on 
the Crauial form of the Lion, 333; 
Parturition in Marsupials and ex- 
characters of new-born 
young, 337; Horns of a castrated 
Bushbuck, and the Antler-growth 
in Pere David’s Deer, 337; Exhi- 
bition of preserved brains, 336. 
Insecta: A Gynandromorph of Pa- 
pilio lycophron, 273; Three Aber- 
rations of Lepidoptera, 275. 
Piscus: Hermaphroditism in Seyllium 
eanicula, 217, 
Vila eueidiformis, sp. n. (Pl. I. 
figs. 1, 1a), 261. 


ternal 


Xylopteryx arcuata, 248. 


| Zonurus polyzonus (Zz. 8, t.), 332. 


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


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1917. 


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AND 57 TExtT-FIGURES. 


MARCH 1917. 


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LIST OF CONTENTS. 


1917, Part I. (pp. 1-103). 


EXHIBITIONS AND NOTICES. 


Page 
The Secretary. Report on the Additions to the Society’s Menagerie during the months 
of November and December, 1916... 4.00. cc ede os ose oreo ou be weve beeen ge ee meee 


Mr. EpwArp Gurrarp. Exhibition of a mounted specimen of a hybrid Thrush and 
LACK DIL sie a's og bib les a etaln me MRE a ane Fane Oa Sree gta  ¢ 60 ie Gina eee 99 


The Secrerary. Report on the Additions to the Society’s Menagerie during the month 
of January, LOU? 0 ka. eS ects wiekt 0 oot ale > Uiatai eo mia cee 


My. D. Srern-Saitn, F.Z.8., Curator of Birds. Exhibition of the Trachea from a male 
ANSEL OWUS SCHUPPOLMOLE “Cars. oy tet eee bk oe oe ee eee ihe nee PPAR a 32 


Mr. C, J.C. Poot. Notes from the Caird Insect House ...........c0c 00x00 IPP ree Ss 


Mr. R. I. Pococn, F.R.S., F.Z.8., Curator of Mammals. Exhibition illustrated by 
lantern-slides of the work of the Beavers in the Scciety’s Gardens, (Text-figure 1.) 100 


Contents continued on page 3 of Wrapper. 


PAPERS. 


Page 


aS 1. On the Pectoral and Pelvic Arches of the British Museum Specimen of Arch@opteryx. 


By Brantstav PetronteEvics, Ph.D., and Axrrnur Suirn Woonwarp, LL.D., F.R.S., 
SERUM re 8 en Phy gos See eo 45 0s wis o's 4 Op euitte 0 spe'Wa sob lees rene eek 


2. On Heude’s Collection of Pigs, Sika, Serows, and Gorals in the Sikawei Museum, 
Shanghai. By Axruvr DE CARLB SoWERBY, F-.Z.8. 1.6. cece cease ee cee eee SEALE 7 


3. Report on the Deaths which cecurred in the Zoological Gardens during 1916, 
together with a List of the Blood-Parasites found during the Year. By H. G. 
Purmimer, F.R.S., F.Z.8., Professor of Comparative Pathology in the a a 
of Science and Technology, London, and Patholoeist to the Society os. ieee ees oT 


4. On the Structure and Function of the Mouth-parts cf the Palemonid Prawns. 
By L. A. Borrapairz, M.A., F.Z.8., Lecturer in Zoology in the University of 
Cambridge, Fellow, Dean, and Lecturer of Selwyn College.’ (Text-figures 1-51.) .... 37 


5. On the Scolex in the Cestcde Genus Duthiersia, and on the Species of that Genus. 
By Frank H, Bepparp, M.A., D.&c., F.R.S., F.Z.S. (Text-figures 1-5.) 


eesee ae ®t eo & 2 


6, 1. The Coleoptera of the Family Cissida found in Britain, with Descriptions of two new 
Species.—2. A new Species of the Coleopteran genus Cryptorrhynchus Illigey. 
By C. J. C. Poot, Assistant Curator Caird Insect House ........ce cc cct eevee eee es -83 


LIST OF; PLAIEs, 


1917, Part I. (pp. 1-103). 


