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PROCEEDINGS

0b THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS |

ZOOLOGICAL SOCIETY

OF LONDON.

1916, pp. 1-448,

with 14 Puares and 120 TExt-FIGURES.

Zaeanian Institugj ;
BeeOUN 7]

: National Museu

PRINTED FOR THE SOCIETY,
SOLD AT ITS HOUSE IN REGENT’S PARK.
LONDON:
MESSRS. LONGMANS, GREEN, AND CO.,
PATERNOSTER ROW.

eS

OF THE

COUNCIL AND OFFICERS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

WS)

Patron.
His Masesry Tue Kina.

COUNCIL.
His Grace Tae Duxe or Beprorp, K.G., F.R.S., President.

Tue Hon. Crcit Barine, M.A.
ALFRED H. Cocks, Esq., M.A.
Toe Rr. Hon. THe Harp

oF Cromer, P.C., G.C.B.,

G-C.M-G., FR: Vice=
President.
CHaRLes Drummonp, Esq,

Treasurer.

ALFRED Ezra, Esq.

Caprain HucH 8. GLaDsTons,
M.A.

SIDNEY Freperic HARMER, Ksq.,
M.A., Sc.D., F.R.S., Vice-
President.

Con. Siz WAuTER R. LAWRENCE,
Br., G.C.1.E.

Srr Epmunp Gites Lover, Br.,
Vice-President.

Prof. Ernest W, MacBnrips,
M.A., D.Se, F.R.S., Vice-
President.

7

oe

Gy PAG. PK.
Esq., D.Sc.

E. G. B. Merapre-WaAtpo,
Ksq. ;

P. Cuatmers MircuHet, Ksq.,
iWhva\es IDSs IbbEIDS. TI RIS
Secretary.

Apert Pam, Hsa.

Tue Karu or PortsMoutH.

OLDFIELD THomas, Ksg@.,

AuByN Trevor-Bartys, Esq.,
M.A.

Antuony H. WHINGFIELD,
Ksa.

Artraur Smita Woopwarpb,
Ksq., LL.D., F.R.S., Vice-
President.

Henry W oopwarb, Hsq., LL.D.,
F.R.S., Vice-President.

MARSHALL,

PRINCIPAL OFFICERS.
P. Cuaumers Mircuent, M,A., D.Sc., LL.D., F.B.S.,

+

Secretary.

R. I. Pocock, F.R.S., F.LS., Curator of Mammals and
Resident Superintendent of the Gardens.

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

Epwarp G, BouLencer, Curator of Reptiles.

Prof. H. Maxwett Lerroy, Curator of Insects.

Prof. Heyry G, Pummer, F.R.S., M.R.CS., Pathologist.

Henry G. J. Peavor, Librarian and Clerk of Publications.

JouNn Barrow, Accountant.
W. H. Corn, Chief Clerk.

LIST OF CONTENTS.

1916, pp. 1-448.

EXHIBITIONS AND NOTICES.

Page
The Secretary. Report on the Additions to the Society’s
Menagerie during the months of November, December,

zane se ene rena Wene HO Hie) = Oicnnan rai arate ers cepts race nve tase 297

Mr. R. E. Houpine. Exhibition of the Skull of a Roebuck. 298

Mr. C. Tate Recan, M.A., F.Z.S. Lantern-exhibition of

draninaston larval) Pishes? 225.053.) eo a. ee 298

Mr. OuprieLpD THomas, F.R.S., F.Z.S. A new Sable
Antelope from Angola. (Text-figure 1.) ............... 298

Mr. R. I. Pocock, F.R.S., F.L.S., F-.Z.8., Curator of
Mammals. Antlers of a Virginian Deer affected by

QW cir Ce Tae AEF eases Pw 3 a cp MONE eT oe ache RN NM UT 301
The Rev. H. N. Hurcuinson, M.A., F.Z.S. Exhibition of
drawanesromexctimet amiinall Sy mpeesaee snes ck tea tee 302

Mr. C. Tare Reean, M.A., F.Z.S. Lantern-exhibition of
a Siamese Fighting-Fish and of a Cat-Fish ............ 302

Mr. R. I. Pococx, F.R.S., F.LS., F.Z.S., Curator of
Mammals. The Tympanic Bulla in Hyenas. (Text-
TSIoATAR ESAS eaure) | oars CURB ERR RE Gib6 2s bon cae book e Ee eRe REaEenn a amiceG 303

Mr. H. K. Eustace, F.Z.S. Bioscope-exhibition of African
SANIT CLS EH LPs! See ote OR OL USSU OL NE a acisciul rani cise 307

The Secretary. Report on the Additions to the Society’s
Menagerie during the month of February 1916 ...... 44]

iv

Mr. EK. T. Newton, F.R.S., F.Z.S. Exhibition of the skin
of a Siberian Black Hare

CeCe en ee ee rea}

Mr. D. M.S. Warson, F.Z.S. Notice of remarks on the
habits of Platypus and Hchidna

CC i i i i i ii ii ii iit

Mr Rev: Pocock, BARES.) (EIS SEZs aC uratommon
Mammals. Lantern-exhibition to show the structure
of the Alisphenoid Canal in some Civets and Hyenas.
(Text-figures 1 & 2.)

Mr. J. T. CunninecHam, M.A., F.Z.S. Exhibition of skins
illustrating results of Mendelian Cross in Fowls

Mr. D. Sers-Smirn, F.Z.8., Curator of Birds. Exhibition
- of a small Intensive Poultry-House

wet eee ae weer oer eres

Iron, Ve. les leben, IDS, WINGS, INAS, Idoxlanloninen Gil
living Cecilians from South America

Mr. G. A. Bounencer, F.R.S., F.Z.5. Notice of a paper
entitled ‘‘On the Lizards allied to Lacerta muralis,
with an account of Lacerta agilis and L. parva”

The Secretary. Report on the Additions to the Society’s
Menagerie during the month of March 1916

efufel eel edalielts

The SecRETARY. Notice of a letter received from Lt.-Col.
R. T. Leper, D.Sc., F.Z.S., R.A.M.C., in reference

to Bilharziosis

Ci i i Cie ii i i i i ei i ii i a ier aay

Mr. C. Tare Ruean, M.A., F.Z.8. Exhibition of lantern-
slides illustrating parental care in Fishes ...............

The Secrerary. Report on the Additions to the Society’s
Menagerie during the month of April 1916 ............

Mr. R. H. Burne, M.A., F.Z.8. Exhibition of preparations
of connections between the Swim-bladder and Ear
IME SES Be Soin. osciel wii. cake a eee ee eee

Correction to Prof, K. B. Pouron’s paper on Moths from
Somaliland

Page

bo

V

PAPERS.

. The Morphology of the Cyprinodont Fishes of the Sub-

family Phallostethine, with Descriptions of a new
Genus and two new Species. By C. Tatz Recan,
M.A.,F.Z.S. (Plates I-IV., and Text-figures 1-15.)

. Ona Collection of Mammals from the Coast and Islands

of South-East Siam. By C. Boprn Kuoss, F.Z.8.,
F.R.G.S. With an Account of the Fruit-Bats, by
Dr. Knup ANDERSEN, F.Z.S. (Plate I., and Text-
AHI OSH ATE OB). eee wt se asic ceae ssis's games PRA oe Moston ate saaaie

. Report on the Deaths which occurred in the Zoological

Gardens during 1915, together with a List oa the
Blood-Parasites found during the Year. By H. G.
Puimer, F.R.S., F.Z.8., Professor of Comparative
Pathology in the Imperial College of Science and
Technology, London, and Pathologist to the Society.

A Frog with symmetrically Abnormal Hind Feet. By
R. W. Harotp Row, B.Sc., F.L.S., F.Z.8., Assistant
Lecturer and Demonstrator in Zoology, University of
London, King’s.College. (Text-figure 1.)...............

. On a Collection of Moths made in Somaliland by Mr. W.

Feather. By Professor E. B. Poutron, M.A., F.R.S.,
F.Z.S. With Descriptions of New Species by Sir G.
F. Hampson, Bart., L. B. Prout, J. H. Durrant,
and Dr. Karu Jorpan. (Plates I. & IT.)

. Further Observations on the Intestinal Tract of Mam-

mals. By P..CHAtmers Mircustt, M.A., D.Sc.,
LL.D., F-R.S., F.Z.8., Secretary to the Society.
(Text- fleur es 1- 30.) STs of MEMES rie et ie oth eta led oe

. Studies on the Anoplura and Mallophaga, being a Report

upon a Collection from the Mammals and Birds in
the Society’s Gardens.— Part I., with a Preface. By
Bruce F. Cummines, British Museum (Natural
History). (Text-figures 1-24.)

ern i ie

Page

no
=I

~T

ba |

87

9]

185

v1
Page
8. Observations on the Cytology of Flagellates and Ameasbze
obtained from old Stored Soil. By 'T. Goopry, D.Sc.,
Protozoologist, Research Laboratory in Agricultural
Zoology, University of Birmingham. (Plates I.-IV.,
and ‘Lext-heure sls) sik. sees eens sans 2 een 309

9. On some Fresh-water Entomostraca from Ceylon. By
Ropert Gurney, M.A., F.Z.8. (Plates I.-II1., and
Pext=figure:].) inci eaceat eis ak eae ee ee eee 333

10. On Specimens of the Perciform Fish Tilapia nilotica
with increased number of anal spines. By G. A.
IBOUTENGER. GHARg Os, ohne cere here ee EERE EEE Ren eee ene 345

11. On the External Characters of the Mongooses (Mun-
gotide). By R. I. Pococn, F.R.S., F.LS., F.ZS.,
Curator of Mammals. (Text-figures 1-10.) ......... 349

12. Notes on the Sitatunga or Marsh Antelope of the
Sesse Islands, Lake Victoria Nyanza. By Major R.
MEINERTZHAGEN, F.Z.S. (Text-figures 1&2.) ...... 375

13. An Experimental Determination of the Factors which
cause Patterns to appear Conspicuous in Nature.

By J.C. Morrram, M.B.(Lond.). (Text-figures 1-20.) 383

14. On a small Collection of Vertebrate Remains from the
Har Dalam Cavern, Malta; with Note on a new
species of the genus Cygnus. By DororHna M. A.
Bare, Hon.M.B.O.U. (Text-figures1& 2.) ......... 421

15. The Poison-Organ of the Sting-Ray (Zrygon pastinaca).
By Tempy.-Major H. Murr Evans, M.D.(Lond.),
R.A.M.C.(T.) (Text-figures 1-7.)

Alphabetical List of Contributors ............ccccecececeseceeee vil

Index

AEE ABETICAT Gis £

OF THE

CONTRIBUTORS,

With References to the several Articles contributed by each.

(1916, pp. 1-448.)

ANDERSEN, Dr. Knup. ‘See Kioss, C. BopEn.

Bate, Miss DorotHEA M. A., Hon.M.B.O.U.

On a small Collection of Vertebrate Remains from the
Har Dalam Cavern, Malta; with Note on a new species

of the genus Cygnus. (Text-figures 1 & 2.) ...............

Boutencer, Georce A., F.R.S., F.Z.S8.
On Specimens of the Perciform Fish Zilapia nilotica

with increased number of anal spines .....................+45

Notice of a paper entitled “On the Lizards allied to
Lacerta muralis, with an account of Lacerta agilis and

LEGG IR ile MSer res "ABR B nn Bion9 386 25 <n so SORA eo RE ee

Burne, Ricuarp H., M.A., F.Z.8.

Exhibition of preparations of connections between the

Sivim-_bladder-and Har m Wishes 2.2 i0c.eockel ode oos we csee

42]

447

vill
Cummines, Bruce F.

Studies on the Anoplura and Mallophaga, being a
Report upon a Collection from Mammals and Birds in
the Society’s Gardens.—Part I., with a Preface. (Text-
MeO IWA'S)) oh oa nccedacoonoosonab06docduodossAo028IC0aN0C~ sees

CunnineHaM, JosepH T., M.A., F.Z.S8.

Exhibition of skins illustrating results of Mendelian
Cross ini“ Owls nate e eee ea eee a cere cence

Durrant, J. H. See Poutton, Prof. E. B.

Eustace, Harry K., F.Z.8.

Bioseope exhibition of African Animals

eeecec ese eo coecesece

Evans, Tempy.-Major H. Murr, M.D.(Lond.), R.A.M.C.(T.).

The Poison-Organ of the Sting-Ray (7rygon pastinaca).

(Mext aio uvesRla 1s) ci. esos. vin. ciecen Bae eRe see eee ee eECer
Goopvey, T., D.Sc.

Observations on the Cytology of Flagellates and
Amcebe obtained from old Stored Soil. (Plates I.-IV.,
and Mextotig ured) "0 a4 1. ssc.eeeeenee ce omen eee eee Eee ee

Gurney, Ropert, M.A., F.Z.8.

On some Fresh-water Entomostraca from Ceylon.

(Plates I.—III., and Text-figure 1.)

Peer een eset one resect oney eee

Hampson, Sir Grorce F., Bart., F.Z.S. See Pouuron,
Proteus:

Hin, Prof. Jauus P., D.Sc, F.B.S., F.Z.S.

Exhibition of living Cecilians from South America ...

Hoxpine, R. EH.

Exhibition of the Skull of a Roebuck

eseccoorecesesscocece

Page

253

446

307

431

309

333

447

1X
Hurcutson, The Rev. Henry N., M.A., F.Z.5.

Exhibition of drawings of extinct animals ...............
Jorpan, Dr. Karu. See Pounron, Prof. E. B.

Kuoss, C. Bopgn, F.Z.8., F.R.G.S.

On a Collection of Mammals from the Coast and
Islands of South-East Siam. With an Account of the
Fruit-Bats, by Dr. Knup Anprrsen, F.Z.S.. (Plate L.,
and Text-figures 1 & 2.)

MerNerrzHAcen, Major Ricuarp, F.Z.5.

Notes on the Sitatunga or Marsh Antelope of the
Sesse Islands, Lake Victoria Nyanza. (Text-figures
SOc) Wee oops ates e hia oalale dnsaisrsyiste os eh rast aa Seleisute wane ee

Mrrcuett, P. Coaumers, M.A., D.Sc., LL.D., F.R.S., F.Z.8.,
Secretary to the Society.

Further Observations on the Intestinal Tract of Mam-
mals. (Text-figures 1-30.)

eee cee ere eee creer se oneeesesesereneee

Report on the Additions to the Society's Menagerie
during the months of November, December, and January,

TRON AG aig tte ca tic 2 2H saan Re as acta va Bich AG tiara hase Aaratg

Report on the Additions to the Society’s Menagerie
during the month of February 1916..............:-...eeeeeee

Report on the Additions to the Society’s Menagerie
during the month of March 1916 .........2..22:eeeeeeee eee

Notice of a letter received from Lt.-Col. R. T.
Leiper, DSc., F.ZS., R.A.M.C., in reference to
Bilharziosis ....... aia atl Garena ee a eee ACES SSO SPN

Report on the Additions to the Society's Menagerie
during the month of April 1916 ..... PEARY cunts oni alsin
Proc. Zoou. Soc.—1916, b

no
“I

375

447

x
Morrram, James C., M.B. (Lond.).

An Experimental Determination of the Factors which
cause Patterns to appear Conspicuous in Nature. (Text-

fiowres L205) oak 2 woes ee eee eae cee a ee

Newton, Epwin T., F.R:S., F.Z.8.

Exhibition of the skin of a Siberian Black Have ......

Pirmer, Prof. Henry G., F.R.S., F.Z.S8., Pathologist to
the Society.

Report on the Deaths which occurred in the Zoological
Gardens during 1915, together with a List of the Blood-

Barasibes ound cuning ath em Vecaaeeeeeneeseeeee ne caerereE eee e

Pocock, Reernatp [., F.R.S., F.LS., F.Z.8., Curator of

Mammals.

Antlers of a Virginian Deer affected by Cancer.........
The Tympanic Bulla in Hyenas. (Text-figures 1 & 2.)

On the External Characters of the Mongooses (Mun-
Somes), (Ciesesnemmas WENO) oo. cccosnccnndpooscosodounoe ease

Lantern-exhibition on the structure of the Alisphenoid
Canal in Civets and Hyznas. (Text-figures 1&2.) ......

Poutton, Prof. Epwarp B., M.A., F.R.S., F.Z.S.

On a Collection of Moths made in Somaliland by Mr. W.
Feather. With Descriptions of New Species by Sir G.
F. Hampson, Bart., L. B. Prout, J. H. Durrant, and
Dr. Karn Jorpan. (Plates I. & II.)

eee seer cases cece seesoese

Correction

aTsielelsieseseisiejeleisecsielsie.e is) ekermalelelelsielellelsielalsialelciullelelelalelsteletalelersierrta

Prour, l. B. See Povnron, Prof. BH. B,

Page

44]

~I

“I

349

xall

Ree@an, C. Tate, M.A., F.Z.8.

The Morphology of the Cyprinodont Fishes of the Sub-
family Phallostethine, with Descriptions of a new Genus
and two new Species. (Plates I.—-IV., and Text-figures
AON 2 orig Reveearerse ies Sacha sa vet oe sete Stele trance atalene a sleuils

Lantern-exhibition of drawings of Larval Fishes ......

Lantern-exhibition of a Siamese Fighting-Fish and of
SO ait Pitis ige ec aaterr ise ators Labs ascicfu Se tbge Nap cin Law al paoui stole a sevcbe iste

Exhibition of lantern-slides illustrating parental care

TAVDL UBLTIGS) SiC GYSy Pk 2 See RRR yr a ne ce eR RL Nr ee Ra a Oe

Row, R. W. Haron, B.Sc., F.L.S., F.Z.8.

A Frog with symmetrically Abnormal Hind Feet.
(CISRGSTSUNES UN) Beadaocasdsoet one asec oossmoeponcn song ceaone ognaonG

Sura-Suirn, Davin, F.Z.S., Curator of Birds.

Exhibition of a small Intensive Poultry-House

THOMAS, OLDFIELD, F.R.S., F.Z.8.

A new Sable Antelope from Angola. (‘Text-figure 1.)

Warson, Davip M.S., F.Z.S.

Notice of remarks on the habits of Platypus and
JSIGIIVEROID See nB he NOC BOR DOR ROSE OER Hoer 30600000 5° cae nonpeR GEDA Nnobuec:

Page

302

4AT

446

298

4 ia ed biti a

ej cad NeX adam wnt, Aah Hd

3 es ocd ue} DP ep aN ee a ia
ee : Or ee ee fraewne an wHidivtnatiays sae xay9, it

SSE Gal eee aa wait

INDEX.
1916.—Pages 1-448.

[New names in clarendon type.

Systematic references in italics.

(4.8.1.) indicates additions to the Society’s Menagerie. ]

Acauthion klossi, 60.
Acantholipes circumdata, 135.
—— triment, 135.

Acanthonyx seriopuncta, sp. n. |

(Pl. I. fig. 41), 125.

Achea catella, 125.

Achthina, gen. n., 179.

——- etenodes, sp. n., 180.

Acidalia actuaria, 149.

— addictaria, 151.

cinerascens, 160.

cenosaria, 152.

consentanea, 149,

— derasata, 149.

—— horiochroea, sp. n., 149.

— luridata, 152.

—— minoa, sp. n. (Pl. II. fig. 20),
148.

—— minorata, 149.

—— pulchellata, 151.

— pyrrhochra, sp. n. (Pl. II.
fig. 18), 152.

— remotata, 149.

—— sagittilinea, 153.

—— spoliata, 149.

—— timia, sp. n. (Pl. II. fig. 19),
151.

—— (Pylarge) nepheloperas,
sp. n,,_150.

Acidaliastis subbrunnescens,
sp. n., 146.

Aclonophlebia inconspicua,
sp. n. (Pl. IL. fig. 9), 139.

Acontia albayo, 125,

gephyrias, 125,

Acrapex albicostata, sp. n. (Pl. I.
fig. 14), 109.

Acroriesis ignifusa, gen. et sp. n.
(Pl. I. fig. 6), 110.

_ Aigocera brevivitta, 103.

Agathodes musivalis, 174.
Aglossa basalis, 171.
incultalis, 171.
ommatalis, 171.

| Alona intermedia, 335.

rectangula, 339,

| Alonella davidi, 335.

excisa, 336,

— karua, 336.

Ameeba agricola, sp. n.: structure,
development (Pl. IV. figs. 66-74),
325.

lawesiana, sp: n.: structure,
development (Pls. IIL., IV. figs. 49-
65), 321.

Amyna octo, 120. -

punctum, 120.

Anadiasa simplex, 166.

Anatomy. See Structurn.

Ancylolomia pectinifera, 169.

Anoa depressicornis (z. 8. L.), 298

Anomis erosa, 135.

Jimbriago, 135.

Antarchea fragilis, 135.

— subflavalis, 135.

Antigastra catalaunalis, 175,

x1V

Apisa canescens, 101.

Arbelodes rufula, 166.

Argadesa materna, 136.

Ariela fasciata: structure (Figs. 2, 4,
7, 103, 849.

Artibeus planirostris: structure (Fig.
2D) 228),

Ascotis selenaria, 168.

Aspilates semispurcata, 157.

Asplenia rubrescens, sp. n. (PI. 11
fig. 1), 135.

Athetis discopuncta, sp. n. (PI. I.
fig. 8), 106.

—— ectomelzna, sp. n. (Pl. L. fig. 9),
106.

leuconephra, 106.

Atilax paludinosus: structure (Figs. 1, |

4,5, 9), 849.

Auchenisa cerurodes, sp. n. (PI. I.
fig. 43), 129.

Aulotarache plumbeogrisea,
sp. n. (Pl. I. fig. 34), 122.

Authadistis camptogramma,
sp. n. (Pl. I. fig. 44), 128. .

AVES:

Fossils from Malta: systematic, 421. |

Axis axis (z.S.L.), 447.
zygophleps inclusa, 169.

Babirussa babirussa: strueture (Fig. 17), |

214.

Balenoptera physalus: structure (Fig.
15), 211.

BaTRACHIA:
Rana temporaria: variation, 87.

Bdeogale puisa: structure (Figs. 6, 9),
349.

Boopia tarsata: structure (Fig. 9), 269.

Bostra pyrochroalis, sp. n. (Pl.
II. fig. 43), 172.

—— tenebralis, 172.

varians, 172.

Bubo bubo (z. s. u.), 298.

Bufo arenarum (2.8. L.), 298.

Calamoschoena, gen. n., 169.
—— ascriptalis, sp.n. (PI. II. fig. 38),
170.

INDEX.

Calpe vagabunda, 136.

Canthocamptus grandidiert, var. (Pls. I.,
IL. figs. 7-9), 337.

Capreolus capreolus: variation, 298.

Casama viiis, 139.

Catephia eurymelas, sp. n. (PI. I.
fig. 49), 132.

mesonephele, sp. n.
fig. 48), 131.

—— pericyma, sp. n. (PI. I. fig. 46),
131.

—— poliochroa, sp. n. (PI.I. fig. 47),
130.

-—— pyramidalis, sp. n. (Pl. I. fig.
45), 120:

Cephalophus dorsalis (z. s. u.), 447.

Cercopithecus preussi (z.s.u.), 441.

Certodaphivia cornuta, 334.

rigaudt, 3d4.

Cerocala albimacula, sp. n. (Pl. I
fig. 42), 126.

—— illustrata, 126.

-—— oppia, 127.

ei, Th

Cervulus cambajensis, 62.
Cervus aristotelis, 62.
unicolor, 62.

subsp. 62.
cambojensis, 62.
equinus, 62.
Cetola pulchra, 105.
Chalciope hyppasta, 126.
Chelecala trefoliata, 125.
Chilena donaldsont, 167.

| Chionoxanthia leucophza, sp. n.

(Pl. I. fig. 23), 115.
Chiromys madagascariensis: structure
(Fig. 29), 237.
Chlorerythra rubriplaga, 148.
Chloridea albivenata, sp. n. (Pl. I.
fig. 3), 103.
— obsoleta, 104.
Chlorissa stibolepida, 148.
Chydorus barroisi (P1. I. figs. 4, 5), 336.
-— parvus (Pl. I. figs. 2, 3), 336. .
Cirphis loreyi, 104.
Citellus mongolicus (z. 8. L.), 297.
Cledeobia radialis, 174.
Coenina tergimacula, sp. n. (Pl. II.
fig. 12), 162.

INDEX.

Ccenobasis chloronoton,
(RIE fie 35); lor:

—— fulvicorpus, 167.

Comibena stibolepida, 148.

Comostolopsis stillata, 145.

Connocheetes gnu: structure, 220.

Constantiodes pyralina, gen. et
sp. n. (PL. I. fig. 85), 107.

Cornifrons albidiscalis,
(Pl. II. fig. 49), 177.

Cortyta canescens, 128.

leucoptera, 127.

rosacea, 128.

Craspedia addictaria, 151.

rujinubes, 151.

sagittilinea, 153.

Cricetulus griseus (z. 8. L.), 297.

Crocidolomia binotalis, 174.

sp. 1).

sp.

Crocuta crocuta: structure (Fig. 2, ©), |

304; (Fig. 2), 444.
Crossarchus obscurus: structure (Kigs,
2, 4,7, 9, 10), 349; (Big. 1), 442:
CRUSTACEA :

Entomostraca from Ceylon: system- |

atic, 333,
Cyclestheria hislopi, 333.
Cyclops distinctus (Pl. I. fig. 6), 337.
hyalinus, 337.
——— Jeuckarti, dal.
prasinus, 337.
VarICANS, DOI.
varius, Var. Proximus, dd7.
Cygnus equitum, sp. n. (Figs. 1, 2),
427.
Cyligrammea latona, 125.
Cynictis penicillata: structure (Figs. 3,
8, 9), 349.
Cynopterus brachyotis angulatus, 40.
brachyotis, 40.
Cypricercus reticulatus, 339.

309.
CyToLoey.
Protozoa :
309.

Flagellata, Rhizopoda,

Daphnia lumholizi, 334.
Darapsa rosé, 141.

KV

Dasychira miserata, 139.

—— remota, 139.

Dasypus villosus: structure (Fig. 11),
201.

Dattinia costinotalis, sp. n. (Pl. II.
fig. 42), 173.

ornata, 173.

——— peratalis, sp. n. (PI. II. fig. 41),
173.

—— perstrigata, sp. n.
fie. 40), 172.

Dendrogale frenata, 37. °

Dendrohyrax dorsalis: structure (Figs.
12, 13), 202:

Dendrolagus ursinus: structure (Fig. 9),
IM.

DEVELOPMENT.
Pisces: Phallostethine, !.
Insecta: Anoplura, Mallophaga, 288.
Prorozoa: Flagellata,

309.
Diacrisia, var., 102.
—— diversata, 102.

(Pl. IL.

Rhizopoda,

Diaphanosoma excisum, 339.

Diaphone eumela, 104.

Diaptomus anne (P1. LI. fig. 10), 338.

— doriai, 338.

—— strigilipes, 338.

—— viduus, sp. n. (Pl. IT. figs.11-14),
338.

Didelphys albiventris (z. s. u.), 297.

—— aurita (z. 8. L.), 297.

Dipus xgyptius: structure (Fig. 21),
223.

| Discalma calvifrons, sp. n. (Pl. II.

fig. 14), 158.
—— puerilis, sp. n. (Pl. IIL. fig. 15),
157.

| —— subcurvaria, 157.
| Duomitus mesosticta, sp. n. (Pl. IT.
Cypridopsis newtoni (Pl, III. fig. 16), |

fig. 30), 165.
-—— simillima, sp.n. (PI. II. fig. 32),
166.

| ---— steniptera, sp. n. (Pl. II.fig. 31),

166,

Earias insulana, 124.
Kehidna hystrix: structure (Fig. 3),
190.

xvl

Elaphurus davidianus (z.s. u.), 298.

Elephas maximus: structure (Fig. 14),
210.

Elydna bisignata, 108.

Ematurga bilineata, 160.

Endotricha consobrinalis, 171.

Enispa flavipars, sp. n. (Pl. I.
fig. 18), 112.

Ephyra rufistrigata, 148.

Epimys berdmorei magnus,
subsp. n. (Fig. 1), 57.

concolor, 57.

—— griseiventer, 57.

—— jerdoni marinus, subsp. n., 50.

pan, Ol.

rattus, subsp., 50.

—_ _—. klumensis, subsp. n., 56.

——. —— kraensis, subsp. n., 57.

—— —— makensis, subsp. n., 56.

—-— —— Trangensis, subsp. n., 56.

—— rufescens, 50.

—— surifer changensis, subsp. n.,
52.
—— —— connectens, subsp. n., 53.

—— —— eclipsis, subsp. n., 53.

—— —— finis, subsp. n., 51.

—— —— kutensis, subsp. n., 52.

—— —— pelagius, subsp. n., 53.

——. tenebrosus, subsp. n., 54.

Epiphora atbarinus, 164.

Equus granti: structure (Fig. 20), 221.

Estigmene griseata, sp. n. (Pl. I.
fig. 1), 102.

Ethiopica ignecolora, sp. n. (Pl. I.
fig. 10), 108.

—— pheocausta,
fig. 11), 108.

Ernouoey.
Mamuatia: Limnotragus, 375.
Insncra: Lepidoptera, 388.

Eublemma admota, 118.

——. arenostrota, sp.n.(PI.I. fig. 21),
114.

— conistrota, 114.

— eremochroa, sp.n (PIl.I. fig. 19),
113.

— nigrivitia, 113.

(PL. 1.

sp. n.

— ochricosta, sp. n. (Pl. I. fig. 20), |

114.

INDEX.

Eublemma reducta, 1138.

scitula, 114.

Hubolia disputaria, 160.

Eucosma somalica, sp. n., 178.

Eucrostes astigmatica, sp. n.
(PI. II. fig. 22), 146.

rubristicta, 145.

— rufostellata, 146.

Eulocastra argyrostrota, sp. n.,
(Pl. I. fig. 38), 120.

Huphractus vellerosus pannosus (2.8. L.),
448.

Kuphyta
156.

Huproctis fasciata, 139.

Eurycypris subglobosa, 339.

Hustrotia mianoides, 120.

Hutelia discistriga, 124.

—— grisescens, sp. n. (PI. 1. fig. 40),
124.

Euterpiodes croceisticta, sp. n.
(Pl. I. fig. 16), 111.

—— pictimargo, sp. n. (Pl. 1. fig. 15),
TG.

Eutrichophilus setosus: structure (Fig.
17), 283.

Huxoa spinifera, 104.

Euzophera stramantella, 170.

—— villora, 170.

(Camptogramma) natalata,

Featheria obvia, gen. et sp.n. (PI. II.
fig. 33), 167.

Felis: structure (Fig. 1), 443.

caracal (z. 8. L.), 297.

—— eyra (z.8. L.), 297.

ocreatus (z. 8. L.), 297.

—— pardus (z. 8. L.), 297.

salinarum (z. 8. L.), 297.

Funambulus berdmorei, 48.

Galidictis : structure (Fig. 1), 442.
Gallus bankiva: variation, 446.
Gargetta xylochroa, 141.

Gavara leucomera, sp.n. (Pl. II

fie. 36), 168.
Gazella rufifrons (2. s. L.), 441.
Genetta: structure (Fig. 1), 442.

INDEX.

GHOGRAPHICAL.

Manmatta: S.H. Siam, 27; Limno-
tragus: Sesse Islands, 875; Fossils
from Malta, 421.

Aves: Fossils from Malta, 421.

Insecta: Moths from Somaliland,
Oil

CRUSTACEA :
Ceylon, 333.

Geometra (Idea) minorata, 149.

Glossotrophia disparata soma-
liata, subsp. n., 153,

Glyphodes indica, \74.

Gnamptoloma neptunaria, 148.

Gnamptonyx innexa, 127.

Goniocotes sp., 286.

gigas: structure (Fig. 18), 286.

microthoraxr, 285.

verrucosus, 286,

Goniodes bicuspidatus: deyelopment,

structure (Figs. 20-22), 288,

colchicus, 286,

— dispar, 286.

falcicornis: structure (Fig. 19),
287.

megaceros, 287.

minor, 286.

Gonodela obliquilineata, 159.
Grammodes netta, 160,

stolida, 126.

Grison allamandi (z. s. L.), 297.

Entomostraca from

Hemoproteus danilewskyi, 80.

Hellula undalis, 174.

Helogale undulata: structure (Migs. 1,
10), 349. ;

Hemidromodes, gen. n., 144,

robusta, 145,

Hemithea albistrigulata, 144,

malescripta, 144,

—— vermiculata, 144.

Hemitragus jemlaicus (z, s. u.), 447.

Herse convolvuli, 140.

Heterostegane indularia, 156.

Hierochthonia featheri, sp. n.
(Pl. IL. fig. 23), 145.

robusta, 145.

XVll

Hippopotamus amphibius: structure
(Fig. 16), 213.

Hippotion celerio, 140.

ros@, 141.

roseipennis somalicum,
subsp. n., 140.

Hippotragus niger
subsp. n. (Fig. 1), 298.

Hoplotarache czruleopicta, sp.
n. (Pl. L. fig. 37), 121.

—— ectorrida, sp. n. (Pl. I. fig. 36),
121.

nubila, 121.

Hyena hyena: structure (ear) (Fig. 2),
308,

variani,

Hybophthirus notophallus: develop-
ment: (Figs. 7, 8), 267.

Hydrocherus hydrocherus (2. s. L.),
448,

Hylobates agilis, var. pileatus, 29.

pileatus, 29.

Hypena abyssinialis, 136.

Jussalis, 156.

masurialis, 136.

strigata, 136.

Hypotacha indecisa, 125,

Hyrax dorsalis: structure (Figs. 12, 13),
205.

Hystrix bengalensis, 60.

Tambiodes incerta, 1\O-4.
Ichneumia albicauda: structure (Figs.

1, 4, 6, 9), 349.

Idea luridata, 152.
INSECTA.

Lepidoptera: analysis of patterns,
383; Moths from Somaliland :
systematic, 91; Anoplura, Mallo-
phaga: structure, development,

systematic, 253.

Lelia testacea, 139,

Lemobothrium titan, 285.

Laphygma exigua, 109.

Lemur sp.: structure (Fig. 30), 237.
Leptodactylus mystacinus (4. 8. L.), 298.

Proc. Zoou. Soc.—1916, No. XXX. 30

XVill

Lepus sp. (black var.), 441.
Leucinodes orbonalis, 174.
Leydigia australis, 335.

, var. ceylonica (Pl. I. fig. 1),

335.
Limnotragus spekei sylvestris,
subsp. n. (Figs. 1, 2), 375.
Linognathus cavie-capensis: structure
(Fig. 1), 257.
gazella, 260.
limnotragt :
259.

structure (Hig. 2),

pithodes, sp. n.:

(Figs. 5-5), 260.

tibialis: structure (Fig. 6),
267.

Lophorache fulvirufa, 120.

LIudia hansali, 164.

Lyncestis amphizx, 133.

—— diascota, sp.n. (Pl. I. fig.
133.

260,

Macaca andamanensis, 30.

cynomolgus, 31.

wus, Ol.

leoninus, 30.

Macalla purpureopicta, sp. n.
(Pl. II. fig. 39), 170.

Macaria obliquilineata, 159.

semtalhida, 159.

Macropus bennetti (z. s. L.), 447.

Macrorhinus leoninus: structure (Fig.
27), 232.

structure |

50), |

Macroscelides sp.: structure (Fig, 22), |

226.
Macrothrix odiosa, 335.
triserialis, 335.
Maenas arborifera, 102.
Magulaba grisea, sp. n.
fig. +), 138.
MamMauta.
From S.E. Siam: systematic, 27;
Intestinal tract of Mammals: struc-

(P1.

Jt, |

ture, systematic, 183; Hippotragus |

niger variani: systematic, geogra-
phieal, 298; Odocoileus
canus: antlers affected by cancer,
301; Mungotide :

ameri-

INDEX.

MAMMALIA (co7.).
(external characters), 349 ; Limno-
tragus spekei sylvestris: geogra-
phical, ethology, systematic, 379;
Fossils from Malta, 421; Lepus
sp. (black var.), 441; Viverridie,
Hyenidex: structure (alisphenoid
canal), 442.
Manis tricuspis: structure, 200.
Martes flavigula indochinensis,
subsp. n., 3d.
peninsularis, 3d.
Matopo heterochroa, sp. n. (PI. I.
fig. 5), 105.
Maurilia arcwata, 125.
Mecyna gilvata, 175.
Melasina psephota, sp. n., 180.
——- recondita, sp. n., 181.
Menetes berdmorei, 48.
mouhotii, 48,
—— —— rufescens, subsp. n., 50.
UMbrFosus, subsp. n., 49.
Metapioplasta insocia, 122.
Metarbela diodonta, sp. n. (PI. II.
fig. 28), 164.
perstriata, sp. n. (Pl. I. fig. 29),
165.
Metarctia burra, 101.
Mocis repanda, 126.
Moina dubia, 334.
Morrnoxrocy. See Structure.
Moschus moschiferus: structure (Fig.
18); 216.
Mungos auropunctatus: structure, 349.

brachyurus: structure, 349.

gracilis: structure (Figs. 1, 8),

349.

mungo: structure (Fig. 9), 349.

smithii: structure (Figs. 38-5),
349.

Muntiacus muntjak, subsp., 61.

Mus concolor, 57.

grisciventer, 97.

rattus, od.

rufescens, Dd.

_Naarda nigripalpis, sp. n. (PI. IT.

structure

fig. 5), 159.

INDEX.

Negeta lwminosa, 125.
Nemoria stillata, 146.
Neostethus, gen. n., 2.
bicornis, sp. n.: morphology,
development (Fig. 11), 14.
lankesteri, sp. n.: morphology
(Pls. I.-IV.; Figs. 1-10, 12), 2.
Nephopteryx emussatatella, 170.
—— eugraphella, 170.
metamelana, 170.
serratella, 1'70.
Neromia malescripia, 144.
-—-— manderensis, sp. n. (Pl. Il.
fig. 24), 144.
Noctuelia globuliferalis, sp. n.
(Pl. I. fig. 50), 177.
Nodaria externalis, 136.
Nola chionea, 102.
Nomima prophanes, gen. et sp. n.,
7S, Age),
Nomophila noctuella, \74.
Noorda blitealis, 175.
Notodromas oculata (Pl. III. fig. 15),
309.
Notoryetes typhlops: structure (Fig. 4),
190.

Odobenus rosmarus: structure (Fig. 26),
232.

Odocoileus americanus: antlers affected

by cancer, 301.
Odontoretha featheri, gen. et sp. n.
(Pl. L. fig. 7), 105, 106.
(Hdicodia limbata, 116.
melanographa, sp. n. (Pl. I.
. 25), 116.
strigipennis, sp. n.
fig. 24), 115.
Oglasa cornuta, 133.
Oncocypris pustulosa, sp. n.
(Pl. LL. figs. 17-21), 340.
Ophideres fullonica, 156.
Orcella brevirostris (Fig. 2), 65.
Ornithorhynchus anatinus: structure

fi

OS

ie

(Fig. 2), 189. |
Orycteropus capensis: structure (Vig.
10), 109.

|

abe

Osteodes procidata, 157.

eritreénsis, 157.

turbulentata, 157.

Ozarba consanguis, 117.

endoplaga, sp. n. (Pl. I. fig. 32),
119.

—— endoscota, sp. n. (PI. I. fig.
INE

—— exolivacea,sp.n. (Pl. 1. fig. 30),
118.

hemimelena, 118.

-—-hemipyra, sp.n. (Pl. I. fig.
117.

—— hemisarca, sp. n. (Pl. I. fig. 29),
118.

—— mesozonata, sp.n. (PI. I. fig.
31), 119.

phea, 120.

27),

28),

—— sancta, 120.
semitorrida, sp. un.
fic. 26), 116.

ds

(QL

2

Pachycoa olivacea, gen. et sp. n.
(BI Tete. 12), 1095110:

Pachypalpia subalbata, 163.

Pachyzancla basalis, 175.

bipunctalis, 175.

pheopteralis, 175.

Pandesma anysa, 128.
Papio leucophzeus (z. s. u.), 447.
Paradoxurus hermaphroditus, 33.
— minor kutensis, subsp. n., 34.
pallasii, 33.
Parallelia algira, 126.
rectifascia, 126.
Paratuerta nana, sp. n.
fice le) Wha
Paryphanta arcuilinea, 168.
jfimbriata, 168.
Pasipeda sambesita, 138.
PaTHOLOGY.
Deaths in the Society’s Gardens, 77.
Pedicinus rhesi, 256.
Pediculus affinis, 255.
capitis, 255.
Pericyma metaleuca, 127.

(Blee

Perigea capensis, 104,

xXx INDEX.

Phalena Geometra selenaria, 163.

Phalena pulchellata, 151.

Phallostethus dunckeri: morphology
(Pls. I., IIT., 1V.; Figs. 12-15), 16.

Phascolarctos cinereus: structure
(Fig. 8), 193.
Phascolomys mitchelli: structure

(Figs. 6, 7), 193.
Phiyctenodes nudalis, 175.
Phthirpedicinus miecrops, 256.
Puysronoey.
Mamata: Mungotide (anal sac),
349.
Pisces: Phallostethine, 1.
Physocypria tuberata,
(PL. IDL. fig, 22: Hig. 1), 341-
Phytometra acuta, 128.
~— limbirena, 128.
mi, 128.
Pionea melanostictalis, sp. n.
(Pl. II. fig. 46), 175.
—— rubritinctalis, sp. n. (Pl. I.
fig. 45), 175.
PIscys :

sp. Nn.

Phallostethinz : morphology: sys-
tematic, 1; Tilapia nilotica athi-
ensis: variation, systematic, 345;
Trygon pastinaca ‘poison-organ),
451; Veleoste1: structure, 448.

Pithecus fascicularis, 31.

Plasmodium kochi, 85.

—— precox, 85.

Plecoptera hypoxantha, 135.

— polymorpha, sp. n. (Pl. II.

fig. 3), 134.

Poliana micra, 140.

Polydesma colutrix, 128.

Polyplax spinulosa, 256.
Prasinocyma perpulverata,

sp. un. (Pl. IL. fig. 25), 148.

—— perscripta, ab. n., 143.
subfasciata, ab. n., 143.
Preshytis germaini, 32.

— mandibularis, subsp. n.,

32
Proconis abrostoloides, 128.

Proteles cristatus : structure (Fig. 28), |

235.

Protozoa:

Parasites in the Society’s Gardens,
84: Flagellata, Rhizopoda: cy-
tology, structure, systematic, 309.

Prowazekia (Bodo) saltans: structure,

development, (Pl. I.), 311.

Pseudosterrha gayneri, 156.

philearia, 156.

Pteropus hypomelanus condorensis, 38.

medius: structure, 230.

vampyrus mataccensis, 39.

Ptychopoda sp., 155.

aperta, sp. n. (Pl. Il. fig. 16),
155.

—— nigrosticta, 155.

—— subtorrida, sp.n. (PI. II. fig.17),

154.

Pyrausta conistrotalis, sp. n.

(Pl. II. fig. 48), 176.

incoloralis, 176.
—— sthenialis, sp. n. (PI. II. fig. 47),

176.

Rabila albiviridis, sp. n. (Pl. I.
fig. 18), 109.

Rana temporaria: variation (Fig. 1),
87.

Ratufa melanopepla, 43.

leucogenys, subsp. n., 43.

—— —— sinus, subsp. n., 44.

Rhamphocelus icteronotus (z. Ss. L.),
297.

Rhinopoma macrophyllum: structure
(Fig. 24), 229.

Rhodoneura hamatipex, sp. n.
(Pl. LI. fig. 37), 168.

Rhopaloceras stylifer: structure (Figs.
23, 24), 292.

Rhynchina albiscripta, sp. n.
(Pl LL. fig. 8), 137.

—— endoleuca, sp. n. (Pl. II. fig. 6),
137.

——perangulata, sp. n. (Pl. I.
fig. 7), 136.

revolutalis, 137.

Rothia aisha, 108.

tucervus schomburghki, 62.

Rusa peronii, 62.

INDEX.

Sameodes ocellata, sp. n. (Pl. II.
fig. 44), 174.

Sarmatia interitalis, 136.

Sceliodes laisalis, 176.

Sciurus albivexilli, sp. n. 47.

bicolor, 43.

—— cinnamomeus, 40.

—— ferrugineus, 45.

—— -— cinnamomeus, 40.

frandseni, subsp. n., 46.

—— finlaysoni, 45.

Javensis, 43.

—— macclellandi rodolphi, 48.

—— mouhotii, 48.

—— pyrrhocephalus, 48.

-— rodolphi, 47.

—— splendens, 45.

Scotinochroa minor, sp. n. (PI. I.
fig. 34), 168.

Scotosia natalata, 15.

—- rubritincta, 156.

Scrancia discomma, sp. un. (Pl. Il.
fig. 10), 141.

Secusio somaliensis, sp. n. (Pl. 1. |

fio. 2), 102:

strigata, 103.

Selagena atridiscata, 104.

eustrigata, sp. n. (Pl. II.
fie. 27), 164.

Semiothisa butaria, 160.

obliquilineata, 159.

Semnopithecus germaini, 32.
Sesamia coniota, 119.
Sesquialtera, gen. n, 162.