Page 


Purronravics & WoopwarpD: Pl. 1. Arch@opteryx 1... -ceescssvevese 1 


NOTICE. 


The ‘ Proceedings’ for the year are issued in four parts, paged consecutively, 
so that the complete reference is now P. Z. 8.1917, p.... The Distribution 
is usually as follows :— 


Part I. issued in March, 


PP ieyty bo ce Son June. 
pth 6) 3 aia September, 
Pees ta. bb oceelas December. 


‘ Proceedings,’ 1916, Part IV. (pp. 553-756), were published ou 
December 15th, 1916. 


The Abstracts of the ‘ Proceedings,’ Nos. 163-165, are 
contained in this Part. 


aati aiiaioae 


PROCEEDINGS 


OF THE 


| GENERAL MEETINGS FOR SCIENTIFIC BUSINESS 


OF THE 


ZOOLOGICAL SOCIETY 
OF LONDON. 
1917. 


PART II. _ 


CONTAINING Paces 105 to 215, witH 5 PLaTEs 


AND 20 'l'EXT-FIGURES. 


JULY 191%. 


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


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


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LIST OF CONTENTS. 


1917, Parr II. (pp. 105-215). 


EXHIBITIONS AND NOTICES. 


Page 
The Secrurary. Report on the Additions to the Society’s Menagerie during the month 
of Hebruary, TOUT. i. «sways cass btera es Slee wm ee ata oc en Ieee ee i eee 209 


Mr. R. I. Pocock, F.R.S., F.Z.S., Curator of Mammals. Exhibition of a young Lion Cub 
recently deposited in the Gardens, and also a Galago from German East Africa.... 209 


Mr. C. J. Ganan. Exhibition of a living example of the “ Death-Watch” Beetle ...... 209 — 


Mr. C. Davies Suzrzorn, ¥.Z.8. Exhibition of an autograph of Captain Bligh of 


‘The Bounty? sa ca cece eee ence winieiesen 9. eee! ais et ule mie aft ale Whe olsen et rr 210 
Mr. C. Tats Ragan, M.A., F.Z.8. Exhibition of a Piraya from the Amazon .......... 210 
Mr. Aurrep Hzra, F.Z.8. Exhibition of lantern-slides of and remarks upon Big-Game 

Shooting tin India‘)... osc. oe geese se Wale wate Kt asia alas ett an ee rr 210 


The Szcreraky. Report on the Additions to the Society’s ae during the month 
of March, 1917.6. cv eetea cose a gee re beds pe one vo 9 oieiual igi <n 


Mr. C. Dayrns SuErRBory, EZS. Exhibition of the piece of hairy Skin of a Gorilla, sent 


to Richard Owen © 2.2.0 2. Plu shen wale vine Ua sete ees Cece oy 
Mr. D. Seru-Suiru, F.Z.S., Curator of Birds. Exhibition of a female Japanese Pheasant. 211 
Exhibition of a ‘male specimen of the Carolina Duck .....0s.0% 5.5 oe ss ee eee edd 
Exhibition of some specimens of the trachea of various species of Ducks!. San Pee 
Mr, E. Heron-Autey, F.L.S., P.R.M.S., F.Z.8. Remarks on the Mussel-fishery and 
Foraminifera of Hsnandes, illustrated by means of lantern-slides .............. v2 ae 
The Szcretary. Announcement of-the death of Mr. Henry Peavyot.. ..............5- 212 
Dr. P. Cuatmers Mircuect, F.R.S., F.Z.8., Secretary to the Society. Exhibition of 
coloured sketch of an abnormal Cape Lourie... .....24+ 2. cccccseeer ss cnsserse ae 212 


Prof. J. P. Hrut, F.R.S. _F. Z.S. Exhibition of ee of and remarks upon a Bars 


dnceiGipieol genie w oe ee Mate te beeen eee atten nee ee cette eereseeeeene Od ha 


Dr, A. Suirn Woopwarp, F.R.8. Exhibition of enlarged models of the first molar tooth ; 
of a large Chimpanzee, Melanesian Man, and Piltdown Man 


Mr. D. M.S. Watson, F.Z.S. Exhibition of models of the teeth of fossil Horses....... 


Contents continued on page 3 of Wrapper. 


a 


Exhibitions and Notices (contenued). 


Prof, H. Maxwe.u Lerroy, M.A., F.Z.S., Curator of Insects. Remarks on the Silkworms 
and Silk industry of India, illustrated with lantern-slides and specimens of the 
RE ee OC UCOUTI UNM OIL. s)shu cl ay. pebgks oo sks suis SY eee eaee v's le ee eee we de ee 


The Sxcrerary. Report on the Additions to the Society’s Menagerie during the month 
Orompr LOI... Fe ee eee ee eee cade dane se Slee det Basis 


Dr. P. Cuatmers Mircustt, F.R.S., F.Z.S., Secretary to the Society. Exhibition of the 
behaviour of Birds and Mammals in the presence of living Serpents ............-- 


Mr. D. Sxrn-Smitu, F.Z.S., Curator of Birds. Lantern-exbibition of Birds now or 
recently living in the Society’s Gardens ......... SPAS 038 i GR ns i ee 


Mr. R. I. Pocock, F.R.S., Curator of Maminals. Exhibition on behalf of Messrs. Row 


land Ward, Ltd., of the skins of two Zebras from British East Africa ............ 
Mr. R. E. Hotpine. Exhibition of and remarks on a Cirriped ......0...... cee ee eee 


Exhibition of two curiously malformed eggs 


Mr. D. Setu-Smrru, F.L.S., Curator of Birds. Account of the Poultry Exhibition now 


being held in the Society’s Gardens......... 


PAPERS. 


4, The Prechordal Portion of the Chondrocranium of Chimera collie. By Epwarp 
Pneuprs Auus, Jun., F.Z.S. (Plates I-III.) 


ee f®@ ee eeese ts * veo ee se ee ee eet eae es © & so 8 o- 


8. On the Lizards of the Genus Philochortus Matschie. ie G. A. DOCuENGER, E.R.S., 
F.Z.S. (Plates I. & IL.) oh aan ios ee i eee 


eore ee ee ee ee adv oe 


9. An Experimental Investigation of the Migration of Woodcock breeding in the West of 
Ireland. By S. R. Doveuas, M.R.C.S., L.R.0.P.Lond., Capt. I.M.S. (retired), F.Z.S., 
Ist Assistant Bacteriological Department, Medical Research Committee, National 
Imsurarice Act ...... 0005. 


10. A Sketch Classification of the Pre-Jurassic Tetrapod Vertebrates. By D. M. 8. 
Watson, M.Se., H.Z.S., Lecturer in Vertebrate Palaontology in University College, 
London. (Text-figures 1 & 2.) 


Peers eer ene i LO Ne ea ae e) kia ies) ew) ew. Teel ete? 6) 16) @.\8) 1”) 8) eg ier ei e88 «6 ew, 0! & 


11. Notes on some of the Viscera of an Okapi (Okapta ee Selater). By R. H. 
oo M.A., F.Z.S. (Text-figures 1-23.) .. Ree e earoiecar |, 


Pitlepage Aree Be aie OZ) Tee eae se ts Fe eb a 
Tastot Council and Officers ...0.......04. 


peose es 2 oe wee we - ew ee es we ww 


Se en 8 eee ee lh BP eh tw Me hw tee OO het 8 


List of Contents 
Alphabetical List of Contributors ........ 
Index 


CC ee ee ed 


Page 


213 


213 


214 


215 


105 . 


145 


159 


ALLIS : ee es es 
II. | Chimera colliet ......++ 60. 


: Boutmneer: Pl. a 1, 2. Philochortus mele saree a 
3, 4. P. Bae sir ag eae erates 


so that the ee reference e 
is usually as follows:— 


: a Part ae issued i in March. 


; se) ” ; gees Orie June. ay ee 
meats sy ee ages September. — . 
, | ” IV. Su Sir 9 December, 


; + Broveedings, 1917, Part 1. Cou. 1-108), » were 
ap 20th, a e 


“contained i in this 8 


Bia waked 


PROCEEDINGS 


OF THE 


| GENERAL MEETINGS FOR SCIENTIFIC BUSINESS 


OF THE 


ZOOLOGICAL SOCIETY | 
OF LONDON. | 


ae ———=S~«~LZ. 