= ridicula, sp: n. (Pl. il. fig. 1);

163.
Siccia sordida, 102.
Simplicia capalis, 136.
Siphonops annulatus (4. s. L.), 447,
448.
Sminthopsis crassicaudata :
Igile
—— larapinta: structure, 191.
Sphingomorpha chlorea, 133.
Sphinx celerio, 140.
convolvuli, 140.
Spironema multiciliatum :
(PI. IIT. figs. 46-48), 318.
Staudingeria sub-oblitella, 170.

structure

structure, |

Slegania indularia, 156.

secularia, 157.
Stenocypris malcolmsoni, 339.
Stenostaura tmpeditus, 141.
Stenosticta grisea, 124.
Sterrha philearia, 156.
Strandesia (Cypris) vittata, 389.
Strix delicatula (zs. rs), 448.
Strongylocotes coniceps, 286.
STRUCTURE.
Mammatta: Intestinal tract, 183;
Mungotids (external characters’,
349; Viverride, Hyxnide (ali-
sphenoid canal), 442.
Pisces: Phallostethinz, 1 ;
pastinaca (poison -organ),
Teleostei, 448.
Insucta: Lepidoptera (analysis of
patterns), 883; Anoplura, Mallo-
phaga, 253.

Trygon
folly,

Prorozoa: Flagellata, Rhizopoda,
309.
Suraitha tnvectellus, 169.
scitulellus, 169.
Suricata suricatta, structure (Figs. 3, 4,
8-10), 319.

Sus cristatus, (4.

——- —— subsp., 64.
Sylepta sabinusalis, 174.

Talpa europa: structure (Fig. 28),
226.

Tamandua tetradactyla :
20M

Tamiops novemlineatus, 48.

rodolphi, 47.

‘arache carnescens, 125.

structure,

hortensis, 125.

—— mesoleuca, sp. n. (PI. I. fig. 38),
Ts,

miogona, sp. n, (Pl. I. fig. 39),
128.

—— opalinoides, 128.

umbrigera, 123.

——- gelleri, 128.

Tathorhynchus exsiccata, 135,

Tegostoma bipartalis, 177.

comparalis, 177.

Proc. Zoou. Soc.—1916, No, XX XI. eal

Xi

Tegustoma subditalis, 177.
egulifera nigricinetalis, 171.
zonalis, 171.
Tephrias trigonosema,
(Pl. IL. fig. 2), 134.
Tephrina butaria, 160.
cinerascens, 160.
deerraria, 161.
—— disputaria, 160.
—— inconspicua, 160.
netta, 160.
prionogyna,
fig. 18), 161.
——— subeurvaria, 157.
Teracotona submacula, 102.
Tetramitus spiralis, sp.n.: struc-
ture, development (Pls. IL., III.
figs. 42-45 ; Fig. 1), 314.

(PL. IL

sp. n.

Thalatha melanostrota, sp. n.
(Pl. I. fig. 4), 104.
Thylacinus cynocephalus: structure

Gigs) LOM

Tilapia nilotica athiensis, var. n.,
345.

Timandra neptunaria, 148.

viridaria, 148.

Toana nigrilineata, sp. n. (Pl. I.
fig, 22), 114.

Tragulus affinis, 63.

javanicus, 63.

-—— kanchil, 63.

—— —— affinis, 63.

prerret, 60.

Traminda neptunaria, 148.

—— rufistrigata, 148.

Tricentroscelis protrusifrons,
gen. et sp. n. (Pl. LI. fig. 21), 147.

Trichiura obsoleta, 166.

Trichodectes sp., 283.

—— breviceps, 271.

cornutus, 273.

crassus, 272.

INDEX,

Trichodectes harrisoni, sp. n.:
strueture (Figs. 13-16), 276.

—— hemitragi, sp. n.: structure
(Figs. 11, 12), 273.

— latus: structure (Fig. 10), 271.

— ovis, 283.

parumptlosus, 271. :

Trigonomelea semifusca, 163.

Trygon pastinaca: structure (poison-
organ) (Pigs. 1-7), 431.

Tuerta trimeni, 103.

Tupaia belangeri, 36.

concolor, 36.

—— —— sinus, subsp. n., 36.

—- frenata, 37.

Tyndis proteanalis, 171.

Ulothrichopus tinctipennis, 125.
Utetheisa pulchella, 102.

VARIATION.
Manmaura: Lepus sp. (black yar.),
441.
Aves: Gallus bankiva, 446.
Barracuia: Rana temporaria, 87.
Pisces: ‘Vilapia nilotica (anal spines),
345.
Victoria sematoperas, sp. n.
(P1. IL. fig. 26), 142.
Viverra hermaphrodita, 33.
Viverricula: structure (Fig. 1), 442.

Zamarada pulverosa, 157.

—-— secutaria, 157.

Zinckenia fascialis, 174.

ZLitha subcupralis, 171.

Zygophyxia tornisecta, sp. n.,
153.

PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STREET.

PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCIETY
OF LONDON,

1916.

PAR.

CONTAINING Paces 1 To 3807, witH 7 PuaTEs

AND 75 TExtT-FIGURES.

MARCH 1916.

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

MESSRS. LONGMANS, GREEN, AND CO.,
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|
On [Price Twelve Shillings.] e

LIST. OF CONTENTS:

1916, Parr I. (pp. 1-307).

EXHIBITIONS AND NOTICKS.

Page

The SucreTary, Report on the Additions to the Society’s Menagerie during the months
of November, December, and January, 1915-6 (...... 1.1 cece ee ce cece ee ee cere 2
Mr. R. BE. Houpine. Exhibition of the Skull of a Roebuck ..--.... +++. -+1e-+ sees pone Age,

Mr. C. Tats Rucan, M.A., F.Z.S. Lantern exhibition of drawings of larval Fishes .... 298

Mr. Oupriztp Tuomas, F.R.S., F.Z.S. A new Sable Antelope from Angola, (Text-
eure) erat NAR cta HOD OO bo DOG So ute Ager dic Ode Fel cliesoy drove eater 298

Mr. R. I. Pocock, F.R.S., F.Z.S., Curator of Mammals. Antlers of a Virginian Deer :
afiected by Cancer sic 2 vise) aici cielare + ipo slalecetstnisusrs ca eeel Gale velet-tetalerehetctaie! t-test ae 301

The Rev. H. N. Hurcuiyson, M.A., F.Z.8. Exhibition of drawings of extinct animals .. 802

Mr. C. Tarr Ragan, M.A., F.Z.S. Lantern exhibition of a Siamese Fighting-Fish and of
PV OPiS non oomaomnee Go OOMEOrorGn NovoAS Gao aicodnb.ads.cudd sanbocODOGCn ca 302

Mr. R. I. Pocock, F.R.S., F.Z.S., Curator of Mammals. The Tympanic Bulla in Hyzenas.
(GIGS fairies Ap Pa AAP Ar ccmejon: cisandnomooualamacadododouon son: 303

Mr. Harry K. Eustace. Bioscope exhibitio: of African Animals ........ ........... 307

Contents continued on page 3 of Wrapper.

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-_-—>-—

PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCIETY OF LONDON

PAPERS.

1. The Morphology of the Cyprinodont Fishes of the
Subfamily Phallostethinee, with Descriptions of a new
Genus and two new Species. By C. Tatrt Ruean,

Mea, ZnS.
[Received November 8, 1915: Read November 23, 1915.]

(Plates I.-IV.* and Text-figures 1-15.)

INDEX.

MorrHoLoey: Page
INCOSCEERWS LOMKESTOTO) pate e eee eee ee,
ENERLAMIGESECTUAT Ge saeco EE eee) ul
IN. bicornis ..... Saas pecan Alte nto acta coe, ae LAL
Phallostethus dunckeri i 18 Peete aucicatanes: eat acetate ame oan! |)
P. dunckeri, 3... . Pea B secrets Ar ORES a oe Coe ae
sep ria piumnitiert tbe hoes ae een ee ea a einen ce 20
Other copulatory organs of fishes ......................... 28

PuystoLtoey: Use of the priapium .......... compe

DEVELOPMENT—of the priapium of iyeoate nies Bicornis, eas 14

SYSTEMATIC:

Neostethus lankesteri, gen. et sp. n. Reece phir

NV. bicarnis, sp. n de eames acetates

2ank and position of the Phallostethine .................. 23
Eyotutrron—Origin of skeletal elements........................... 24

1. Introduction.

In 1913 I described an extraordinary little Cyprinodont fish
from Johore, and named it Phallostethus dunckeri (Regan, 11).
Some more fishes from the same locality have been sent to me

* For explanation of the Plates see p. 25.

Proc. Zoou. Soc.—1916, No. I. 1

2, MR, C. TATE REGAN ON THE

for description ; they represent a new genus, Veostethus, related
to Phallostethus, and belong to two new species, JV. lankesteri
and JV. bicornis.

Of Neostethus lankestert there are six specimens, all about
30 mm. in total length; five are adult males and the other is an
adult female. They come from the Muar River (brackish-water)
and from Singapore. Some features in their structure, such as
the number of vertebre and the general relations of the viscera,
have been elucidated by prolonged clearing with oil of cloves, but
the detailed account of the structure of the male fish is based on
the study of a series of transverse sections. Of JV. bicormis there
are three examples, two males (21 and 25 mm. long)—one imma-
ture, the other nearly adult—and a female of 24 mm.; these are
from Kuala Langat (brackish-water),

Phallostethus is redescribed and is compared with WVeostethus.

2. Structure of Female NeosTEYHUS LANKESTERI *, gen. et sp. n.

a. EXTERNAL CHARACTERS.

Form elongate, strongly compressed. Head rather small ;
mouth terminal, strongly oblique, protractile, with one or two
series of conical teeth in the jaws; eyes large, lateral. Scales
very similar in structure to those of Panchax; 34 to 36 in a
longitudinal series. Dorsal fin of 5 or 6 rays, above the end of
the rather long anal, which has 15 or 16 rays; caudal emar-
ginate; pectorals 10 or ll-rayed, placed rather high. Anus
{text-fig. 12, B,a.), genital aperture, and urinary opening behind
each other in middle line below bases of pectoral fins; behind
~them abdomen compressed to an edge bearing a rayless fringe
(text-fig. 12, B, f.); just behind anus a pair ‘of papillee (text-
fig. 12, Be p. ) (2 Sagal pelvic fins tT), one much larger than the
other, that partly cover a depression into which the oviduct and
ureter open.

6. SKELETON.

The skeleton is typically Cyprincdont and essentially similar
to that of Panchax, except that the hemal arches of the caudal
vertebrae are not expanded, as the air-bladder does not extend
back into the tail. The vertebree number 34 or 35 (15-16+19).

c. VISCERA.

. ‘
The air-bladder occupies the posterior part of the abdominal
cavity ; it is large, simple, and thin-walled, except an anterior

* T have ventured to name this species in honour of Sir Ray Lankester, K.C.B.,
F.R.S., to whom J am indebted for many acts of kindness and much sound advice.
Moreover, it seems to me not inappropriate that this little fish, whose structure
presents more than one problem tor the consideration of students of animal mor-
phology, should bear the name of the most distinguished mor phologist of our time.

+ In Phallostethus the postanal papillae are supported by a pair of minute
skeletal elements that may be vestigial pelvic bones.

MORPHOLOGY OF CYPRINODONT FISHES. 3

part, tapering forwards, that has thick glandular walls. ‘There
is no trace of a pneumatic duct.

The short cesophagus leads into the stomach, which is simple,
without cecum or pyloric appendages; the intestine forms a
single coil just in front of the air-bladder, and then runs down-
wards and forwards to the anus. The liver is large, and there
is a well-developed spleen situated on the anterior part of the
intestinal coil.

The kidneys extend from below the basioccipital to the posterior
end of the abdominal cavity; they are paired, but not enlarged,
anteriorly, unpaired posteriorly ; the ureters leave the kidneys

Text-figure 1.

ae if FE
ae —
NN

Neostethus lankesteri, 2. Part of head and abdominal region cleared and viewed
as atransparent object (X 10). ‘The ovary is clearly visible, lying in front of
the air-bladder and below the alimentary canal.

h., heart; a., anus; p., postanal papille; o., opening of the oviduct; w., opening of
ureter.

above the anterior part of the air-bladder and soon unite to
form a single duct that runs downwards and forwards below
the intestine. The ovary is unpaired and lies in front of the
air-bladder and below the intestine; it narrows forwards, and
the very short oviduct arises from its anterior end. The ova are
comparatively few and large.

Except for modifications correlated with the thoracic position
of the anus and urino-genital apertures, e. g. that the air-bladder
is posterior instead of superior, the intestine runs forwards instead
of backwards, etc., the visceral anatomy is essentially similar te

that of the Funduline,
1*

4 MR. C. TATE REGAN ON THE

3. Structure of Male NEosTETHUS LANKESTERI.

a. EXTERNAL CHARACTERS.

The male differs from the female externaily in the presence of
the priapium (text-fig. 2), a fleshy appendage that lies below the

Text-figure 2.

Neostethus lankesteri, $. Head and priapium (X 10). A, proctal side;
B, aproctal side.
et., ctenactinium; ctm., ctenactinial muscle; a., anus; 7., end of priapial rib;
v.d., terminal coil of vas deferens; pv.; pulvinulus; pva., pulvinular ap-
pendage ; s., pulvinular spine; g., glandular groove; ef, efferent groove ;
sp., seminal papilla; p., infrasulcar prominence.

head and the anterior part of the body, to which it is attached
for the greater part of its length, only the posterior end being
free. Anteriorly the priapium is confluent with the isthmus ;

MORPHOLOGY OF CYPRINODONT FISHES. 5

further back it increases in size and is well-defined ; owing toa
strong constriction of the body just behind its attachment, the
posterior part of the priapium projects but little beyond the
general outline of the abdominal region.

On one side, the proctal side, which may be either right or
left, may be seen the anus (a.); above and behind it can be seen
the outline of the enlarged terminal coil of the vas deferens (v.d.),
occupying most of the free posterior part of the priapium, which
ends in a membranous fringe, produced into some. half-dozen
slender processes. On the proctal side a shallow groove marks
the boundary between the priapium and the body of the fish, but
on the other side, the aproctal side, there is a much deeper groove,
margined above by a thick fold of the integument; this groove is
lined by a glandular epidermis, and may be termed the glandular
groove (g.). Posteriorly a dermal fold arises from the inner wall
of the groove, and this fold is continued backwards on the free
part of the priapium as the roof of another groove, leading from
the glandular groove to the end of the priapium ; this may be
termed the efferent groove (éf.).

The enlarged part of the vas deferens lies below the floor of the
efferent groove; here it is running backwards, and at the end of
the priapium it curves round from the proctal to the aproctal side
and then runs upwards and forwards, ending in a seminal papilla
(sp.), which opens into the glandular groove, the terminal aper-
ture being a wide slit. Directly in front of the seminal papilla
is a papilliform projection (p.), which may be termed the infra-
sulear prominence. Below the infrasulcar prominence is the
articulation of the ctenactinium (ct.), a long and slender movable
bony appendage that curves backwards and upwards, then for-
wards to below the eye, and, finally, downwards and across beneath
the chin ; a short pointed process, directed outwards and down-
wards, arises from its concave edge above the infrasulcar
prominence.

Further forwards, a rather soft appendage, subconical in form
and with its apex directed backwards, lies in the glandular groove ;
at its base it is separated by a deep constriction from a lateral
mass of tissue that tapers forwards to the anterior end of the pri-
apium. ‘This mass of tissue may be termed the pulvinulus ( pv.)
and its appendage the pulvinular appendage (pva.); a small
antrorse spine (s.) projects from the posterior part of the pul-
vinulus, and a branch of the glandular groove runs forwards
between the priapium proper and the lower part of the pul-
vinulus; this infrapulvinular groove narrows forwards and
disappears a little in advance of the level of the pulvinular
spine.

b. SkEvETON (PI. J. B, and text-fig. 3).

This differs from that of the female in that one of the cleithra,
the third vertebra, and the first pair of ribs are modified in

6) MR. C. TATE REGAN ON THE

connection with the priapium, whilst the priapium itself has a
special skeleton.

Cleithra.—The cleithrum of the proctal side is normal and ends
anteriorly below the angle of the preoperculum. ‘That of the
aproctal side is produced into a slender process (¢el.) that extends
forward to below the attachment of the urohyal; this process lies
between the isthmus and the priapium ; it is somewhat expanded
transversely (PI. I. A), and anteriorly it spreads downwards on
each side of the priapium, almost enclosing it.

Text-figure 3.

Neostethus lankesteri. Skeleton of priapium from the aproctal side (x 12)
(diagrammatic).

cta., ctenactinium; w., urohyal; cl., cleithrum; ¢7., ¢7.’, transverse processes of
third vertebra; c., cartilage; 7., 7°’., first pair of ribs; ae., antepleural carti-
lage; va., pulvinular appendage; pus., pulvinular spine; pv., outer, and
pv.’, inner pulvinular bones; és., anterior, and is.’, posterior infrasulcar bones;
p., papillary bone; a.7., vertical, and a.r.’, horizontal anterior ridges of axial
bone: 77, its lateral ridge; cv., its main crest; and s¢., its terminal style.

Third vertebra.—The transverse process of the aproctal side (¢7".)
is normal, but that of the proctal side (é#.) is much stronger ;
proximally it is directed outwards at right angles to the centrum,
then it runs forwards below the transverse process of the second
vertebra (text-fig. 4B), until it reaches the level of the first
vertebra, when it curves downwards and ends.

First pair of ribs.—These are attached proximally to the
transverse processes of the third vertebra; the rib of the aproctal
side (7.') is not particularly strong, but it is very long and runs
downwards and forwards into the priapium; the rib of the
proctal side (7.) is much stouter and runs downwards right to
the ventral surface of the priapium below the anus (text-
fig. 2, 7.); proximally a nodule of cartilage (c.) intervenes

MORPHOLOGY OF CYPRINODONT FISHES. 7

between the end of the rib’and the posterior face of the de-
curved part of the transverse process. A mass of cartilage, the
antepleural cartilage (ap.), lies in front of the distal ends of this
pair of ribs and embraces them laterally; this mass is to a large
extent composed of parenchymatous cartilage, but in the centre
approaches true hyaline cartilage in structure (cf. text-fig. 4, B).

Axial bone of the priapium.—This extends nearly the whole
length of the priapium ; in front of the articulation of the cten-
actinium it has the form of a rod of cartilage enclosed in a cylinder
of bone (PI. IIT. A, a.) that bears certain crests and ridges, namely,
(1) the anterior ridges, the upper (a@.) vertical, the lower (ar.')

Text-figure 4.

Neostethus lankesteri, 6. ‘Transverse sections (x 18): A, through postorbital
part of head and base of pulvinular appendage ; B, through second vertebra and
antepleural cartilage.

ao., aorta; g., glandular groove; pva., pulvinular appendage; a., axial bone;
is., infrasulear bone; e¢m., ctenactinial muscle; pm., muscle of proctal side;
apm., muscles of aproctal side; ppm., pleuro-priapial muscle; ac., antepleural
cartilage; +., priapial rib; 7sc., cartilage; ¢7., transverse process of third
vertebra; &., kidney; c., cesophagus; sv., sinus venosus.

nearly horizontal and aproctal; these increase in height back-
wards and end abruptly at the level of the pulvinular spine.
(2) The main crest (cr.): this rises obliquely from the proctal
side of the axial bone and then curves upwards until it is
vertical (text-fig. 4 A, a@.); it commences a little behind the end

8 MR, C. TATE REGAN ON THE

of the anterior ridges and ends in front of the priapial ribs.
(3) The lateral ridge (Jr.), on the aproctal side from the level
of the end of the pulvinular appendage to the level of the priapial
ribs ; this ridge almost reaches the surface at the lower margin
of the glandular groove. The axial bone passes backwards on the
aproctal side of the priapial ribs and antepleural cartilage, and in
the region of the anus it becomes stouter and expands down-
wards; the ctenactinium (cta.) is attached to the aproctal side
of this part of the axial bone. In the region of the seminal
papilla the axial bone extends upwards nearly to the glandular
groove, and behind this it contracts to form a terminal style (sé.)
that runs backwards and ends just in front of the transverse
portion of the terminal coil of the vas deferens.

Anterior infrasulcar bone (is.).—This is a laminar bone that
lies near the surface on the aproctal side. It is broad posteriorly
and tapers anteriorly ; behind it extends from the floor of the
glandular groove under the base of the infrasulcar papilla to out-
side the proximal end of the ctenactinium ; for the greater part
of its length its upper edge is just below the edge of the lateral
ridge of the axial bone: anteriorly it hes within the inner surface
of the infrapulvinular groove (PI. II. B, zs.) and is quite a slender
bone. Near its posterior end, where its lower edge overlaps the.
end of the ctenactinium, it bears an inner knob to share with the
axial bone in supporting that appendage.

Posterior infrasulear bone (is.')—A bone whose expanded
upper surface lies just below the floor of the glandular groove,
below the anterior part of the seminal papilla; it is a solid bone
that extends downwards and inwards on the aproctal side of the
axial bone: from its upper surface it sends forwards a laminar
process that runs below the anterior infrasulear bone into the
base of the infrasulecar prominence.

Papillary bone (p.).—A bone that supports the seminal papilla,
which it enters from behind and below, and then divides into
three branches that expand into lamine lying just below the
skin, one on the inner side of the papilla and two, an upper and
a lower, on its outer side. Before entering the papilla the bone
is a slender rod that curves downwards across nearly to the
ctenactinium and then tapers forwards and inwards, ending a
little in advance of the seminal papilla on the aproctal side of the
axial bone (cf. Pl. II. A, and text-fig. 7 A, p.).

The pulvinular appendage (pva.) is subconical, somewhat com-
pressed ; it is a mass of parenchymatous cartilage, but has the
structure of true hyaline cartilage on the inner side near the
base; in this region it is hollowed out for the reception of a
cartilaginous peg that arises from a bone in front of it (Pl. II. B).

The pulvinulus consists of fibrous connective tissue sur-
rounding two longitudinal bones, the inner and outer pulvinular
bones.

The inner pulvinular bone ( pv.') is largest posteriorly, where

MORPHOLOGY OF CYPRINODONT FISHES. 9

it bears a backwardly directed process for the support of the
pulvinular appendage; it tapers anteriorly and ends below the
axial bone at about the middle of the length of the anterior
ridges. This bone is rounded or oval in cross-section, posteriorly
deeper than long; it has a cartilaginous core.

Text-figure 5.

Neostethus lankesteri, 6. ‘lransverse sections cutting the infrasulcar prominence,
A, near its anterior end, and B, at its posterior edge, also just cutting the
seminal papilla (X 18).

&., kidney; 1., liver; sv., sinus venosus; @., cesophagus; 7., intestine; w., ureter;
v.d., vas deferens; pm., muscle of proctal side; apm., inner muscle of aproctal
side; s., seminal papilla; ip., infrasulear prominence; is., infrasulecar bone;
a., axial bone; c¢., ctenactinium.

The outer pulvinular bone (pv.) is rather similar to the inner
in form and structure (PI. IIT. A, pd.); posteriorly it bears the
pointed antrorse process which appears externally as the pulvi-
nular spine; it runs forwards outside the inner bone, but in front
of the end of the latter becomes median and ventral, and extends
forwards in front of the end of the axial bone to the extreme
anterior end of the priapium.

10 MR. C. TATE REGAN ON THE

c. VIscERA (text-fig. 6).

Air-bladder (a.), alimentary canal, kidneys (4.), etc., as in the
female, except that the intestine (7.) and ureter (w.) end in
the priapium.

The intestine (7.) enters the priapium near the posterior end
‘of its junction with the body (text-fig. 5), and runs downwards,
somewhat forwards, and across to the proctal side, ending at the
anus.

The ureter (w.) enters the priapium just below and behind the
intestine (text-fig. 5 B) and runs downwards into the middle of
the priapium and across until it meets the enlarged part of the
vas deferens, here running backwards on the proctal side; the
ureter now runs upwards and backwards and towards the proctal
side, always in contact with the vas deferens, and ends by opening
into the efferent groove, not far from the proximal end of the
latter (text-fig. 7 A).

Text-figure 6.

Visceral anatomy of Neostethus lankesteri, 8 (X 8). From the proctal side;
the liver, etc., removed.

@., esophagus ; s., stomach; z., intestine; &., kidney; w., ureter; ¢., testis;
vd., vas deferens; a@., air-bladder.

The testis (¢.) is unpaired (text-fig. 8 A) and corresponds to the
ovary in form and position ; the vas deferens (v.d.) arises from the
middle of the upper surface of the testis and runs forwards, at
first at the side of the intestine and then partly above it; it
curves downwards to enter the priapium just behind and on the
aproctal side of the intestine, and runs downwards and a little
backwards until it reaches the terminal style of the axial bone ;
the vas deferens runs across to the proctal side below this bone
and then expands to form the large terminal coil that runs back-
wards, then across, and then upwards and forwards, ending in
the seminal papilla.

MORPHOLOGY OF CYPRINODONT FISHES. im)

The vas deferens has a thin wall, comprising an outer fibrous
layer and an inner layer of mucus-secreting cells; within the
testis the tubules that unite to form the vas deferens have a similar
structure (PI.IV.C). The epidermis of the glandular groove also
secretes mucus, and in the neighbourhood of the seminal papilla
and on the papilla itself the area of secreting-cells is increased
by folding.

Text-figure 7.

Neostethus lankesteri, 8. Transverse sections (X 18): A, passing through the
posterior part of the seminal papilla, and B, through the priapium near “its
posterior end.

k., kidney ; U., liver; s., stomach; 7., intestine; v.d., vas deferens; w., ureter ;
a., axial bone; p., papillary bone; cé., ctenactinium, e., efferent groove.

The tubules of the testis, the vas deferens, and the glandular
groove in the neighbourhood of the seminal papilla hold a mucus
secretion, in which appear numerous spermatophores; these are

12 MR. CG. TATE REGAN ON THE

subspherical, with the heads of the spermatozoa at the periphery
and their tails curled round inside (text-fig. 9).

Text-figure 8.

Neostethus lankesteri, §. ‘Transverse sections (X 18): A, passing through testis ;
b, through air-bladder.
k., kidney; J., liver; 7., intestine; ¢., testis; v.d., vas deferens ;
a., air-bladder ; r., rib.

Similar spermatophores have been described in the Peeciliine
by Philippi (Philippi, 6), but I have not been able to detect
them in other Cyprinodonts.

d. Muscuxs (text-fig. 10).

Pleuro-priapial muscle (pp.).—It has been mentioned that the
first pair of ribs enter the priapium, and that the rib of the proctal
side is much enlarged and is attached proximally to the enlarged

MORPHOLOGY OF CYPRINODONT FISHES. We

Text-figure 9.

aN

“Hi((

WE

o
4
ff
6
‘1

4

oh See

Spermatophores of Neostethus lankesteri (X 750

a=

and forwardly directed transverse process (é.) of the third ver-
tebra. To this process is attached also the proximal end of a
muscle that runs downwards into the priapium on the inner side

Text-figure 10.

Neostethus lankesteri. Muscles of priapium, from the aproctal side (Xx 20).

t., transverse process of third vertebra; ct., base of ctenactinium ;*aa., axial bone;
pp., pleuro-priapial muscle: p., longitudinal muscle of proctal side’; ap., outer,
and ap.’, inner longitudinal muscles of aproctal side; ctm., ctenactinial muscle.

14 MR. C. TATE REGAN ON THE

of the rib, and then backwards on the proctal side of the axial
bone, ending in a nodule of bone that lies near the axial bone at
the level of the articulation of the ctenactinium.

Longitudinal muscles of the priapium.— These are four in
number and each,is more or less fusiform in shape. In front
of the priapial ribs these muscles, with the intermuscular connec-
tive tissue and the axial bone, constitute the whole body of the
priapium (excluding the pulvinulus) (text-fig. 4 A).

Ctenactinial muscle.—This is the largest of the longitudinal
priapial muscles; it originates at the posterior end of the upper
anterior ridge of the axial bone and is inserted on the proximal
end of the ctenactinium. For most of its length its inner surface
lies against the whole proctal face of the main crest and the lower
face of the lateral ridge of the axial bone.

Outer muscle of the aproctal side.—This originates at the
anterior extremity of the axial bone and runs back on the
aproctal side between the anterior ridges of that bone, and then
on the aproctal side of the main crest until the inner muscle
intervenes; it ends posteriorly at the level of the anus in the
connective tissue that lies between the floor of the glandular
groove and the lateral ridge of the axial bone.

Inner muscle of the aproctal side.—Anteriorly this is inserted
between the outer muscle and the main crest of the axial bone;
it runs back in contact with the crest and above the lateral ridge,
‘and then on the aproctal side of the priapial ribs, pleuro-priapial
muscle, and intestine; it is attached posteriorly to the lower and
proctal surface of the terminal style of the axial bone just above
the vas deferens, which is here crossing to the proctal side below
the axial bone.

Muscle of the proctal side.—This originates on the anterior
end of the upper edge of the main crest of the axial bone, and
runs backwards at first above and then at the proctal side of the
ctenactinial muscle; it lies on the proctal side of the priapial ribs
and intestine and behind them at the side of the inner aproctal
muscle; it ends in the connective tissue that lies above the ter-
minal style of the axial bone and between the descending portion
and the enlarged terminal part of the vas deferens.

4, NEOSTETHUS BICORNIS, sp. n. (text-fig. 11).

This species is more slender than JV. lankesteri and has 36 ver-
tebre (16-17+ 19-20) instead of 34 or 35. There are 13 to
15 anal rays, and 35 to 37 scales in a longitudinal series.

A male of 25 mm. is not fully adult, but its priapium (text-
fig. 11, B) differs from that of WV. lankesterit in three important
characters: (1) there are two ctenactinia (ct.), both on the
aproctal side; (2) the efferent groove (éf.) extends downwards
to the ventral surface of the posterior end of the priapium; and
(3) the seminal papilla (p.) opens into the efferent, not the
glandular groove. The ctenactinia have not attained their full

MORPHOLOGY OF CYPRINODONT FISHES. 15

development and are cartilaginous and enclosed in skin; their
final shape is uncertain; the testis (¢.) is fairly large.

A male of 21 mm. (text fig. 11, A) has the priapium but little
developed and without trace of ctenactinia, but showing a differ-
entiation into an anterior muscular and a posterior visceral
portion ; in this fish the testis is quite small.

Text-figure 11.

Neostethus bicornis : immature males ; head and priapium from aproctal side (X10).
Total length of fish: A, 21 mm.; B, 25mm. The testis (¢.) is shown separately.

pv., pulvinulus; pva., pulvinular appendage ; c¢., ctenactinia; p., seminal papilla;
ef., efferent groove.

A female of 24 mm. is very similar to the female 1. lankesteri,
except for the more slender form; the postanal depression is less
developed than in JV. lankesteri, probably because the specimen is
not fully adult.

The male examples are of interest as indicating that the pri-
apium develops only as maturity approaches ; presumably males,
only a little smaller than the smaller one, would be almost indis-
tinguishable from immature females.

16 MR. C. TATE REGAN ON THE

5. Structure of Female PHALLOSTETHUS DUNCKERI *
(text-fig. 12, A).

Total length 29 mm. The mouth is less oblique than in
Neostethus, and the body is less compressed; the abdominal
profile is not convex, but nearly straight, the rayless fringe (/.)

Text-figure 12.

A, Phallostethus dunckeri, 2. B, Neostethus lankesteri, 2.
Head and abdomen from below (x 8).

@.,anus; p., postanal papille (in Neostethus covering the depression into which
oviduct and ureter open); o., opening of oviduct; w., opening of ureter;
F., dermal fold.

lies in a groove instead of at the edge of the abdomen, and the
genital opening (0.) does not le in a depression; the postanal
papille (p.) are minute. The dorsal fin has more rays (8 to 10)

* T have already given some account of the structure of Phallostethus dunckeri
(Regan, 11); the sections of the male fish are too thick and somewhat overstained,
but with the much better sections of Neostethus lankesteri at hand for comparison
I have been able to make out certain details that were difficult to see without this
help. So far as I can see, the most important error in my former description was
that the priapial ribs, displaced forward and separated from the vertebral column,
were interpreted as elements of the pectoral arch and the antepleural bone, which
embraces their ends, was not recognized as a separate bone distinct from them.

MORPHOLOGY OF CYPRINODONT FISHES, A

than in Neostethus, and the anal fin is much longer, having

26 to 28 rays. Correlated with this is the shorter abdominal

region, the smaller number of preecaudal vertebre (11 or 12), and

the larger number of caudal vertebre (26 or 27) (cf. Pl. I. A).
The visceral anatomy is as in Veostethus.

6. Structure of Male PHALLOSTETHUS DUNCKERI.

Tn addition to the differences from Weostethus described above
for the female, the male Phallostethus has many distinctive
features.

a. HXTERNAL CHARACTERS.

Total Jength 25 mm. The priapium (text-fig. 13) is much
more prominent than in WVeostethus; the grooves between it and

Text-figure 13.

Phallostethus dunckeri, 6. ead and priapium from the proctal and
aproctal sides (X 10).
tx., toxactinium; ef., ctenactinium; pv., pulvinulus; a., anus; w., opening of
ureter; v.d., terminal coil of vas deferens.

the body of the fish are not glandular and are of equal size; they
increase in depth posteriorly and meet behind the priapium to
Proc. Zoou. Soc.—1916, No. II. 2

18 MR. CG. TATE REGAN ON THE

form a median groove, which extends to the anal fin, decreasing
in size backwards (text-fig. 15 B); there is no efferent groove.

The urinary opening (w.) is immediately behind the anus (@.).

The genital opening is ventral and posterior, behind the
articulation of the ctenactinium (ct.).

The ctenactinium is short and nearly straight, with denticu-
lated upper edge; its articulation is ventral, and it lies on the
aproctal side of the priapium.

Anteriorly the attachment of the priapium to the isthmus ends
at the level of the posterior edge of the eye; in front of this is a
free terminal portion that ends in a second movable bony ap-
pendage, toxactinium (tw.); this is rounded in cross-section, tapers
forwards, and curves towards the aproctal side, ending below the
extremity of the lower jaw.

The pulvinulus (pv.) appears to be represented by a rounded
shield, with thick edges, that covers the basal part of the tox-
actinium below and on the aproctal side.

b. SKELETON (PI. I. A).

Third vertebra and first pair of ribs.—The transverse processes
of the third vertebra are normal and symmetrical, but bear no
ribs. The first pair of ribs are nearly symmetrical, but they are
not articulated with the vertebral column ; proximally they end
at the level of the second vertebra, but at some distance from it
on each side; the proximal part of the rib of the proctal side
curves forwards to give attachment to the pleuro-priapial muscle.
These ribs run downwards and forwards, meet below the peri-
cardium, and enter the priapium, where they run downwards and
towards the proctal side in front of the intestine.

Cleithra.— Both cleithra are prolonged forwards and enter the
priapium, ending between the pulvinulus and the basal part of
the toxactinium (PI. III. B, el., ed.’).

Priapial skeleton.—The axial bone (text-fig. 15, a.) is com-
paratively simple; the toxactinium articulates with its anterior
end superiorly and proctally (Pl. III. B, tx.,a.), the ctenactinium
(text-fig. 15, ct.) with its posterior end inferiorly and aproctally.

The antepleural cartilage of Weostethus is represented by an
antepleural bone that embraces the distal ends of the priapial
ribs and runs forwards on the proctal side of the axial bone,
ending a short distance behind the base of the toxactinium.

The pulvinulus has no pulvinular appendage and no bones; it is
aring of parenchymatous cartilage, but seems to have the structure
of true hyaline cartilage in the middie (Pl. IIT. B, pa., pe.).

There are no infrasulear. bones, but the terminal part of the
vas deferens appears to be supported by a bony lamina, as in
Neostethus.

c. MUuscLEs.

The muscles correspond to those of Weostethus, except that
(1) the pleuro-priapial muscle is attached proximally to the

MORPHOLOGY OF CYPRINODONT FISHES. 19

proximal part of the first rib of the proctal side instead of to
the transverse process of the third vertebra; and (2) there isa
toxactinial muscle (text-fig. 15, twm.), that lies outside the longi-
tudinal muscle of the proctal side, ends posteriorly between the
coils of the vas deferens, and anteriorly is inserted on the base of
the toxactinium.

d. ViscerA (text-fig. 14).

A difference from Veostethus 1s that the ureter (w.) accompanies
the intestine (7.) and opens behind the anus; of more importance
are differences in the testis and vas deferens (vd.). The testis
(Pl. IV. A) has no tubules lined with a distinct epithelium, even
near the origin of the vas deferens, which leaves the testis (¢.)
anteriorly and at once becomes coiled up into a mass that lies in
front of and to the right side of the testis ; this “‘ epididymis”’ is
similar to the testis in form and is not much less than it in size

Text-figure 14.

Visceral anatomy of Phallostetius dunckeri, &; the liver, etc., removed (X 10).

c., esophagus; s., stomach ; 7., intestine; &., kidney; w., ureter; ¢., testis;
v.d., vas deferens; a., air-bladder.

(text-fig. 15, B); the vas deferens emerges from its narrowed
anterior end, enters the priapium, and runs backwards on the
aproctal side to above the base of the ctenactinium, then across
to the proctal side before coiling round in a complete circle and
opening ventrally to the exterior in a short seminal papilla behind
the base of the ctenactinium.

The vas deferens is lined with a glandular epithelium ; in the
“epididymis ” this is formed of long columnar cells with basal
nuclei (Pl. IV. B). The outer fibrous layer is quite thin, except
in the terminal coil, where it is thick. The lumen contains
spermatozoa, which seem to adhere together, their heads forming
more or less convex plates, whilst their tails are dependent from
the concave surfaces. However, this structure cannot be very
definitely made out from the sections, but it seems clear that
there are no spermatophores of the type described in Veostethus.

2*

20 MR. C. TATE REGAN ON THE

7. Remarks on the Structure of the Priapium.

Many of the peculiarities of the priapium of Phallostethus, as
compared with that of Veostethus, seem connected with the two
main differences, that the vas deferens opens directly to the
exterior instead of into a glandular groove and that a tox-
actinium is present. To the absence of a glandular groove may
be due the great length of the vas deferens, which coils to form an
“epididymis”; the secretion supplied in Veostethus by the testis,

Text-figure 15.

Phatllostethus dunckeri, 8. ‘Transverse sections (X 18): A, showing the intestine,
vas deferens, and ureter entering the priapium, and also the ureter opening to the
exterior; B, showing the “ epididymis” and the posterior thick-walled portion
of the vas deferens above the articulation of the ctenactinium.

k., kidney; J., liver; s., stomach; 7., intestine; w., ureter; v.d., vas deferens;
a., axial bone; cf., ctenactinium; ppm., pleuro-priapial muscle; tam., tox-
actinial muscle; etm., ctenactinial muscle; pm., muscle of the proctal side;
apm., inner muslce of the aproctal side.

the vas deferens, and the glandular groove comes in Phallostethus
from the vas deferens alone. ‘The efferent groove, infrasulcar
bones, and pulvinular appendage of Veostethus are all directly
connected with the glandular groove, and it is not surprising that,
they are absent in Phallostethus.

MORPHOLOGY OF CYPRINODONT FISHES. 21

The approximate symmetry of the priapial ribs and cleithia in
Phallostethus, as compared with their marked asymmetry in Veo-
stethus, is no doubt due to the symmetrical attachment of the
priapium in the former (text-fig. 15), and its asymmetrical
attachment, owing to the great development of the glandular
groove, in abe bier (text- -fig, 4).

With the presence of a toxactinium as a free appendage may
be correlated the freedom of the anterior part of the priapium in
Phallostethus, whereas in NVeostethus it is attached to the isthmus
right to the end. It seems probable that the toxactinium may
replace functionally the anterior part of the ctenactinium of
Neostethus, and that the shortness of the ctenactinium of Phallo-
stethus may be connected with this.

In both genera the ureter enters the priapium and runs down-
wards and across to the proctal side behind the intestine; in
Phallostethus it opens to the exterior just behind the anus, but
in .Veostethus, owing to the shortness of the free posterior part of
the priapium and the great size of the terminal coil of the vas
deferens, it finds the latter in its way and has to surmount it
before reaching the surface at some distance from the anus.

In Phallostethus, as compared with WVeostethus, the great length
and coiling of the vas deferens, the separation of the first pair of
ribs from the vertebral column, and perhaps the presence of a
toxactinium, may be features of specialization ; in iVeostethus the
development of the glandular groove and the structures associated
with it may be similarly vegarded,

When I first described Phallostethus I suggested that the axial
bone of the priapium might be pelvic and the ctenactinium and
toxactinium fin-rays. This inter pretation seems much less likely
to be correct when the structure of Veostethus is considered also.
The antepleural cartilage of Veostethus has developed in Phallo-
stethus into a long bone with a cartilaginous core; conversely,
the presence in Veostethus of pulvinular and infrasulcar bones,
absent from Phallostethus, seems to show that the skeletal ele-
ments of the priapium develop when and where they are wanted,
and are not to be homologized with any other parts of the
skeleton. Moreover, the pulvinular appendage of WVeostethus
seems to represent a stage of development intermediate between
a simple dermal papilla and movable bony appendages, such as
the ctenactinium and toxactinium, and suggests that these may
have originated as outgrowths whose skeleton changed from con-
nective tissue to cartilage and then to bone as they grew longer
and formed more definite proximal articulations with the axial
bone. Their development in WVeostethus bicornis supports this
view.

In fact, the whole priapium seems to be an entirely new forma-
tion; its appendages, bones, muscles, and glands are not to be
homologized with any structures found in the female fish or in
other Cyprinodonts.

22 MR. C. TATE REGAN ON THE

8, Use of the Priapiwmn.

Jn oviparous Cyprinodonts (Mundulus and Cyprinodon), New-
man (5) has observed the intercourse of the sexes; the male and
female lie side by side and looking in the same direction, and the
male clasps his mate by folding his dorsal and anal fins across.
her, whilst the paired fins also may interlock ; in this way the
eggs and sperm are extruded in such close proximity that fertili-
zation is assured.

It seems likely that in the Phallostethine also the male and
female take up a similar position, the female lying on the
aproctal side of her mate.

In Phallostethus the toxactinium, which curves towards the
aproctal side, may grip her under the chin or even be held in
her mouth, whilst the serrated edge of the ctenactinium may
give it a firm hold on the pectoral region in front of and on the
far side of the genital orifice, in order that the seminal papilla
may be placed against or introduced into the latter.

In Weostethus in seems likely that during intercourse the female
may be held by the ctenactinium across the back of the head, the
anterior descending part lying on her distal side and the terminal
part that at rest curves across under the chin of the male now
curving towards him, perhaps under hers. The spinous process
of the ctenactinium and the pulvinular spine would stick into
her on the side near the male. The posterior end of the priapium
may be held in the depression into which the oviduct opens, whilst
the terminal projecting part of the fold that roofs the efferent
groove and the membranous fringe below it may form a sort of
tube for insertion into the genital aperture of the female.

The asymmetry of the postanal papille of the female may be
due to the side by side position presumably adopted; if so, it.
may be supposed that a male with the right side aproctal would
pair with a female that had the smaller papilla on the left side,
and vice versa.

It is possible that no part of the priapium is actually intro-
duced into the oviduct, and that the spermatophores, first
discharged into the glandular groove and then ejected through
the efferent groove, may adhere to the surface of the postanal
papillee and of the depression into which the oviduct opens, and
that they may be introduced into the oviduct by the action of
the papillee.

The infrasulcar prominence may prevent the flow of the
seminal fluid outwards or forwards at the place where it exerts
the greatest pressure, and the pulvinular appendage may possibly
help to drive the glandular secretion backwards.

In both Phallostethus and Neostethus the probable effect of the
contraction of the longitudinal muscles of the aproctal side and
of the pleuro-priapial muscle would be to move the posterior end
of the priapium aproctally; an additional effect in WVeostethus
would he to close the glandular groove.

MORPHOLOGY OF CYPRINODONT FISHES. 23

It is only by a study of the actual behaviour of these fishes
during the breeding-season that one can hope fully to understand
the purpose for which this elaborate apparatus has been evolved.

9. Comparison of the Priapium with other Copulatory
Organs of Fishes.

In many bony fishes a papilla bearing the genital aperture
forms a simple but efficient intromittent organ; in others the
anal fin is utilized, the vas deferens either opening at its base or
being prolonged on the fin, which may form a copulatory organ
of considerable complexity of structure, as in the viviparous
Cyprinodonts, Peeciliine, Anablepine, etc. (Regan, 10; Gar-
man, 1; Langer, 4).

In its asymmetry and in being either dextral or sinistral
the priapium agrees with the copulatory organ of Anableps
(Garman, 1).

In the Phallostethinz the remoteness of the genital opening
from the anal fin explains why the latter has not been involved,
but does not explain the extraordinary complexity of the pri-
apium, which parallels the mixopterygia of the Selachians in its
specialized skeletal and muscular system (Jungersen, 3).

In the mixopterygia new skeletal elements are developed, and
may either margin a groove or may become movably articulated
with the main axial piece; one may project as an external spine,
in this case formed of calcified cartilage, not of bone. Another
parallel with the priapium is that the mixopterygia attain their
full development rapidly as the individual becomes sexually
mature.

In the Selachians the mixopterygium has a nearly uniform
structure in members of the same family, or even of the same
suborder (Huber, 2; Regan 7), and the differences between the
priapia of Phallostethus and Neostethus are as great as between
the mixopterygia of the subclasses Holocephali and Euselachii.

10. Rank and Position of the Phallostethine.

There can be little doubt but that Veostethus and Phallostethus
belong to the family Cyprinodontide, and they seem to agree in
every way with the most generalized subfamily, the Funduline,
except for three features of specialization, namely, the anterior
position of the anus, the absence of pelvic fins, and the develop-
ment of a priapium in the male. This view as to their relation-
ship is best expressed by placing them in a distinct subfamily,
Phallostethine, of the Cyprinodontide, a family that already
includes both oviparous and viviparous fishes, the latter with
intromittent organs of three different types (Regan, 8).