- PARTS III. & IV. 


CONTAINING Paces 217 to 338, witH 5 Puates 


AND 82 'l'Ex?-FIGURES. 


FEBRUARY 1918. 


PRINTED FOR THE SOCIETY, 
SOLD AT ITS HOUSE IN REGENTS PARK. 
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MESSRS. LONGMANS, GREEN, AND CoO., 
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LiIsY. OF CONTEST. 


1917, Parts III. & IV. (pp. 217-338). 


EXHIBITIONS AND NOTICKS. 


Page 

The Szcrurary. Report on the Additions to the Society’s Menagerie during the months — 

of May, June, July, August, and September, 1917 .......... «7s sos le whclatin ls ea ale 331 
Prof. F. Woop Jonzs, MB.. D.Sc., F.Z.8. Exhibition of the Skull of a Lioness, showing 

the effect: of captivity ........... faeces net ee ye tee eels telnie salt alee 333 
Miss E. CuzresMAan. Exhibition of, and remarks upon, the feeding habits of the Long 

Water-Scorpion, on the burrows made by the Mole-Cricket, and on the formation of 

the case by the Caddis-worm Phryganea grandiosa, based on observations made in 

the Caird Insect House. (Text-figure 1.) ....-... 00 .. cess ween sioie: vidio aarp 833 — 
Mr. C. Tarn Ruaan, M.A., F.R.S., F.Z.8. Description of the present state of knowledge | 

regarding the life-history of the Common Eel, illustrated by means of lantern- 

SLOTS SS a aoe pe aN becca s he sth eo ar tee Gar Rea eee or eee ta PPR PIA ee ro 
Mr. R. H. Burne, M.A., F.Z.S. Exhibition of a Common Frog, showing the Spalteholz 

method of preparation AY et ood RS aot eta tit pares Ws feds oe ee Soir denen ee hers «- Boo 
Prof, F. Woop Jongs, M.B., D.Sc., F.Z8. Exhibition of a series of brains prepared in the 

PEOSOCHOPIUMT 9: sce oie fare Slee ie sale a a ae oo ete ele eon aierd 7 at eens Pera tC: 
Dr. P. Cuatuers Mrrcuait, F.R.S., F.Z.8., Secretary to the Society. Exhibition on behalf 

of Mr. D. Sxrn-Smitn, F.Z.S., Curator of Birds, of a set of lantern-slides prepared 

from photographs of mammals iy alain. eats pao a: sclera bere ene «aaa eter -» 336 
The Sucrerary. poe on the Additions to the Society’s ge during the month 

of October, 1917 .- \ aul damicigr cates ee oo eae eee “ 336 
Mr. B. I. Pocock, F.B.8., F.L,8., FZS., Curator of Mammals, Exhibition of fae aaa 

a castrated Bushbuck, and of three successive pairs of antlers of an adult Pére 

David's Meer soc. oa es ee oe ee tials phuivatieteus iate cn aie ee eae PEP ert Wurake = 7 
Prof. J. P. Hint, D.S8e., F.R.S., F.Z.S. Lantern-exhibition of new-born Marsupials, inh : 

an aecount of the mode of parturition and external characters ......... tS aha Gernot 337 | 


Contents continued on page 3 of Wrapper. 


PAPERS. 


Page 
12. Note on a Hermaphrodite Dogfish. By Rutn C. BamsBer, M.Sc. (Plates I. & IL.) .. 217 


13. On the Use of the Names Plesiosauria and Sauropterygia. By G. A. BouLEnGrER, 
aN ee oleae ili Ghee ties oes cine erellle cals mses ses eae acd ep ac estes 221 


14. Some Additions to the known Dragonfly Fauna of Borneo, with an Account of new 
Species of the Genus Celiccia. By F.F. Larpuaw, M.A.,F.Z.S8. (Text-figures 1-10.) 228 