Other examples of animals which retain the general structure
of the group to which they belong, whilst one particular organ or
- system is profoundly modified or some new feature of importance

24. MR. C. TATE REGAN ON THE

is developed, can readily be found, although few cases are so
striking as the Phallostethine.

Among fishes, one may recall the curious Cyprinid Gyrinochilus
(Regan, 9, p. 29), which has the form, fins, scales, etc., of Crosso-
chilus and Discognathus, to which it is certainly closely related,
yet it has the mouth, gills, and pharyngeals so modified in con-
nection with its peculiar methods of breathing and feeding that
some ichthyologists have regarded it as the type of a separate
family

Because WVeostethus and Phallostethus so obviously belong to the
large and varied family Cyprinodontide, one attaches but little
classificatory importance to the development of the priapium and
its evolution along two very distinct lines. But if these were the
only known Cyprinodonts they would certainly form a separate
order, and the differences in structure of the priapia would be
regarded not merely as generic, but as subordinal, and the much
longer anal fin, the abdominal groove, ete., of Phallostethus would
be held to support the view derived from the structure of the
priapia that it and Veostethus had diverged widely and through
a long period of time from their common ancestor.

Were these the only living Teleosts many zoologists would
regard them as a separate class, comprising two well-marked
orders, just as some have suggested that the Dipnoans should be
removed from the Pisces, mainly on account of the isolated
position of their living representatives, Ceratodus and the Lepido-
sirenide, and have given these ordinal rank with the names
Monopneumones and Dipneumones.

These somewhat fanciful considerations are put forward merely
to suggest that the rank given toa group depends on several
factors, and that the degree of differentiation is one of the least
of these.

11. Note on the Origin and Homologies of Skeletal Hlements.

The priapium appears to be an entirely new organ, and it has
a highly developed skeleton, comprising a number of new ele-
ments that cannot be homologized with any parts of the skeleton
of other fishes ; this suggests that the intermuscular connective-
tissue may give rise to cartilaginous or bony elements whenever
and qlhemeren the necessity may arise. This is, of course, not
new, but it is a point of view not always kept in mind by mor-
phologists, as could be illustrated by numerous examples, one of
which may be adduced.

In certain Selachians, and especially in the Hypotremata, there
is a median series of vertical cartilaginous plates above the verte-
bral column, and the question has been raised whether these
belong to the vertebral column or to the fin-skeleton. Thus
Goodrich (Lankester’s ‘Treatise of Zoology,’ pt. ix. figs. 50, 52)
has figured them in Squalus and Squatina, and has “described
them as either modified radials or neural spines. I have long
thought it probable that they were neither, but autogenous

MORPHOLOGY OF CYPRINODONT FISHES. 25

structures developed in the intermuscular septum, and I feel
this opinion strengthened as the result of my work on the
Phallostethine.

BIBLIOGRAPHY.

1. Garman, S.—‘ The Cyprinodonts.” Mem. Mus. Comp. Zool.
xix. 1895, 179 pp., 12 pls.

2. Huser, O.—‘ Die Kopulationsglieder der Selachier.” Zeitschr.
f. wiss. Zool. lxx. 1901.

3. JuNGERSEN, H. F. E.—‘“‘ On the Appendices Genitales in the
Greenland Shark, Sommiosus microcephalus, and other
Selachians.” Danish Ingolf.-Exped. u. pt. 2, 1899,
SS) pps Or pls.

4, Lancer, W. F.—“ Beitriige zur Morphologie der viviparen
Cyprinodontiden.” Morph. Jahrb. xlvii. 1913, pp. 193-
307.

5. Newman, H. H.—‘‘Spawning Behaviour and Sexual Di-
morphism in Pundulus heteroclitus and Allied Fish.” Biol.
Bull. xii. 1907, pp. 314-348, pls. xxvii.—xxviil.

6. Puitrer1, EH.—‘‘Spermatophoren bei Fischen.” Verh.
Deutsch. zool. Ges. xvii. 1907, pp. 105-108.

7. Regan, C. T.—“ A Classification of the Selachian Fishes.”
P.Z.S. 1906, i. pp. 722-758.

8. Reaan, C. T.—‘The Osteology and Classification of the
Teleostean Fishes of the Order Microcyprini.” Ann.
Mag. N. H. (8) vii. 1911, pp. 320-327, pl. viii.

9. Reean, C. T.—‘‘ The Classification of the Teleostean Fishes
of the Order Ostariophysi.—I. Cyprinoidea.” Ann. Mag.
N. H. (8) viii. 1911, pp. 13-32, pl. 1.

10. Recay, C. T.—“‘ A Revision of the Cyprinodont Fishes of the
Subtaraily Pecline..’ PZ) 1903) pp. If —101s;
pls. xcix.-ci.

11. Reean, C. T.—‘‘ Phallostethus dunckeri, a remarkable new
Cyprinodont Fish from Johore.” Ann. Mag. N. H. (8)
x. 1913, pp. 548-555.

EXPLANATION OF THE PLATES.
Puate I.

Skeletons of Phallostethus dunckeri (A) and Neostethus lankesteri (B),
reconstructed ; A 6 and B 5 times the actual size.

Puate II.
Neostethus lankesteri.

A. Transverse section through anterior part of seminal papilla and adjacent portion
of glandular groove (X 130). e., glandular epidermis lining folds and
pockets on surface of papilla; p., papillary bone; sp., mass of mucus con-
taining spermatophores, lying in the vas deferens, where this opens into the
glandular groove (g.) ;.a., process of axial bone.

B. Transverse section through glandular groove and pulvinulus at base of pul-
vinular appendage (X 130). g., glandular groove; 7., infrapulvinular groove ;
is., anterior infrasulcar bone; pvd., cartilaginous terminal process of inner
pulvinular bone, for articulation of pulvinular appendage (pva.).

26 ON THE MORPHOLOGY OF CYPRINODONT FISHES.

Prats III.

A. Neostethus lankesteri. Transverse section through isthmus and anterior part
of priapium (X 130) ; the right side is aproctal. w., urohyal; cl., cleithrum
of aproctal side; a., axial bone; pé., outer pulvinular bone; g., glandular
epidermis ; m., outer muscle of aproctal side.

B. Phallostethus dunckeri. ‘Transverse section through isthmus and anterior part
of priapium, showing the toxactinium articulating with the axial bone
(X 130); the left side is aproctal. w., urohyal; a., axial bone; ¢a., tox-
actinium ; e/., cl.’, cleithra; pe., pulvinular cartilage; pa., parenchymatous
cartilage. 5

Puate IV.

A. Phallostethus dunckeri. Transverse section through part of testis near origin
of vas deferens (X 200).

B. Phallostethus dunckeri. Transverse section through “ epididymis” (X 170).

C. Neostethus lankesteri. Transverse section through upper part of testis (X 170).

IVA AS. AUS. UWOLOS iS), Jel IL.

102° 30 E.

Jl
Reg. No. 231-15. Drawn at the C.D.O. K.L. F.M.S.
Scale, 21 Miles to an Inch : i
Miles 10 5 0 10 20 30 40 Miles
poe eet st ———|

SKETCH MAP OF S.E.SIAM

ON MAMMALS FROM SIAM. 27

2. On a Collection of Mammals from the Coast and Islands
of South-East Siam. By ©. Bovey Kuoss, F.Z.5.,
F.R.G.S. With an Account of the Fruit-Bats, by
Dr. Knup AnpirseEn, F.Z.8.

[ Received August 30, 1915: Read November 9, 1915.]

(Plate I.* and Text-figures 1, 2.)

INDEX.
GEOGRAPHICAL : Page
Description of districts visited ...............-:.s::00-. 27
SYSTEMATIC:
Presbytis germaini mandibularis, subsp. n. ......... 32
Paradoxurus minor kutensis, subsp. n. ............... 84
Martes flavigula indochinensis, subsp. n. ............ 35
Tupaia concolor sinus, SUbSp.D. ...........02..ee eee. 86
Ratufa melanopepla leucogenys, subsp.» TV etciqaecee rane)
RK. m. sinus, subsp. n. ........ be eee eee
Sciurus ferrugineus fr andseni, subsp. 1 1B cananpeoeas: | aio)
S. albivewilli, sp. n. : BREE RE STEREO eee,
Menetes berdmorei umbr osus, subsp. MF esse et. ce Let RAD
M. b. rufescens, subsp. Ts Wesson PAGDRE Nace ans eases £ LO)
Hpimys jerdoni marinus, subsp. Ba hel endeswra sis Atscecis cs O)
JOG SOMFOFEP THOUS, SUDO Wo pooscuaedssosddcascce oodossono sen! Ol
JBL, 8 OPGHOG ESOS: SOUNDS Tc. s-o505 de cspocncavonaceecanoep oo D4
IIL 3 TEOEDSOS, SUD}, Te, Soeceoces sanc5oanoea -os00cenaGd0ue Oe
196 Gs OANAG UG, SUONGD Me aconsocunoreavacnauvesecs8ecodconsee | OE
[Bis 8s GOW ZQIAISy SOI Os We GoosrncneMieduisocassedcsodondonces OS)
JI, 83 GROSS, SUOID5 Wo | Gapsgsussdcasendadcconegcoscenscnsna | 2
II), Ss GAMADPOSIS, SUN 05 We bos beeche soca eeorascoasnonnnaes OE
Hi. rattus rangensis, subsp. n. ............ 56
H. 7. klumensis, subsp. n. a 56
IIL, 7s TOU EPOSOSS SOO We oon sovesdccosoasensononndosnacneacss Bw
IDG FP LERCIAISAS, OUOGV Wo 253 sondossoscsbosoconossesocpaegsien BTL
EE. berdmorei magnus, subsp. n. 57

Towards the end of 1914 I went on short leave to Siam with
three Dyak assistants, my object being zoological collecting
in the extreme south-east of the country, and having reached
Chantabun by steamer via Bangkok, I hired a small native
sailing-vessel (“rua pet”) and passed six weeks cruising and
camping on the coast and islands beyond (see Pl. I.), setting in
altogether about thirty-three working days. The result was
a set of rather over 500 mammals, 300 birds, and 250 reptiles
and batrachians.

Mammals were the principal object of the excursion, and |
chose this district of the mainland because very little investigation
of it had taken place, while the islands were quite untouched ;
for although Captain Stanley Flower appears to have been in
Chantabun in 1898 (P. Z. 8. 1900), no other naturalist has
followed Mouhot, whose collections were reported on by Gray
and Gunther in the ‘ Proceedings’ of 1859 and 1861.

In recent years Mr. T.. H. Lyle has sent home specimens

* Kor explanation of the Plate see p. 66.

28 MR. C. BODEN KLOSS ON

from the better-known parts of Siam—the basin of the Menam
and its head-waters near Chiengmai and Nan (¢/. Bonhote,
P. Z. 8. 1900, 1901, 1902); and the ornithologist Count Nils
Gyldenstolpe, of the Royal Swedish Museum, has lately collected
mammals in much the same area and also in the province of
Korat (cf. Gyldenstolpe, Arkiv fér Zoologi, Stockholm, 1914);
but still the remote south-east has remained unvisited, nor can
I find that its French possessors have carried out any investiga-
tions in the adjacent parts of Cambodia. So to the zoologist
the region between Cochin-China and the better-known districts
of Siam was largely a no-man’s-land.

The mainland visited requires no description ; it is undulating
coastal country, covered with forest except round villages, and
with mountains in the distance. Lem Ngop * les on the north
shore of Koh Chang f Strait, and Ok Yam (ov Jam) is eastward
of Koh Kut, just within French territory, as the present
boundary of Cambodia comes out on the coast half a mile to the
west of it (lat. 11° 40’ N.). Klong Yait and Klong Menao are
estuaries to the north of Ok Yam.

Of the islands, Koh Chang is about 15 miles long and 7 wide ;
it is very hilly, and its highest peak, one of a number, rises to
2446 ft. The strait which separates it from the mainland.
narrows in part to 3 miles, though, since much of the adjoining
province is an alluvial plain, the island was probably at one
time more isolated. It is the northernmost of the Chantabun
Archipelago.

The two little islands of Mehsi (935 ft. high) lie close together,
two miles off the south-east end of Koh Chang; they are called
on the chart (Admiralty 2721) “Te du Pic” and ‘‘ Le Chameau ”
respectively. A little farther from the southern shore le Koh
Klun (600 ft.) and Koh Kra (800 ft.).

Koh Kut, 15 miles 8.8.E. of Koh Chang and about 18 miles
from the Siamese-Cambodian coast, is the southernmost of the
group; it is some 13 miles long and 5 wide, elevated, with a
peak of 1171 ft. It is uninhabited, but swarms with ticks,
which infest every animal upon it and made our week’s residence
there a most uncomfortable experience.

Between the northern islands and Koh Kut, from east to
west, are Koh Mak and Koh Rang (800 ft.), the first the larger,
being about 4 miles long, but, except for one small hill, very low.
Three miles or so west of it is Koh Rang (Koh Loi of the chart),
the most seaward of the group, and, like the other smaller
islands, a little under two miles long. There are, further, a few
little islets which I did not visit.

Koh Chang and the Mehsi Islands are separated from the
mainland by depths of between 3 and 4 fathoms; Kra, Klum,
and Mak rise from 6 to 7 fathoms of water; Koh Kut stands in
9 to 10 fathoms, and Koh Rang is on the 12-fathom contour-

* Lem=Cape. + Koh=Island. t Klong= River.

MAMMALS FROM SIAM. 29

line. All are covered with dense evergreen tropical forest, and
Koh Chang and Koh Mak alone are inhabited.

The only terrestrial mammals on the smaller islands are forms.
of Epimys surifer and H. rattus. Koh Mehsi, though nearer the
mainland, higher, and in shallower water, has a form of the
latter only.

Koh Chang and Koh Kut vary in the composition of their
faunas, for while the former lacks a Ratufa, a species which
occurs on Koh Kut, the latter is without any Presbytis, Tupaia,
or rattus vat, all of which are found on the larger island. On
both a form of Hpimys jerdoni is very common, though it was
not met with on the mainland.

When we got back to Bangkok visits were paid.to Koh Si
and Koh Phai in the Inner Gulf, and accounts of the collections
made on them and of the reptiles and batrachians obtained in
the south-east appear in the ‘ Journal’ of the Natural History
Society of Siam, while a report on the birds is contributed to
‘The Ibis.’

The first set of all collections has been given to the British
Museum (Nat. Hist.) at South Kensington, and the second to
the U.S. National Museum at Washington.

Ridgway’s colour-names used here are those of his second
publication, ‘ Colour Standards and Nomenclature,’ 1912

Though in the title of this paper I have mentioned only one
locality for the sake of brevity, it may be said that most of the
specimens obtained at Ok Yam and many from Klong Yai
actually came from the Cambodian side of the boundary, as it
now runs between Siam and that country.

1. HyLopatss PILEATUS Gray.

Hylobates pileatus Gray, P. Z. 8. 1861, p. 135, pl. xxi.; de Pou-
sargues, Mission Pavie, Indo-Chine, Etudes Diverses, iii. p. 511
et seq. (1904).

Hylobates agilis, variety pileatus Flower, P. Z. S. 1900, p. 313.

5 adult males, 2 adult females, 1 female juv., from Klong
Menao and Len Ngop, 8.E. Siam.

Tt is probable that these animals are practically topotypes, as
Mouhot’s specimens came from some part of Cambodia. The
statement that they were met with on a small island near the
coast must be received with hesitation until their occurrence in
an insular locality is confirmed.

The original description of the species applies excellently to
the present series of this individually variable genus. The five
males are (though paler on the dorsal region and the lower half
of the legs) black or brownish black throughout except for a ring
round the face, the hands and feet, and a genital tuft, which are
dull white or buffy-white. The pale band running from the
temples round the back of the head, greyish, ashy or brownish
posteriorly, is not always complete behind, and then the dark

30 MR. C. LODEN KLOSS ON

patch on the crown (which gives its name to the species) is not
always isolated. The pale area on the hands and feet varies in
extent ; sometimes it covers the extremities from the wrists and
ankles and sometimes is confined to the fingers and toes.

The two adult females are drab above, the rump and outer
side of limbs somewhat warmer in tone, being slightly tinged
with ochraceous. ‘fhe crown, cheeks, throat, and a diamond- or
shield-shaped area over the breast and abdomen extending to
the axille and almost to the genital region are blackish. The
hands and feet, inner side of limbs, and the pelage bordering
the black areas are somewhat lighter than the rest of the body.

The young female is drab to buffy throughout except for a
slightly indicated dark cap on the top of the head.

Gray was only able to state that the pale examples examined
by him were ‘“ probably female.” I was told by Siamese villagers
who saw my specimens that females were always pale on the
back and limbs, while the abdominal surface is variable (cf. de
Pousargues, op. cit. p. 516).

I am of opinion that this animal should be given only sub-
specific rank and treated as a geographical race of H. lar, which
occurs as far east as Luang Prabang (de Pousargues, op. cit.
p. 511) and in Central Siam, east of the Menam (Gyldenstolpe,
Arkiv for Zoologi, Band 8, no. 23, p. 6, 1914).

Though I only obtained these animals at two collecting-stations,
we heard them calling every morning from the hills all along the
coast from Ok Yam on the Franco-Siamese boundary to the
mouth of the Ban Yao River near Chantabun. Their ery did
not appear to differ in any way from that of H. lar or H. agilis.

Measurements.—See table, p. 67.

9. Macaca ANDAMANENSIS Bartlett.

Macaca andamanensis Bartlett, Land and Water, vol. viii.
p- 57 (1869); Sclater, P. Z. 8. 1869, p. 467 & figure.

Macaca leoninus Sclater, P. Z.S. 1870, p. 663, pl. xxxv.;
id., op. cit. 1898, p. 280; de Pousargues, Mission Pavie, Indo-
Chine, Etudes Diverses, iii. p. 517 (1904).

A single adult female of this species was obtained at Klong
Menao. This sex does not appear to have been fully described,
so that an account of the present specimen may not be out of
place.

Colour.—Crown, nape, and entire upper surface, outer side of
forearms, hands, and feet mummy-brown to olive-brown, slightly
lighter and yellower across the shoulders and lighter on the
upper part of forearms and on the sides, the hairs annulated with
dull pale buff. Thighs and buttocks paler and greyer (near
drab-grey), devoid of annulations, the hips somewhat more buffy.
Face, temples, sides of head and neck, and buttocks buffy-silvery
or dirty brownish white, underside of body and inner side of
arms more silvery. ‘Tail mummy-brown below, blackish brown

MAMMALS FROM STAM. 31

above, this colour not continued on to the rump, which is
scarcely perceptibly darkened mesially. Inner side of ears
silvery, a few dark hairs round the eyes. Skin of orbital region
in living animal pale lilac-blue; nose, lips, palms, and soles
fleshy brown.

Comparison with Pacneles of M. nemestrina from the Malay
Peninsula shows that the female andamanensis differs in the
absence of any rufous or ochraceous tone in the pelage and in
its undarkened rump: while, as regards the skull, the muzzle is
much reduced in size and tapers both upwards and forwards and
the interpterygoid space is wider, embracing laterally a greater
portion of the bulle.

Considering the individual variability of macaques, the
dimensions of the skull are in strikingly close agreement with
those of the female recorded by Anderson in ‘ Zoological Re-
searches,’ p. 52, which probably came from the country west of
the Irawadi River.

Measurements.—External measurements, taken in the flesh:
total length, 640 mm. ; tail from angle formed with rump above,
200; head to vent, 480; hind foot, 158; ear, 38. Skull: greatest
length, 117°5(113°8); basal length, 77°7 (79-2); palate to foramen
magnum, 32 (34:5); anterior edge of auditory opening to
gnathion, 86:2 (81:2); occiput to narion, 85 (87°6); narion to
gnathion, 49-3 (51:7); orbit to gnathion, 39; breadth of muzzle
at pm' 30°5 (30-4), at m® 35°6 (34:3), at roots of zygomata 40;
facial breadth at fronto-malar suture, 61 (62); post-orbital
breadth, 47-6 (48:2); zygomatic breadth, 77 (76); mastoid breadth,
64 (62:7); orbit, 25x 25°5 (24x 28); maxillary tooth-row ex-
clusive of incisors, 88; length of mandible in alveolar plane,
77 (75).

Elliot has pointed out (‘ Review of the Primates,’ vol.

p. 209) that Blyth’s name for this monkey is preoccupied ie
leonina Shaw, applied to J. albibarbatus, so that andamanensis
Bartlett, based on an example introduced into the Andamans
from Burma, must unfortunately be used.

3. Macaca irus Cuvier.

Macacus irus F. Cuv. Mém. Mus. Hist. Nat. Paris, iv. 1818,

120.
r; Macacus cynomolgus Flower, P. Z. 8. 1900, p. 316; de Pou-
sargues, Mission Pavie, Tindle-Olhine. Etudes Diversions iol Oe) DIG
(1904).

Pithecus fascicularis Gyldenstolpe, Arkiv for Zoologi, Band 8,
No. 23, p. 3 (1914).

3 adult males, 2 adult females, 1 immature female, from Koh
Kut Id., S.E. Siam.

3 immature females, from Koh Chang Id., S.E. Siam.

I have compared these animals with a large number from
the Malay Peninsula and adjacent islands which show a great

32 MR. C. BODEN KLOSS ON

amount of individual variation from each locality. The only
difference I can detect is that, series for series, the Siamese
animals have perhaps a somewhat greater quantity of black hairs
on the forehead and top of head, but the distinction is un-
important, and I have therefore placed them under the name
first definitely applied to the crab-eating monkey of the S.E.
Asiatic mainland.

It may be said that the blackish hands and feet which Elliot
(‘ Review of the Primates,’ vol. 11. pp. 189, 230, 231, 233 (1913))
emphasizes as the distinguishing characters of J. irws do not,
so far as my experience goes, exist.

On the whole the series is very uniform, save that adult females
have no trace of rufous on the dorsal surface.

We met with a large herd of these monkeys on the mainland
on one oceasion while sailing up a river at daybreak, but were
not prepared for obtaining them.

x

Measurements.—See table, p. 67.

4. PRESBYTIS GERMAINI M.-E.

Semmnopiihects germaint Milne-Edwards, Bull. Soc. Philom.
1876, Feb. 12; Flower, P. Z. 8. 1900, p. 319; de Pousargues,
Mission Pavie, Indo-Chine, Etudes Diverses, TAO. aL (1904)
(mis-spelt ger mani).

Presbytis germaini Gyldenstolpe, Arkiv for Zoologi, Stockholm,
Band 8, No. 23, p. 5 (1914).

One adult male of this handsome silvery-black leaf-monkey was
obtained at Klong Yai, 8.H. Siam, where, and in Cambodia and
Cochin-China, it is the representative of the cristata group.

Measurements.—See table, p. 67; and also under P. g. man-
dibularis.

5. PRESBYTIS GERMAINI MANDIBULARIS, subsp. n.

Type. Adult male (skin and skull), No. 1483/C.B.K. B.M.
No. 15.11.4.5. Collected on Koh Chang Id., $.E. Siam, 7th
December, 1914.

Characters.—A form of P. germaini characterised by smaller
size, paler legs, more slender zygomata, broader interpterygoid
space and basioccipital, and by the higher and narrower ascending
ramus of the mandible.

Colowr.—Resembles P. germaini, except that whereas in the
mainland animal the buttocks, outer and posterior sides of legs
are pure silvery white, sharply contrasting with the black feet
and back, in the island form these areas are black or greyish,
only frosted with silver, and thus less contrasting with the feet
and back.

Skull and Teeth.—As compared with an adult but less aged
male from the mainland, the skull is smaller ; the zygomatic arches
much less deep (4: 7 mm.), the lower edge being concave instead
of straight; the basioccipital is relatively broader; the inter-
pterygoid space broader and more oblique, the sides more spread

MAMMALS FROM SIAM. 33

out, and the tips of the pterygoids farther apart (36 : 33 mm.).
The ascending ramus of the mandible is steeper, being both
higher and narrower, with the sigmoid notch less broad; a line
passing down the back of the condyle and angle of the mandible
is almost perpendicular to the plane of the base of the mandible
in P. germaini, while in its subspecies, at their junction, these
planes form an acute angle only.

Measurements *.—External neu Ree, taken in the flesh:
total length, 1260 (1300) mm.; tail from angle formed with
rump aioe. 720 (755); head ‘to vent, 540 (570); hind foot,
160 (160); ear, 40 (42). Skull: greatest length, 99-5 (106) ;
basal Eee 72:5 (80); zygomatic breadth, 77 (78); maxillary
tooth-row, 35 (36); greatest length of mneraiblle. 73 (81): per-
pendicular height of coronoid process, 47 (44): perpendicular
height of condyle, 44 (42); breadth of ascending ramus from
anterior root opposite centre of m, to angle, 32 (34); coronoid
process to back of condyle mesially, ae (18: B).

Specimens examined.—Seven (the type, 3 adult females, 2 im-
mature females, 1 male juy.)T

Remarks.—Though, as listed above, I have only one adult
male from the mainland for comparative purposes, I have re-
garded it as a typical example, and while the above details record
differences between it and the insular male, the characters of the
latter are completely confirmed by the remainder of the series
from Koh Chang.

The colour of the infant male is ochraceous orange above,
ochraceous buff below, with traces of darkening on forehead,
temples, and tip of tail.

The two immature females, both partially retaining their
milk-dentition, resemble adults in every respect in colour of
pelage. No form of Presbytis occurs on the neighbouring large
island, Koh Kut.

6. PARADOXURUS HERMAPHRODITUS Pall.

Viverra hermaphrodita Pallas, Schreber, Saugeth. ii. p. 426
(1778).

Paradozurus pallasii Gray, P. Z.8. 1861, p. 136.

Paradoxurus hermaphroditus Flower, P. Z.8. 1900, p. 329;
de Pousargues, Mission Pavie, Tndo-@hine! Etudes Diverses, iii.
p. 522 (1904).

A half-grown female was obtained on Koh Chang Id. which
differs from P. m. kutensis in its lighter, more bufty colour,
narrower stripes, and paler shoulders; the muzzleand extremities
are less black, and the chest is darker; a broad pale band extends
uninterruptedly across the forehead, much reducing the brownish
area on the top of the head. Save for this latter, it appears to
bear some likeness to P. cochinensis of Cochin-China (Schwarz,

% Measurements in parentheses those of an adult male from the adjacent main-

land, No. 1839/C.B.k.
+ See table, p. 67.

Proc, Zoon, Soc,—1916, No, III, 5)

34 MR. C. BODEN KLOSS ON

Ann. & Mag. Nat. Hist. ser. 8, vol. vii. p. 635, 1911), but it also
exactly resembles externally a specimen from the State of Perlis,
in the Malay Peninsuia, and others from islands north of
Penang. The skull is too young for comparison.

PARADOXURUS MINOR KUTENSIS, subsp. 1

Type. Adult male (skin and skull), No. 1749/C.B.K. B.M.
No. 15.11.4.39. Collected on Koh Kut Id., 8.E. Siam, 27th
December, 1914.

Characters.—A race of P. minor Bonhote (‘ Fasciculi Malay-
enses,’ Zoology, Part 1, p. 9, 1903), characterised by blacker
(less brownish) muzzle, crown, tail, and extremities, less fulvous
fur above, paler nape and chest, the latter concolorous with the
rest of the buffy abdomen, by the great breadth across the
zygomata, and by smaller bulle.

Colour.— Muzzle, chin and throat, top of head and ears, distal
half of fore and hind limbs extending higher on under surface,
and greater part of tail brownish black. Five black stripes or
rows of spots on the back and a few on the flanks, the inner
three extending on to the base of the tail and becoming somewhat
broken and ecoulan on the shoulders, which, with the neck, are
somewhat clouded with blackish. A few black spots on sides
and thighs. General ground-colour of the body, extending to the
throat below and over the neck above, pale smoke-grey, slightly
tinged with buff on the rump and thighs. A broad whitish
band from above the eyes passes below the ears to the sides of
the neck, but is interrupted on the forehead, where the black
area of the muzzle is connected with that of the crown. Base of
tail pale fulvous clouded with longer black-tipped hairs.

Skull and Teeth.—Yhe skull agrees with those of P. minor,
from Peninsular Siam, but is rather larger, the zygomatic
breadth in particular being greater. The bullz, on the other
hand, are considerably saellen The teeth also are in general
agreement, but there is a greater deflection in the maxillary row
owing to the fact that the posterior cusp of m‘ is placed much
nearer the centre of that tooth.

Measurements *. 8, taken in the flesh :
total length, 480 (460) mm. tal, 420 (450); hind foot, 74 (64) ;
ear, 40 (39). Skull : ereatest length, 101 (96); basal length, S10 3
leneth of palate, 45 (43) : breadth of palate between carnassials,
16 (4); ereatest breadth of brain-case, 30°D (32); zygomatic
breadth, 58°5 (53); intertemporal constriction, 13; length of

maxillary tooth-row, exclusive of incisors, 37.

Specimens examined.—The type and a half-grown female with
exactly similar coloration.

Remarks.—I\ts rather larger size, paler, less fulvous ground-
colour, smaller bulle, and broader skull serve to distinguish this -
animal from P. minor of the type-region ; in addition, the tooth-

* Measurements in parentheses those of the type of P. minor, an adult female
from Jalor, Peninsular Sian.

MAMMALS FROM SIAM. 35

row is a little more deflected. This latter character, apart from
smaller size, may serve to separate the minor from the herma-
phroditus section, in which the maxillary tooth-row is less bent,
since the outer lobes of m' are more in line. In the Koh Kut
animal the great breadth of the skull near the posterior roots of
the zygomata causes it to have a very heart- or pear-shaped
outline.

The only examples of P. minor hitherto recorded from Indo-
China are two young individuals collected by Dr. Vassal in
Annam (Bonhote, P. Z. 8. 1907, vol. i. p. 6).

8. MARTES FLAVIGULA INDOCHINENSIS, subsp. n.

Martes flavigula de Pousargues, Mission Pavie, Indo-Chine,
Etudes Diverses, iii. p. 521 (1904) 2

Type. Adult female (skin and skull), No. 1860/C.B.K. B.M.
15.11.4.40. Collected’ at Klong Menao, S.E. Siam, on %th
January, 1915.

Characters.—Closely resembling in’ colour Martes flavigula
flavigula, but considerably smaller, with naked soles, harsh fur,
and broad-lobed posterior molar, agreeing in these respects with
the equatorial members of the group.

Colowr.—Entire upper surface of head and neck to shoulders,
including ears, hind feet, and tail, deep brownish black; distal
half of fore legs, rump, base of tail, and hind limbs blackish
brown, these colours gradually changing into honey-yellow be-
tween the shoulders and hind quarzters, the median dorsal line
being washed throughout with mummy-brown, most strongly
posteriorly. Sides of neck between ears and shoulders bright
buft-yellow; sides of upper lip, chin, and throat white; under-
side of neck and fore-chest dirty white tinged with yellow; chest
and abdomen cartridge-buff. Inner sides of ears mummy-brown
edged with whitish and with an indistinct ceutral patch of the
same colour.

Skull and Teeth.—Do not appear to differ from those of JZ. f.
peninsularis Bonhote (8 specimens examined), except that the
skull is a trifle larger and the bulle a little larger and more
dilated.

Measurements *.-—Kxternal measurements, taken in the flesh :
head and body, 480 (430) mm.; tail, 385 (383) ; hind foot, 96 (85) ;
ear, 38 (32). Skull: greatest length, 91 (90); basal length,
85:5 (83°5); length of palate from henselion, 42 (41); least
palatal breadth between carnassials, 13°5 (15); breadth at post-
orbital constriction, 24°5 (21-5); zygomatic breadth, 51 (49°5).

Specimens exanuned.—One, the type.

Remarks.—In colour this animal seems closely to resemble
M. f. flavigula, ranging from Nepal to Burma, but its naked
soles, short harsh fur, small size, and broad inner lobe of the
posterior molar place it in the equatorial section of the group

* Measurements in parentheses those of an adult female IL. f. peninsularis from

Trang, Peninsular Siam: F. M.S, Mus. No. 1142/10.
)
4 wo

36 MR. C. BODEN KLOSS ON

and separate it from the true J/. flavigula according to Mr. J. L.
Bonhote in Ann. & Mag. Nat. Hist. ser. 7, vol. xu. p. 342 et seq.
(1901), who there reviewed the group.

Since writing the above paper Mr. Bonhote has recorded two
examples of the true J. f. flavigula from Chiengmai, Northern
Siam (P. Z. 8. 1902, part i. p. 38), thus increasing the range of
this form. These examples are much larger than the individual
under discussion here, so that it would appear that there are in
Siam two races similar in colour bus differing in size and in the
characters given above. Were it not for these latter it would be
most convenient to regard the south-eastern animal simply as a
small race of I. flavigula.

TuUPATA CoNcCOLOR Bonhote.

Trupaia concolor Bonhote, Abstract Pe Ze Ss. 190 Soares
12 oe Wo KONG job S$ Iivom, (Peoe, WSs INA. Wine. vol, adh [Do De)
(OILS),

Tupara belangeri de Pousargues, Mission Pavie, Indo-Chine,
Etudes Diverses, iii. p. 520 (1904); Gyldenstolpe (partim), Arkiv
for Zoologi, Band 8, No. 23, p. 9 (1914).

2 males, 2 females, Ok Yam, Franco-Siamese Boundary ;
3 males, 5 females, Klong Yai; 2 males, 2 females, Klong Menao,
S.E. Siam.

These animals are intermediate between 7’. helangeri Schreb.,
of Pegu and Tenasserim, and 7’. concolor Bonhote, known from
Southern Annam and Cochin-China. On the whole, however,
they most closely approximate to the latter, though the pale
shoulder-stripe is perhaps a little more marked, and so I have
assigned them to it on geographic grounds. Several of them
which are faintly washed on the rump with ochraceous, nearly
resemble examples of 7. belangert from Southern Tenasserim, but
the annulations anteriorly are somewhat coarser, while the buff is
of a slightly deeper shade. There are three pairs of mamme as
in 7. belangeri; in 1’. concolor the number is unfortunately un-
known, as are the external measurements. The present animals
ave somewhat larger than 7. belangeri, while the dimensions of
the skulls ave similar to those of skulls from Annam and Cochin-
China.

Measurements.—Kars of the series, 15-20 mm. For other
measurements see table, p. 68. ;

10. TuPAIA CONCOLOR sryus, subsp. n.

Type. Adult male (skin and skull), No. 1422/C.B.K. B.M.
No. 15.11.4.31. Collected on Koh Chang Id., 8.E. Siam, 7th
December, 1914.

Characters. —Like T. concelor, from the adjacent mainland, but
smaller, darker, with yellower under surface and a conspicuous
shoulder- -stripe.

Colowr.—KEntive upper surface a grizzle of ochraceous buff and

MAMMALS FROM SIAM. Si

blackish, the crown more ochraceous, the tail blacker, and the
sides and limbs more buffy, but no difference in tone between
the shoulders and rump ; shoulder-stripe conspicuous cream-buff.
Chin, throat, chest, and median abdomen buff-yellow ; underside
of thighs btffy- grey, of fore limbs buffy. Hairs of tail below
with two distinct buffy annulations and a narrow subterminal
one slightly deeper in shade. Ears dark.

Skull and Teeth.—Do uot differ from 7. concolor except in
size.

Measurements.—Kars of the series, 15-18 mm. For other
measurements see table, p. 68.

Specimens examined.—Seven, 4 males and 3 females, from the
type-locality.

Remarks.—Only one island race of 7 paia,the present form, was
met with during the excursion, the family being unrepresented
on the large island of Koh Kut. 7c. sinus is a well-marked
insular race, clearly differentiated by smaller size, concolorous
dorsal area, and conspicuous shoulder-stripe.

11. DENDROGALE FRENATA Gray.

Tupaia frenata Gray, Ann. & Mag. Nat. Hist. ser. 3, vol. vi.
p. 217 (1860).

Dendr ogale frenata de Pousargues, Mission Pavie, Indo-Chine,
Etudes Diverses, ili. p. 520 (1904) : 2 ‘eoralmone., IAS. ISOM.
p. 8; Lyon, Proce. U.S. Nat. Mus. vol. xlv. p. 128 (1913).

1 male, 1 female, Klong Menao; 1 female, Klong Yai, 8.E.
Siam.

Recently Dr. M. W. Lyon has so carefully characterised this
species in his monograph on the Tupaiidee (loc. cit. supra), that
there is little more to add. Hs description, however, apparently
applies to the female, as the two examples of that sex now
obtained most closely agree with it, while the male is both larger
and of richer colouring, with darker tail, the upper surface of
the body being more tawny, the under surface of a richer buff,
while the bright areas of the head are buff-yellow (Ridgway
LOW): These differences have not hitherto been properly
allocated, as the sex of the few specimens previously collected
was unknown,

With regard to the head-markings, it may be further noted :—
The black line through the eye is continued over the inner side
of the ear, and the buffy cheek-colour runs round the lower edge
of the ear to the back of this. There occurs also below the eye
a narrow black line running from the upper lip to the roots of
the cheek-vibrisse. The buff of the thigh continues along the
outer side of the foot and spreads over part of its upper surface,
while the toes are largely buffy.

Dr. Lyon remarks of this and the allied D. murina from
Western Borneo, that their scarcity in collections may probably
be due to some peculiarity of habit making them ditticult to

38 MR. CG. BODEN KLOSS ON

secure. My experience of D. frenata is that it does not come
to traps, as do other Tupaiide. It runs about on the ground or
along the roots of large trees, and being an extremely shy and
inconspicuous little animal, is not easy to see or to shoot.

Measurements.—Ears of the series, 12-13 mm. For other
measurements see table, p. 69.

12. PreROoPUS HYPOMELANUS CONDORENSIS Pet.

1602. 9 ad. Koh Mak. 19th December, 1914.

1628. 2 juv.; 1629. 2 subad.; 1630. 2 subad. Koh Mak.
20th December, 1914.

IGob, iad. WhGDi oad. 1658203 msulbadaaiue leaiuenmce
21st December, 1914.

“Tn the three fully adult specimens, one from Koh Mak and
two from Koh Rang, the forearm measures 138-142 mm., the
skull (total length) 62-67, the lower jaw (condyle to front of
meisors) 49°8—53°5, and the maxillary tooth-row (c'—m’*, crowns)
24—25-5,

“Though not very large, this series beautifully illustrates the
individual colour-variations in this bat. I should not be sur-
prised if they represent approximately the extremes in colour,
besides, of course, several intermediate stages. The mantle varies
from a tinge a little paler than “ hazel” (Ridgway, pl. iv. no. 12*:
g subad,, Koh Rang, 1658), through several darker tinges, to
warm glossy seal-brown (pl. iii. no. 1: 9 ad, Koh Mak, 1602).
The underparts are, in two individuals (@ juv., Koh Mak, 1628,
and gd ad., Koh Rang, 1656), quite or very nearly uniform dull
seal-brown ; in one (@ ad., Koh Mak, 1602) there is just a faint
suggestion of a deep chocolate tinge in the central area of the
breast and belly, in others this same area becomes gradually
lighter in colour, the extreme being a quite pale shade of russet
(considerably paler and more buffy than Ridgway’s pl. 11, no. 16:
@ subad., Koh Mak, 1629). The back varies from blackish
seal-brown to a tinge almost like ‘ burnt umber, and is some-
times nearly uniform, but more often thinly, sometimes rather
thickly, sprinkled with greyish hairs.

‘The variations as described above are entirely independent
of the sex and age of the individuals.

“These are the first specimens of condorensis I have seen
since working out the genus Péeropus for the British Museum
Catalogue of Chiroptera. My material then was the type in
Berlin and the paratypes in Paris, from Pulo Condor (off S.E.
Lower Cochin-China), all of which are mounted and faded, and
an old and much faded skin in the British Museum labelled
‘Siam’ (Finlayson). Judging from the present fresh and well-
preserved series, 1 am inclined to think that condorensis will
have to be put down as asynonym of Pt. hypomelanus tomesi.

* The colours in my ‘Catalogue of Megachiroptera’ weré named from the old

edition of Ridgway’s : Nomenclature of Colours’ (1886). To avoid confusion I use
the same hook for my description of the colours here,

MAMMALS FROM ‘SIAM. 39
But before giving any definite opinion I should like to see fresh
material from Pulo Condor.”—Knud Andersen.

Bats of the hypomelanus species are apparently strictly con-
fined to small islands, and do not occur on the mainland or on
large islands, however closely their homes may be situated to
such regions,

The vampyrus species, on the other hand, is largely of con-
tinental habitat, or, if living on islands, the animals are of greater
size and more nearly related to the mainland forms than ave those
of the hypomelanus species; and except on one of the Natuna
Islands, representatives of the two species never seem to occur
side by side.—C. B. X.|

13. PreRopUS VAMPYRUS MALACCENSIS K. And.

LOGE Osuva wl OiZens ade-wlOls cr ada5) VO%4. Oradea Kol
Kut. 23rd December, 1914.

1690. g ad. Koh Kut. 24th December, 1914.

1732. 2 subad.; 1748. d subad. Koh Kut. 26th December,
1914.

‘““No. 1671 is not nearly full-grown. . No. 1732 is perhaps
externally full-sized, and the skull very nearly so.

“Tn the four fully adult specimens the forearm varies between
190-204 mm. (this lowers the minimum given in my Catalogue
by 5 mm., but it is easy to see that my material did not show the
real extremes in size; the then available measurements of the
forearm were 195-209, but as the variation in length of forearm
in any species or subspecies of Pteropus is usually at least ten per
cent., 190-209 comes probably nearer the true extremes); the
skull (total length) measures 77°5-81, the lower jaw (from
condyle) 60-5—64, and the maxillary tooth-row (crowns) 30-30-7.

“The six specimens are not peculiar in any respect. In colour
they exhibit no more variation than usual in P¢. v. malaccensis,
i.e. a lighter, or deeper tinge of the mantle and head, and a
thinner or heavier admixture of greyish hairs on back and under-
parts.

“Pt. v. malaccensis was known to be generally distributed over:
Sumatra, including Banka and the Linga Archipelago, and north-
wards through the Malay Peninsula to Jalor and Patani. The
present series extends its range across the Gulf of Siam to the
island of Koh Kut, off S.E. Siam.

“Three years ago, in the new British Museum Catalogue
of Chiroptera (vol. i. p. 325, in the paragraph ‘ Differentiation
of species’) I hinted at the possibility that a completed material
might show a gradual transition from Pt. giganteus (* Pt. medius’
of Dobson’s Catalogue) to Pé, vampyrus (‘ Pt. edulis’). But
time was not ripe then for any final conclusion on this subject,
and I preferred, in order not to prejudice matters, simply to
record the distinguishable forms under four headings :—

(1) Pt. giganteus (two races) from India and Ceylon, north
and north-east to Nepal, Assam,and Manipur—represented in the

40 MR. C; BODEN KLOSS ON

Maldive Archipelago by (2) Pé. ariel, in Tenasserim by (3) Pé.
intermedius, and in the Malay Peninsula and Indo-Malayan
Archipelago by (4) Pt. vampyrus (six races). A fifth form,
Pi. lylei (Siam and Saigon) appears to be a perfectly distinct
species.

“‘ Since then the Mammal Survey of India, carried out under
the auspices of the Bombay Natural History Society, has enor-
mously increased our knowledge of the range of variation of
Pt. giganteus, and everything I have seen so far tends to con-
firm my belief that future systematists will be compelled to
regard giganteus, ariel, intermedius, and vampyrus as _ local
representatives *(‘subspecies’) of one species, Pt. vampyrus—
intermedius being (as the technical name was intended by me to
indicate), both geographically and in its characters, the con-
necting-link between the western (Indian and Indo-Chinese)
giganteus and the eastern (Indo-Malayan) vampyrus, and ariel,
an only slightly and probably imperfectly differentiated island
form of giganteus.’—Knud Andersen.

[The occurrence of this animal on Koh Kut is interesting, as
it must apparently have arrived there by flight from the Malay
Peninsula across the Gulf of Siam—a journey over water of
about 250 miles—since the only flving-fox known on the main-
land from Bangkok to Siam is P. lylei, a very distinct species,
while the continent west of Bangkok is apparently occupied by
another species, P. ditermedius,—C. B. K, |

14, CYNoPTERUS BRACHYOTIS ANGULATUS Miller,

15. CYNOPTERUS BRACHYOTIS BRACHYOTIS S. Miller.

1437-1440, All g ad. Koh Chang. 7th December, 1914.

1508. gad, Koh Mehsi Hast. 13th December, 1914.

1579. g ad.; 1580, ¢ ad.; 1581, 92 ad.; 1582. Q ad,; 1583.
@ ad. Koh Kra. 17th December, 1914,

1599. g ad.; 1600. 2 subad.; 1601. 9 ad, Koh Klum:
18th December, 1914.

1786, g ad. Koh Kut, 30th December, 1914.

“The four specimens from Koh Chang I refer to C. 0.
angulatus ; all the others are undoubtedly C, b, brachyotis.