15. Notes on a Collection of Heterocera made by Mr. W. Feather in British East Africa, 
ett. By Wt-Ook J. M. Faweurr. (Plate 1.) 0... 00. c cee ce cee ee -- 233 


16. Deformity of Os penis in a Seal (Phoca caspica Nilsson). By Srerervs Aupn&Raxy. 
NI ye oN og li ei sial sp te oden a iad ooo nec s anne es 251 


17, Some Observations upon Concealment by the apparent Disruption of Surface in a 
Plane at Right Angles to the Surface. By J. C. Morrram, M.B.(Lond.). (Text- 


NR cg rae aa de ac  wiallpsa lan Wieliens, o weia a e/hielg 6 wa"e pelea skews es os 253 

18. New South-American Rhopalocera. By J. J. Jotcny, F.LS., F.Z.8., F.ES., and 
Smet S abe I) oe panies lb as Vin be ia On 'mp leads ae eee a Ives rt Seales a 259 

RO eae Aiterican Arctiide, By J. J. Jorny, F.LS., F.ZS., FES, and G. Ta.zor, 
ee (UEC. De. oe ee te wee ee Ree a ui iw A eam icra rs ar al aie ob w 6%, eles oie as 265 

20. New Butterflies from Africa and the East. By J.J. Jorcny, F.LS., F.Z.8., F.E.S., 
and G. TaBor, Er Cea ee nh Lanse ell gL Selena Melee Vee e bee ale 8 271 

21. A Gynandromorph of Papilio lycophron, Hbn. By J. J.I OICEY, E.L.S., F.Z.S., F.E.S., 

Memes emit, PHS. (Plate asc Ge. occ five cas ye we de ce enh geeiet cn ee esas 273 

22. Three Aberrations of Lepidoptera. By J. J. Jorcry, F.LS., F.Z8., F.E.S., and 
Peeerer eee Eate My ae Pern Seana ses by lain Goal seinen ee mei esveainn vee’ 275 

23. Ant-like Spiders from Malaya, collected by the Annandale-Robinson Expedition, 
fut. by HD. Bancoon, M.A. (YLext-figures 1-12.) 22... ee eee ee 277 

24. The Structure of the Orbito-temporal Region of the Skull a Lemur. By F. Woop 
Paton voc: P25. (Rext-feures 1-5.) 0. sce cae. sc cae s as cevtteenaeses 323 
Vitlepage .. 2... eee e eee ete eee ee et eee te cae eet teens icteceye, elcce'a oie i 
dass of Council.and Officers ......7....... eee sae Cae aN AMS Nas. fe Wiles a) lis sie wine G5 ghd ii 
Be OOMECHLS, caylee s Cele voc wees cn ea cows Oe Mitte erie dan We c00y aioe lara: atc G'n’ e's a seco 9% lii 
Alphabetical List of Contributors ........ ... NEN TMEE thie SN Beta eee Soren Rr a Re ee vii 


ieee Syren aE Se CRN oi SN Sp CN Se a ect eae ac a Rs a xi 


LIST OF PLATES, 


1917, Parts lil. & TV. (pp. 217-338). 


BampBer: Pl. I. 
i. 


Fawcerr: Pl. I. Heterocera from British Hast Africa.......cececece 


\ Seyltiutm camiculd ..-...sveveecessernssesandins 217 


233 


Pl. I. Rhopalocera from 8. America .... 259, 265, 275 


Jorcey & Tansor: 
Pl. Ti, «Papilio lyeophrom .i045 000 ce4s oe, cee eee 273 


NOTICE. 


The ‘ Proceedings’ for the year are issued in four parts, paged consecutively, 
so that the complete reference is now P. Z. 8. 1917, p.... The Distribution 
is usually as follows, but on account of war conditions Parts III. & IV. of 1917 
were issued as one Part, and a similar course may be followed in 1918 :— 


Part I. issued in March. 

EY a June, 

erage 1) Gl Ee 5, . September. 
1, Gia December. 


‘ Proceedings,’ 1917, Part IT. (pp. 105-215), were published on 
July 18th, 1917. 


The Abstracts of the ‘ Proceedings,’ Nos. 172-174, are 
contained in this Part. 


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