‘“‘T have carefully examined and measured all the specimens.
Unfortunately, all being skins*, I have been unable to verify
the collector’s measurements of the ears, as given on the label of
each specimen, Not that I have the slightest doubt of the
accuracy of his measurements, but what I do have is a strong

* J should like to take this opportunity of urging on collectors the advisability of
preserving én alcohol a fairly good number of the bats obtained. Skins are in-
dispensable for a study of the colours of the fur, but the shape and size of the ears
and (an leaf-nosed bats) the details of the nose-leaves are in this group of mammals
such important items that I often, during my work for the Catalogue, have had to
deplore the now almost universal habit of experienced collectors of making nearly
every good specimen of a bat into a skin. My earnest advice is, if only one
specimen is obtained, put it in alcohol, if several, put about half of them (and not
only the most hadly damaged ones) in aleohol.—K, A,

MAMMALS FROM SIAM. 41

suspicion that his method of measuring the ears of a Cynopterus
is different from mine, and his measurements, therefore, not
directly comparable with those given by me in the new ‘Cata-
logue of Chiroptera.’ If they were, then the four specimens
from Kohn Chang, though having a cranial rostrum perfectly
similar to that of C. b. angulatus, would possess ears as long
as, or (in three out of four cases) conspicuously longer than,
any ©. sphinw sphinw I have seen. It is only natural, I think,
that before admitting the existence of such specimens £ should
like to verify their characters on alcohol material. Also the
‘ collector’s measurements’ of the ears of the ten C. b. brachyotis
are unusually large.

“In the specimens of brachyotis the forearm measures 58-
66 mm. (57-66: I add everywhere in parentheses, for com-
parison, the corresponding measurements taken by me on the
large series examined for the ‘Catalogue of Chiroptera’), in
the four angulatus 66-70 (65-72); ear, collector's measur ements,
brachyotis ‘15°5-18’ (15-17), angen thin ‘183-21’ (16 6-18);
skull, lambda to gnathion, brachyotis 28°5-29°8 (27-30°7), angu-
latws 32-33 (30°5-33°2); rostrum, orbit to nares, brachyotis 6-7—
73 (6-7-4), angulatus 7°2-7:5 (6: 5-8:2); mandible, brachyotis
21-°5—22°8 (20°2-22°8), angulatus 24-2—-25 (22°8-25°5); maxillary
teeth (crowns), brachyotis 9:2—-10:4 (8°8-10°4), angulatus 10°2—
10-8 (10-2-11°3),

“Tf all the fourteen specimens are placed in a row the
practised eye will easily pick out the four angulatus, owing to
a different, but hardly describable, tinge of the colour of the
upper side. If, similarly, the skulls are placed in a row those of
angulatus ave, of course, distinguishable at a glance by their
conspicuously longer size (see measurements above).

‘“¢ Of course, if a form really does exist, in the north of the
Malay Peninsula, in the islands off 8,K. Siam, and possibly some-
where else, which possesses the skull of angulatus*, but the ears
of sphinwt, then an entirely new and unsuspected element is
introduced into the genus. But unless and until the existence
of such a form is properly established, I should think it yather
premature to discuss its probable effect on our arrangement.”—
Knud Andersen.

| Dr. Andersen’s notes seem to eall for some remark, With
regard to the measurement [ am satisfied that that used by me
is the same as his, 2, e. ‘ from orifice’ (to the extreme tip under-
stood). It is the only one of the outer external side that can be
taken with any certainty and uniformity, and is so obvious that
it suggests itself to every collector, The only possible alternative
is the length of the taner external side from tip to base on the
crown—quite another thing and not to be confounded with the
former.

* Cranial rostrum (orbit to nares) less than one-fourth of skull (lambda to
gnathion).

7; Kars from orifice (18-205 mi.).

49 MR. C. BODEN KLOSS ON

Now ©. angulatus Miller, does have long ears, for the measure-
ments of the type series are given as 18-21 mm.* The type-
locality is Trang, S. Peninsular Siam or, to put it another way,
Central Malay Peninsula. Recently Messrs. H. C. Robinson
and K, Seimund obtained a series of bats from Bandon (about
100 miles to the north of this) and the adjacent islands of Koh
Samui and Koh Pennan, with ears which they found to range
between 18°5-21 mm.‘7, while I, again, consider my Koh Chang
specimens to have ears of 18°5-21 mm. It is impossible to
ignore the evidence of so many independent observers, which
goes to prove that a bat with the long ears of Dr. Andersen’s
sphinw veally does occur in this region.

The question then arises as to what is the angulatus of
Andersen, based on a large mass of heterogeneous material from
an extensive region, ranging from Assam and Annam to Sumatra
and the islands off its western coast. Though it includes six of
Miller’s type series, three of which have the ear-length recorded
as above, our author does not seem to have taken this statement
into consideration.

Now Dr. Andersen recorded C. brachyotis brachyotis as also
occurring throughout Sumatra and the Malay Peninsula as far
north as Trang (and now in the islands of 8.E. Siam), so that if
angulatus is to be accepted as a form of brachyotis, as he desires,
we have an instance of two subspecies of the same species living
side by side; or, in other words, two geographical races or local
forms occurring in the same place—a thing which most zoologists
will flatly refuse to admit: they must either, be the same thing
or forms of two species.

Again, if on account of the long ears (which I think must be
accepted as occurring in the Malay Peninsula and islands of Siam
at any rate) we regard angulatus asa form of sphinw, we should
have, if the long-eared angulatus occurs there too, a similar
questionable state of affairs existing in Sumatra, which is in-
habited by titthecheilus, also, according to Andersen, a form of
sphinx. So we are left with three alternatives: either angulatus
has no real existence, the material forming it being part sphina
and part bracha yotis—not very probable; or it is a very plastic
and comprehensive form of the latter, of which the typical race
is non-existent in Sumatra and the mainland; or it is an in-
dependent species. In the last case its central position is good
reason for the possession of characters appertaining to both the
other species :—long ears of sphinw, short rostrum of brachyotis,
and medium size. And to explain the occurrence of all species in
one locality to-day we may imagine sphinw extending eastward
from Ceylon, ang gulatus southward from Indo- China, and brachy-
otis westward from, for present purposes—say, Borneo: all con-
verging on Sumatra—probably the home of the other section
of the genus WViadius. Or conversely, all species of Cynopterus

* Miller, Proc. Acad. Nat. Sciences, Philadelphia, 1898, p. 316.
ap Robinson & Kloss, Journ. F. M.S. Museums, vol. y. pp. 115, 1384 (1915).

MAMMALS FROM SIAM. 43

originated in the latter locality, and in the race for expansion
the last got left at the post.—C. B. XK].

16. RaruFA MELANOPEPLA LEUCOGENYS, subsp. n.

Sciurus javensis Gray, P. Z. 8. 1861, p. 137.

Sciurus bicolor de Pousargues, Mission Pavie, Indo-Chine,
Etudes Diverses, ili. p. 582 (1904).

Ratufa melanopepla Gyldenstolpe, Arkiv fér Zoologi, Band 8,
No. 23, p. 15 (1914).

Type. Adult female (skin and_ skull), No. 1912/C.B.K.
B.M. No. 15.11.4.48. Collected at Lem Ngop, 8.E. Siam, on
loth January, 1915.

Characters.—Like R. m. peninsule Miller (Proc. Washington
Acad. Sci. vol. ii. p. 71, 1900; id., Smithsonian Miscellaneous
Collections, vol. lxi. No. 21, p. 25, 1913), but yellow of cheeks,
fore limbs, and under surface markedly paler than the respective
areas in that form, yellow on thighs more extensive and con-
tinued along the sides of the feet on to their upper surfaces,
where it occupies a considerable area, while the yellow of the
fore limb extends to the base of the toes above.

Colour.—Upper surface and entire tail brownish black; a
fairly conspicuous russet patch on the nape. Under surface pale
orange-yellow. Cheeks to base of ear, but not reaching the eye,
lower sides of neck, greater part of upper side of fore limb to base
of toes ivory-yellow, becoming cream-colour on the posterior part
of the fore limb. The yeilow area of the under thigh continued
as ivory-yellow along the outer side of the foot and on to the
upper surface of the latter, where it occupies almost half of the
area between the ankle and bases of the toes. Sides of muzzle
like throat, but chin black.

Skull and teeth.—Possess apparently no constant features
which will separate them from topotypes of FR. m. peninsula,
from Trang, and others from Banden, 100 miles to the nortli in
Peninsular Siam. On the whole, the bulle appear to be a little
longer.

Measurements.—Kiars of the type 30:5, of the series 29:5—
30°5 mm. For other measurements see table, p. 69.

Specimens examined.—Three, the type and two adult males
from the same locality.

Remarks.—This race is separated from that of the Malay
Peninsula by its pale yellow coloration and large yellow patch
on the hind foot, while the yellow of the fore limb is greater in
extent, reaching the base of the toes.

The individual obtained by Gyldenstolpe (loc. cit. swpra) in
Eastern Siam, south of Korat. is, judging from its measurements,
of this form. Ratufa pheepepla Miller (Smithsonian Miscel-
laneous Collections, vol. Ixi. No. 21, p. 25, 1913), from South
Tenasserim should occur also in Western Siam. It is like the
animal of the Peninsula, but larger (hind foot 80-89, skull 74—
78 mm.). Flower (P. Z. 8S. 1900, p. 355) records a black-and-

44 MR. CU. BODEN KLOSS ON

yellow Giant Squirrel from Phrabat, and observed another near
Paknam Kabin which he believed had tufted ears. This feature
would apparently indicate the presence near Bangkok of Ratufa
gigantea McClelland, which, according to Wroughton (Journ.
Bombay Nat. Hist. Soc. vol: XK. ps 890, 1910), also occurs 1n
N. Siam. It is a large black-and-buff animal (head and body
417 mm., hind foot 87, skull 80), with the yellow on the fore
limb confined to the inner side.

17. RAtUFA MELANOPEPLA SINUS, subsp. n.

Type. Adult female (skin and skull), No. 1733/C.B.K.
B.M. No. 15.11.4.41. Collected on Koh Kut Id., S.E. Siam, on
26th December, 1914.

Characters.—Like It. m. peninsula Miller, but uniform black
above with the under surface rather more ochraceous and ex-
tending slightly to the upper surface of the hind foot, while the
yellow of the fore limb extends to the base of the toes above.
Nasals rather longer,

Colour.—Upper surface and entire tail clear black. Under
surface varying from ochraceous buff to ochraceous orange and
ochraceous tawny in centre of abdomen, Cheeks to base of ear,
sides of neck, and upper and inner side of fore limbs pale orange-
yellow, deeper om the inner side of fore limbs, A small area of
ochraceous buff on the outer and upper sides of the hind foot.
Sides of muzzle like throat, but chin black,

Skull and Teeth.—-Resemble those of R. m. peninsule and
Rh. m. lewcogenys, except for the greater length of the nasals, the
posterior terminations of which are more in line with those of
the premaxillaries. The bulle are apparently a little shorter
and broader.

Measurements.— Ears of the type 30, of the series 27-30 mm.
For other measurements see table, p. 69.

Specimens examined.—Six adults, 3 males and 3 females.

Remarks.—This form differs from that of the adjacent main-
land in being uniformly clear black above without any nuchal
spot. The yellow areas are deeper in shade, those of the head,
neck, and fore limbs being of about the same tone as the abdomen
of fi. m. lewcogenys, while the patch of yellow on the upper side
of the hind foot is smaller and somewhat disconnected from the
yellow of the thigh.

This and the preceding race both differ from the Peninsular
form in the constant presence of a considerable amount of yellow
on the hind and fore feet. Jnéer se, the differences in colour are
very marked, though those of the dorsal surface may be only
seasonal. Size is about the same in all (a trifle larger in the
S.E. Siamese forms), but the longer nasals of the island race
further serve to distinguish it from both the others.

Tt may be pointed out that the forms from the islands of the
Malayan part of the Peninsula (Teratau, Langkawi, Penang, and
Tioman), 7. é., southern island forms, are all instantly separated

MAMMALS FROM SIAM. 45

from the mainland races and from Siamese island forms (Telibon,
Samui, Pennan, and Kut) by their deep ochraceous-tawny under
surfaces and by the greater amount of black on the inner side of
the hind limbs.

Though the island of Koh Chang is larger than Koh Kut,
nearer to the mainland, and in much shallower water, no form
of Giant Squirrel cceurs on it. Yet the Ratufa of Siam was
found immediately opposite the former, whereas all along the
coast in the vicinity of Koh Kut it was not met with.

18. SclURUS FERRUGINEUS CINNAMOMEUS Temm.

Sciurus cinnamomeus Temminck, Esq. Zool. Guiné, 1853, p. 250;
Wroughton, Ann. & Mag. Nat. Hist. ser. 8, vol. ii. p. 396 (1908) ;
Gyldenstolpe, Arkiv for “Loologi, Band 8, No. 23, p. 12 (1914).

Sciurus splendens Gray, P. 7.8: 1861, p. 137.

Sciurus ferrugineus Anderson, Zoological Researches, p. 244
(1878); de Pousargues, Mission leew ie, Indo-Chine, Etudes
Diverses, ii. p. 526 (1904),

Sciunes finlaysoni Flower (partim), P. Z. S. 1900, p. 355.

Sciurus finlaysoni, Type B (Se. ne Bonhote, IP, 3 Sie
1901, vol. 1. p. 53.

6 males, 7 females, Ok Yam, Franco-Siamese Boundary ;
2 males, 3 females, Klong Yai; 2 males, 1 female, Klong Menao;
3 males, 2 females, Lem Ngop, $.E. Siam.

The above specimens show a certain amount of variation in
colour. On the whole, however, they most nearly approach
Sciurus splendens, var. 2 of Gray (=cinnamomeus ‘emm.).
“Top of head (and top of back by inference) and tail dark and
very intense red-bay ; side of the back, under sides of the body,
and tip of tail paler red-bay.” A few approximate to var. 3.
** Uniform pale bay, like the side of var. 2; tail and middle of
the back rather darker and brighter; tail without pale tip,”
though the latter shows some -signs of “bleaching” at the
extremity. The majority of the collection have the sides of
the head grizzled greyish, and there are traces of grizzling on the
fore limbs: these latter features , together with an indication of
grizzling on the thighs, being most pronounced among the five
exainpies from Lem Ngop, the western extremity of the series.
Again, two or three approach var. 1, “all over dark and very
intense red-bay,” except that they have no “ white spot on each
side of the base of the tail.”

Measurements.—Kars of the series 19-23 mm. For other
measurements see table, p. 70.

I do not know from what locality Temminck’s Seiurus cinna-
momeus came, and we have also no details as to the provenance
of the animals on which Gray founded his species, beyond the
fact that their collector, Mouhot, travelled widely in Siam and
Cambodia ; but 1t appears to me that when the Red Squirrel is
fully known over the whole of its range it will be necessary
to recognise several geographical races; to all of which the

46 _ MR. C. BODEN KLOSS ON

opinion of Anderson (op. cit. p. 245) with regard to iS. eina-
momeus will likewise apply, 7. é., that they are only local forms
of S. ferrugineus.

Even in the present series, collected along a 50-mile stretch of
coast, there are indications of geographical variation ; for the
eight specimens from the northern stations (Lem Ngop and
Klong Menao) most nearly resemble Gray’s var. 3, while, with
the exception of three or four individuals like them, the eighteen

darker southern animals come nearer var. 2.

19, ScluRUS FERRUGINEUS FRANDSENI, subsp. n.

Type. Adult male (skin and skull), No, 1502/C.B.K. B.M.
No. 15.11.4.85. Collected on Koh Chang Id., S.E. Siam, 12th
December, 1914.

Characters.—Like S. f. cinnamomeus of the adjacent main-
land, but with chin, throat, sides of head, outer sides of fore and
hind limbs grizzled blackish or olive-brown.

Colour.—Top of head, upper part of body, and tail intense
shining red-bay, becoming more fulvous towards the sides and
on the thighs and behind the ears, the hairs black-tipped except
on the distal half of the tail, which is clear reddish chestnut,
somewhat bleached to fulvous at the extremity.

Muzzle, sides of head, chin, throat, shoulders, and sides of body
olive-brown variably annulated with buff-yellow, strongest on
threat and flanks; the shoulders, outer sides of fore limbs and
thighs becoming black, finely annulated with buff. Entire
under surface, except chin and throat, rich tawny, this colour
extending to the fore feet and also to the hind feet, where it is
mingled with black, There is a faintly indicated grizzled line
down the centre of the chest and abdomen. Kars like the hind
feet, their bases posteriorly dull ochraceous buffy.

Skull and Teeth.——As in the mainland race.

Measurements.—Type: Kar, 22 mm. Skull: basilar length,
44-4; brain-case breadth, 20; proximal breadth of nasals, 4 ;
distal breadth of nasals, 8. Hars of the series, 19-22 mm. For
other measurements see table, p. 70.

Specimens examined.—Sixteen, 9 males and 7 females, all from
the type-locality.

Remarks.—In some of these animals the extent of black on
the head is almost sufficient to form a black line between the
crown and the grizzled portion of the face, and the fore feet are
also partially black; in others the under surface is somewhat
more orange than in the type.

Phe definition of Sciur us splendens var. 4, Gray (P. Z.8. 1861,
p. 137), apples to some extent, as does alls Anderson’s s deserip-
tion (Zool. Res. pp. 245-6) of an example considered by him to
be S. siamensis Gray. The locality from which these latter two
specimens came 1s unknown and, though I am unaware that
Koh Chang has been previously visited by any naturalist, such
may have been the case. The characters of S. f. frandseni are

MAMMALS FROM SIAM. AT

so regular throughout in the series of sixteen as to render it an
extremely distinct race, even though, as recorded above, animals
from Lem Negop (the nearest point of the mainland), having
indicatious of grizaling on the thighs, tend to connect it’ with
the more typical cennamomeus animal, It is named after
Captain H. EH. Frandsen, R.N.R., Denmark, to whom I am
indebted for much assistance and interesting information while
in §.E. Siam.

20. ScruRuS ALBIVEXILLI, sp. n.

Type. Adult male (skin and skull), No. 1724/C.B.K. B.M.
No. 15.11.4.46. Collected on Koh Kut Id., S.E, Siam, 25th
December, 1914.

Characters and Colour.— Black throughout except the extremity
of the tail, which is white.

Skull and Teeth.—As in S. cinnamomeus.

Measurements.—Vype: Har, 20°5> mm. Skull: basilar length,
436; brain-case breadth, 25°2; proximal breadth of nasals, 7 ;
distal breadth of nasals, 4:4. Ears of the series, 19-21°5. For
other measurements see table, p. 70.

Specimens examined.—Twenty-three, 12 males and 11 females,
all from the type-locality.

Remarks.—S. albivewilli is somewhat variable in respect of the
white tail-tip. In some animals the last 3 to 4 inches of the tail
are white, and there is a white ring close to the bases of the hairs
for the distal three-fourths of the tail; in others there are no
annulations, and the pale tip is reduced to a bunch of greyish
hairs at the extreme end.

Two other forms of Black Squirrel occur in Indo-China: S. noa
Wroughton (Ann, & Mag. Nat. Hist. ser. 8, vol. ii. p. 397, 1908),
in the neighbourhood of Siracha, on the eastern shore of the
Inner Gulf near Bangkok, and S. germaint Milne-Edwards (Rey.
Zool. 1867, p. 193) on Pulo Condor, south-east coast of Cochin-
China: both are black throughout, but the latter is much smaller
than the other. On distributional grounds it seems impossible
to treat them and the present form as local races of one species,
as there is no geographical connection, the mainland every-
where in the vicinity of Koh Kut being occupied by the red
AS. clnnamomeus.

The latter is, however, known to develop a white tail-tip, and
it is possible that through S. f frandseni, with its black-tipped
upper pelage, blackish fore limbs and thighs, a connection may be
traced between the Koh Kut animal and S. einmamomeus. It is,
however, very slight,

21. Tamtors Ropoutpent M.-H.

Sciurus rodolphi Milne-Kdwards, Rev. et Mag. de Zool. xix.
p. 227 (1867); id., Rech. Mamm. 1871, p. 162; de Pousargues,
Mission Pavie, Indo-Chine, Etudes Diverses, iii. p. 528 (1904),

48 MR. C. BODEN KLOSS ON

Sciurus macclellandi rodolphi Bonhote, Ann. & Mag. Nat. Hist.
Sela (Vole vip. 04 (1900); id PaZaSesl 90M ep lO:

1 male, 1 female, from Lem Ngop, 8.E. Siam.

I have not seen specimens of this squirrel from the type-
locality (Cochin-China), nor is Milne-Edwards’s description ac-
cessibie to me, but Mr. Bonhote’s remarks (loc. cit. swpra) on
examples from Cochin-China and Annam appear to apply to the
individuals obtained in 8.E. Siam.

The dark stripes are all grizzled with brown, the median black
one being divided down the centre by a grizzled brown line, and
the four light dorsal stripes are of equal breadth and distinctness,,
as stated by Mr. Bonhote; but while the outer two are cream-
coloured, the inner pair are buff-yellow and show none of the
pink tinge noted by him. The underparts are buff-yellow, not
ferruginous, but this may be a matter of terms. The white tufts
of the ears are black at their bases.

Measurements.—Ears, 13 mm. For other measurements see
table, p. 72.

Another form of Tamiops found in Siam is 7. novemlineatus
(Miller), which inhabits the Malay Peninsula, certainly as far
north asthe Isthmus of Kra. Bonhote (P. Z. 8. 1901, i. p. 54) has
described, under the name songensis, animals obtained at
Raheng and Nan, but these appear to differ from the Tenas-
serim barbei by just the same characters as does novemlineatus,
so that it is doubtful whether they are really distinct from the
latter.

22. MENETES BERDMOREI MOUHOTII Gray.

Sciurus mouhotit Gray, P. Z.S. 1861, p. 137.

Sciurus pyrrhocephalus Milne-HKdwards, Rev. Zool. xix. 1867,
p- 225; de Pousargues, Mission Pavie, Indo-Chine, Etudes
Diverses, ii. p. 528 (1904).

Funambulus berdmorei Flower, P. Z.8. 1900, p. 359.

Menetes berdmorei Gyldenstolpe, Arkiv for Zoologi, Band 8,
No. 23, p. 15 (1914).

Menetes berdmorei mouhoti Thomas, Journ. Bombay Nat. Hist.
Soe. vol. xxiii. p. 23 (1914).

3 males, 3 females, Lem Ngop: 2 males, Klong Menao:
3 males, 2 females, Klong Yai, S.E. Siam; 1 male, 4 females,
Ok ‘Vem, Franco-Siamese Boundary.

I must confess that I find these specimens somewhat difficult
to place. The only material available to me for comparison
consists of four examples of Jf. berdmorei berdmorei from Mar-
taban and Moulmein (Cat. Mamm. Indian Mus. specimens a, b,c, @)
and a series of thirteen from Bandon, Peninsular Siam, whieh
are apparently conspecific with those (cf. Robinson & Kloss,
Journ. F. M.S. Museums, vol. v. p. 121, 1915), all of which are
clearly distinguishable from the present series by the conspicuous-
ness of the median dorsal and upper lateral blackish lines and
somewhat smaller size.

MAMMALS FROM SIAM. 49

Thomas, however, has recently reviewed the races of this
squirrel (loc. cit. supra) and, though one would, on geographical
grounds, place the present animals in the form mowhotiz, one can
also regard part of the series as of that subspecies as defined
by him and allot the remainder to his new race consularis of
Northern Siam. If we only knew the exact type-locality of
M. 6. mouhotir, given vaguely as Cambodia (Gray, foc. cié. supra),
we should probably find that these S.E. Siamese animals are
geographically, as they are in appearance, intermediate between
the two.

Gray, who only had ene specimen when describing mouhotiz,
makes no mention of any dark stripes between the upper pale
ones, and consularis resembles his type in that respect. Thomas,
however, in extending the range of mouhotii from south of
Bangkok to Cochin-China, reports three inconspicuous black
stripes on the back between the upper pale ones as in berdmoret
berdmorez: mouhotii is white or whitish below, consularis
yellowish white. ‘The series of 16 animals from 8.E. Siam have
backs ranging from three to no dark stripes and under surfaces
from white to yellowish white.

The individuals from Eastern Siam (Korat) referred by
Gyldenstolpe (doc. cit. supra) to M. berdmorei require further
examination.

Thus, exclusive of the two island races described below, the
following forms seem to inhabit our region: the true berd-
morei in Peninsular and perhaps Western Siam ; consudaris in
Northern and perhaps Eastern Siam; and mouwhotii in South-
eastern Siam.

The species seems to increase in size from west to east; the
three forms mentioned here are all larger than those from Burma
and the Malay Peninsula, and Jf. 4. moerescens from Annam
(Thomas, loc. cit. supra) is believed to be larger also.

Measurements.—Ears of the series, 18°5=21 mm. For other
measurements see table, p. 71.

23. MENETES BERDMOREL UMBROSUS, subsp. n.

Type. Adult female (skin and skull), No, 1449/C.B.K.
B.M. No. 15.11.4.97. Collected on Koh Chang Id., S.E. Siam,
8th December, 1914.

Diagnosis.—Like M. 6. mouhotit from the adjacent mainland,
but darker above; the three upper dark stripes very faintly
indicated by a blackish wash, the upper pale stripe narrower and
‘a deeper buff, the lower also deeper in tint but less distinct,
approaching in colour the outer side of the thighs; the sides of
the abdomen between the limbs also darker, Under surface
somewhat more deeply buffy. ‘Tail more ochraceous, blacker and
much less hoary.

Skull and Teeth.—As in the mainland animal.

Measurements.—Wars of the type, 20, of the series, 17°5-
20 mm. For other measurements see table, p. 71.

Proc. Zoo, Soc.—i916, No. TV. 4

50 MR. CG. BODEN KLOSS ON

Specimens examined.—-Six, 3 males and 3 females.

Remarks.—There is extremely little variation in the series
from Koh Chang, which is_ easily separated on the above
characters from the mainland form. The dark dorsal stripes,
though obsolete, are indicated by an increase in the amount of
black annulation, but are not so intense in colour as the area

between the lateral pale stripes.

24. MENETES BERDMOREI RUFESCENS, subsp. n.

Type. Adult female (skin and skull), No. 1740/C.B.K.
B.M. No. 15.11.4.93. Collected on Koh Kut Id., 8.H. Siam,
26th December, 1914.

Diagnosis.—Lighter and more rufous above than the neigh-
bouring continental form, dark dorsal stripes absent, dark lateral
stripe not deeper in colour than the back; upper pale stripe
a little less intense, the lower much less distinct ; sides of body °
bordering the abdomen considerably darker. Under surface
slightly richer buff. Tail darker and much less hoary.

Skull and Teeth—As in the mainland animal.

Measurements.—Ears of the type, 17, of the series, 17-22 mm.
For other measurements see table, p. 71.

Specimens ecamined.—Nineteen, 11 males and 8 females.

Remarks.—As in the series of 18 examples from the adjacent
mainland dealt with above, there is a certain amount of variation
in the dorsal area of this race, some examples having the dark
lateral dorsal stripes present toa slight degree and the median
one just indicated: when this is the case the area between the
pale lateral stripes is proportionately darker also. The other
differences, however, are consistently maintained and the series
further includes the largest animals obtained in this region.

25, EPIMYS JERDONI MARINUS, subsp. n.

Type. Adult male (skin and skull), No. 1455/C.B.K. B.M.
No. 15.11.4.160. Collected on Koh Chang Id., S.E. Siam,
9th December, 1914.

Characters.—Resembles Epimys jerdoni bukit (Bonhote), but
with the white of the under parts hardly ever extending to the
foot, and tail rather shorter. Skull with smaller bulle, larger
palatal foramina, and broader interpterygoid space.

Colour.—Above ochraceous tawny, much darkened or streaked
by the exposed tips of the numerous stiff spines which have
gveenish horn-coloured bases. Base of fur grey. Below yellowish
white to the base of the hairs, extending over the fore limbs to
the hands, but not quite reaching the hind feet, which are white
with brownish centres. Tail bicoloured with a dark tip.

Skull and Teeth.—Like those of H. 7. bukit, but with smaller
and more flattened bulle; broader interpterygoid space, the
outline of which iy more angular owing to the straighter anterior
margin; palatal foramina larger, nearer the incisors, and the

MAMMALS FROM SIAM. yt

nasals slightly more projecting anteriorly. In all except the
first of these characters the skull more nearly resembles Z. j. pan
Robinson & Kloss, from Koh Samui Id. of the opposite side of
the Gulf, but differs in the bulle, which in that race agree with
EL. j. bukit.

Measurements.—Kar of type,20 mm, For other measurements
see table, p. 72.

Specimens examined.—Iwenty-two from Koh Chang and
twenty-three from Koh Kut.

Remarks.—Besides externally closely resembling Z. j. bukit,
this race is also very similar in appearance to #. y. pan. While,
however, in the former the white of the under parts nearly
always reaches to the foot, in marinus it generally just fails to
do so, and in the latter it always stops considerably short of the
ankle.

Although these rats were exceedingly common on the two
islands none was met with on the mainland, so that I have had
to compare them with animals from the Malay Peninsula, which
Bonhote states (Fasciculi Malayenses, Zoology, part 1, p. 27)
exactly agree with those of Siam: in which case L. lepidus
Miller, founded on a single adult individual from Southern
Tenasserim (an intermediate locality), is also probably an example
of H.j. bukit. The position of this latter with regard to the
true #. jerdoni (Blyth) of Sikkim is not fully known, but from
the few details recorded of Darjiling specimens (‘Thomas, P. Z. 8.
1881, p. 5388; Blanford, Faun. Brit. India, Mammals, p. 411),
it would appear that the typical animal is a smaller form having
« tail actually, and so relatively much, longer than bukit (and
therefore still longer than marinus).

In my experience the yerdont rat is by no means common in
the Malay Peninsula, and I failed to meet with it in 8.E. Siam.
Of the small islands of these areas, #. j. pan had oniy recently
been discovered on Koh Samui, and I was therefore much
surprised to find a form occurring in great abundance on the
two larger islands of the Chantabun Archipelago, where it used
to come into my camp at twilight in search of food.

The Koh Kut animals seem to be a trifle smaller than those
of the type-locality, but apart from that I can detect no difference
whatever.

26. EPIMYs SURIFER FINIS, subsp. n.

Type. Aged male (skin and skull), No, 1885/C.B.K. B.M.
No, 15.11.4.117. Collected at Klong Menao, 8.E. Siam, 11th
January, 1915.

Characters.—Like Epimys surifer Miller, from Peninsular Siam
(Trang), but duller; white of under surface normally extending
to the ankle and over the bases of the vibrissee.

Colouwr.— Upper surface ochraceous tawny, clouded on the back
by the dark tips of the flattened spines. Under parts white,
extending to the hind feet and to the roots of the vibriss, but

A*

52 MR. C. BODEN KLOSS ON

not always to the hands. Tail averaging longer than head and
body, bicoloured with white tip. Hands and feet white.

Skull and Teeth.—Resemble those of the typical race and show
no characters which will consistently serve to distinguish them
from it.

Measurements,—Ear of type, 245 mm. For other measure-
ments see table, p. 73.

Specimens examined.—Twenty-six: 3 from Ok Yam, 5 from
Klong Yai, and 18 from Klong Menao.

Remarks.—The above series has been compared with a large
series of topotypes from Trang, Peninsular Siam, and also with
a number recently obtained in Bandon (about 100 miles to the
north of that locality), and distinctly differs as pointed out ;
though, as is always the case with two neighbouring continental
races, the extremes of the two series closely resemble each other.
The extension of the white area to the foot occurs in the great
majority of examples from 8.E. Siam, whereas in series from
Peninsular Siam the contrary is the case.

Epimys surifer has only previously been recorded from Siam
by Gyldenstolpe (Arkiv for Zoologi, Band 8, No. 23, p. 16, 1914),
who obtained a single example on the Korat Plateau.

27. EPIMYS SURIFER CHANGENSIS, subsp. n.

Type. Aged male (skin and skull), No. 1492/C.B.K. B.M.
No. 15.11.4.142. Collected on Koh Chang Id., 8.E. Siam, 11th
December, 1914.

Diagnosis.—Like H. s. jinis, but with tail averaging shorter
than head and body, the dark speckle of the upper surface much
coarser and the white of the under parts more extensive on
limbs, broadening, in some examples across the body and
spreading up the sides of the muzzle and over the upper side
of the fore limb.

Measurements.—Kar of type, 25mm. For other measurements
see table, p. 73.

Specimens examined.—Thirty-three from the type-locality.

Remarks.—The tendency in this race to develop a white fore
limb and a short tail approximates it to #. s. manicalis Robinson
& Kloss *, from Koh Pennan on the opposite side of the Gulf
of Siam, from which, however, it is clearly distinguished by its
much duller upper colour.

28. EPIMYS SURIFER KU'TENSIS, subsp. n.

Type. Aged male (skin and skull), No. 1710/C.B.K. B.M.
No. 15.11.4.151. Collected on Koh Kut Id., 8.E. Siam, 25th
December, 1914.

Diagnosis.—Like EH. s. changensis, but a trifle less tawny and
with less tendency for the white areas to increase (in this coming
nearer to the mainland form); anterior root of the zygomatic

* Ann. Mag. Nat. Hist. ser. 8, vol. xiii. p. 230 (1914).

MAMMALS FROM SIAM. 53

arch ‘much narrower than in the two preceding races, this
diminution markedly increasing the size of the infraorbital
foramina as seen from above.
Measurements.—Har of type, 24 mm. For other measurements
see table, p. 73.
Specimens examined,

Twenty-nine from Koh Kut.

29, KPIMYS SURIFER PELAGIUS, subsp. n.

Type. Adult male (skin and skull), No. 1659/C.B.K. B.M.
No. 15.11.4.109. Collected on Koh Rang Id., S.E. Siam, 22nd
December, 1914.

Diagnosis.—Colour bright clay, duller than the foregoing races,
and further differing from the mainland animal in having the
tail shorter than the head and body, from Z. s. changensis in
showing no tendency to extension of the white area, and from
EL. s. kutensis in the broader anterior zygomatic root.

Measurements.-—Kar of type, 25mm. For other measurements
see table, p. 73.

Specimens examined.—Twenty-one, all from the type-locality.

30. HEPIMYS SURIFER CONNECTENS, subsp. n.

Type. Adult female (skin and_ skull), No. 1613/C.B.K.
B.M. No. 15.11.4.135. Collected on Koh Mak Id., 8.E. Siam,
19th December, 1914. °

MNagnosis.—Clay-coloured above, closely resembling LZ. s. pela-
gius, though with the white on the hind limb in some instances
a little reduced towards the ankle. Skull with slightly broader
nasals and rostrum, and profile less curved. This latter character
is difficult to define, but if skulls are compared when resting on
their upper sides, connectens skulls will be seen to have the
anterior palate and incisors on a lower plane than those. of
pelagius or finis; or if placed end to end the tips of the nasals
of the latter two always enter the nasal cavity of the other.
Zygomatic breadth is little greater, and palatal foramina are
shghtly larger.

Measwrements.—Kar of type, 24mm. For other measurements
see table, p. 74.

Specimens examined.—Twenty, all from the type-locality.

Remarks.—This race is somewhat intermediate between the
preceding and following forms, in that the continuation of
the white area to the foot is rather indistinct in several
examples, while in two specimens (an adult and a subadult
female) the under side of the fore limbs, a band across the chest,
and the lower part of the hind limbs are pale ochraceous tawny.

31. EPIMys SURIFER ECLIPSIS, subsp. n.
Type. Adult male (skin and skull), No. 1540/C.B.K. B.M.
No. 15.11.4.125. Collected on Koh Kyra Id., S.E. Siam, 16th

December, 1914.
Characters.—Clay-coloured ; white of lower side much reduced

DA MR. C. BODEN KLOSS ON

in breadth, not very sharply margined, and not extending to the
limbs. Jail dark with a white tip.

Colour.—A. variable clay, the dorsal area everywhere much
darkened by the bistre tips of the spines; this colour extending
over the whole of the limbs, across the chest in the form of a
broad gorget, the hairs of which have distinct grey bases, and
over the entire muzzle, which is rather browner. White of
under surface reduced to a comparatively narrow band running
from the axille to groin, 25-30 mm, wide, and to an isolated
patch confined to the throat. Fore and hind feet pale. Tail
about the same length as head and body, dark for the basal
two-thirds or three- fourths, the tip white, not sharply defined
from the rest,

Skull and Teeth.—Generally resemble the mainland race, but
with the nasals broader posteriorly, their outer margins
straighter; the palatal foramina distinctly larger, being both
longer and broader; anterior zygomatic plate ivalan « and the
zygomatic breadth a little greater,

Measurements,— Ear of type, 24-5 mm, For other measurements
see table, p. 74.

Specimens ewamined.— Nineteen from the type-locality,

Remarks.—\ts dull colour, reduced white areas, and non-
bicoloured tail render this race of Hpimys surifer the most
distinct known to me, All the Indo-Chinese forms here
deseribed are less brilliant in colour than any of the Malayan
races of the rajah-surifer group, and their tendency to dullness,
manifesting itself strongly in the island forms, culminates so
notably in the Koh Kra animal, that, with its other characters in
addition, one would readily accept it as a distinet species were
it the inhabitant of a large Jand-mass and not of a little satellite
islet.

32, HEPIMYs SURIFER TENEBROSUS, subsp, n,

Type, Adult male (skin and skull), No, 1586/C.B,.K. B.M.
No, 15,11,4.121, Collected on Koh Klum, 8.E. Siam, 18th
December, 1914.

Magnosis— Like EL. s. eclipsis above, but rather more darkened
down the median line of the back. Below the white area rather
broader, but not to the extent of the more typical forms, and
extending a little way across the groin on to the thigh. Wrists
very pale, but separated from the chest ; white area of neck a little
larger, but confined to the throat. Gorget clearer in colour and
rather less distinct, the grey bases of the hairs not visible. Tail
shorter than head and body ; bicoloured with a white tip. -

Skull and Teeth,—Like those of £. s. eclipsis, but the palatal
foramina smaller, about the same length as the mainland form,
but broader; the interorbital breadth markedly greater than
either, and the anterior zygomatic root broader.

Measurements.— Kar of type, 24mm. For other measurements
see table, p, 74.

MAMMALS FROM SIAM, dd
Specimens examined.— Hight from the type-locality.
Remarks.—This form is an intermediate stage between con-

nectens and eclipsis. The former in isolated individuals is
beginning to show the tendency towards increase in the yellow
areas, while in the present animal this, as well as the darker
coloration, is now well and constantly established. Both, how-
ever, still retain the normal bicolored tail.

It is interesting to note that in this small group of islands two
opposite types of deviation occur: in 1. s. changensis and kutensis
the trend is towards an increase of the abdominal white area,
while in this race and Z. s. eclipsis it is the dorsal colour that has
spread until it reaches a climax in the last form in conjunction
with extreme dullness of tint and blackened tail.

33. EPiIMys RATTUS, subsp.

Mus rattus Bonhote, P. Z.S. 1900, p. 194; id., op. cit. 1901,
vol. i. p. 56; Flower, op. cit. 1900, p. 361.

Mus ratius rufescens de Pousargues, Mission Pavie, Indo-
Chine, Etudes Diverses, iii. p. 528 (1904).

Epimys rufescens Gyldenstolpe, Arkiv for Zoologi, Stockholm,
Band 8, No. 23, p. 18 (1914).

I obtained at Ok Yam and Klong Yaia series of 13 rats, which
are apparently indistinguishable from the common Epimys rattus
of the Malay Peninsula, except that the white underparts more
frequently assume a light silvery shade. I have had no oppor-
tunity of comparing them with Epimys rattus robustulus (Blyth)
from Tenasserim, and therefore place them under the specific
name. The tail is longer than the head and body and slightly
paler below proximally ; the feet are whitish.

One female from Ok Yam (No. 1797) is abnormal in having
the tail slightly shorter than head and body and concoloured, the
feet dark; the upper side blackish brown and the underparts of
a colour intermediate between mouse-grey and neutral-grey.

With these I would associate four examples from Koh
Chang Id.

From the two islands, Koh Mehsi East and West, series of 13
and 15 respectively were obtained. While showing much varia-
bility among themselves, all are apparently conspecific with the
above. They range from melanotic individuals having backs
strongly suffused with blackish brown to others having that
surface of warm grizzled-brown, while underparts vary from
white to grey.

This difference in colour is not a question of sex or age, for
though juveniles generally (not invariably) have greyish under-

sides, yet these are by no means of so dark a shade as the extremes
of the adults; neither do the darker-backed individuals always
have darker underparts, though, again, this is generally the case.

The effect is to make the insular series much darker than the
mainland one, but as this character is apparently transitory, it
cannot be used for the purpose of differentiation.

Measurements.—See table, p. 79.

56 MR. C. BODEN KLOSS ON

34, EPIMyS RATTUS RANGENSIS, subsp. n.

Type. Adult female (skin and skull), No. 1669/C.B.K.
B.M. No. 15.11.4.208. Collected on Koh Rang Id., 22nd
December, 1914.

Diagnosis.—Closely resembles the adjacent mainland normal
form of #. rattws, but with the pelage a little coarser. Skull
broader throughout—rostrum, palate, interpterygoid space, basi-
occipital and zy gomata—but with smaller palatal foramina, which
do not reach a line joing the anterior ends of the molar rows.
Nasals shorter and more truncate, so that when the skulls are
reversed and resting on their upper surfaces the ends of the
nasals are not visible from above.

Measurements.—EHar of type, 22 mm. For other measurements
see table, p. 75.

Specimens examined.—Six from the type-locality.

Remarks.—Though differing very little bodily from the main-
land animal, this race is easily separated from it on cranial
characters, the short nasals and blunt muzgle being very
distinct.

35. EPIMYS RATTUS KLUMENSIS, subsp. n.

Type. Adult female (skin and skull), No. 1596/C.B.K.
B.M. No, 15,11,4.207, Collected on Koh Klum Id., 5.H. Siam,
18th December, 1914.

Diagnosis,—Size larger and pelage coarser than the previous
forms, with numerous long black piles on the rump. Oolour
above grizzled-brown and buff, darkest on the rump; below
ivory-yellow, an indistinct greyish band along either side of the
abdomen, separating it from the colour of the upper parts. Feet
parti-coloured ; tail considerably longer than body, relatively
longer than any of the other local races,

Skull generally resembling that of the mainland animal, but
more robust and with the nasals much narrower posteriorly and
longer, prolonged well behind a line jojning the anterior edges
of the orbits.

Measurements,—See table, p, 75.

S\pecimens examined.—Five from the type-locality.

Remarks,—The larger size, relatively long tail, and long,
posteriorly narrower, nasals clearly distinguish this race from
either of the preceding; particularly the latter with its short
square muzzle. It belongs, with the following forms, to the
section of the ratius group consisting of large animals heavily
sprinkled on the rump with long piles, and having large robust
skulls, which includes the similar races of Epimys pannosus and
mara Miller, H, remotus Robinson & Kloss, and is largely of
insular habitat,

36. HPiIMyS RATTUS MAKENSIS, subsp. n.

Type. Adult male (skin and skull), No, 1616/C.B.K. B.M,
No. 15.11.4,211. Collected on Koh Mak Id., 8.E, Siam, 19th
December, 1914, ;

MAMMALS FROM SIAM, OM

Diagnosis.—Closely resembles EL. r. klwmensis in colour, but
with the under surface generally slightly silvered, especially in
immature individuals. Size slightly larger, but tail considerably
shorter. Nasals relatively rather narrower posteriorly, but skull
otherwise apparently not differing from the mainland race except
in greater size and robustness.

Measurements.— Kar of type, 23 mm, For other measurements
see table, p. 75,

Specimens examined,—¥ifteen from type-locality,

37. EPIMYS RATTUS KRAENSIS, subsp. n.

Type. Adult female (skin and_ skull), No. 1550/C.B.K.
B.M. No. 15.11.4.203. Collected in Koh Kra Id., 8.E. Siam,
16th December, 1914,

Diagnosis.—The largest of the known local forms of F. rattus,
but with tail relatively shorter than in £. s. klumensis. Colour
like #. r. makensis, but the grey edges of the abdomen more
intense and skull with broader rostrum and nasals: anterior
zygomatic roots heavier; the zygomatic plate broader and more
convex, projecting further forwards; the plate laterally com-
pressed and more vertical, resulting in a narrower infraorbital
foramen.

Measurements.—See table, p. 75.

Specimens exramined.—Twenty-one from the type-locality.

38. EPIMYS GRISEIVENTER Bonhote.

Mus griseiventer Bonhote, Fasciculi Malayenses, Zoology,
Part 1, p. 30, pl. it. fig. 3, and pl. iv. fig. 5 (1903).

A single example only of a rat that appears referable to this
species was obtained on Koh Chang. As my camp was close to
a village, which is a port of call for steamers, 1t is quite possible
that the species has been introduced. The uniform upper surface,
smoky-grey underparts tinged with buff, dark feet, and black tail
distinguish it from forms of B. rattus.

Head and body 181 mm., tail 212, hind foot 34, ear 22.

39. Kreimys concotor Bonhote.

Mus concolor Bonhote, P. Z.S, 1900, p. 195; id., op. cit. 1902,
voli. p, 39; Flower, op, cit. 1900, p. 361 ; de Pousargues,
Mission Pavie, Indo-Chine, Etudes ieee ili. p. 528 (1904);
Gyldenstolpe, Arkiv for Zoologi, Stockholm, Band Sy Nowe pels
(1914).

Numerous specimens of this little rat were brought to me by
the children of Klong Yai for the sake of a cent. or two, but all
were immature, and I only preserved five examples.

40. EPIMYS BERDMOREI MAGNUS, Subsp. n. (Text-fig. 1.)

Type. Adult female (skin and skull), No. 1890/C.B.K.
B.M, No, 15.1].4,157, Collected at Klong Menao, 8.H. Siam,
12th January, 1915.

58 MR. C, BODEN KLOSS ON

Characters. —A large form of Mus berdmorei, with smaller ears
and tail considerably shorter than head and body, bicoloured with
dark tip. Pelage of two elements :—slender, very flexible spines,
with light bases and dark brown tips, and soft under-fur with
neutral grey bases and dirty-white or drab tips. Mamme
3-2 = 10,

Colour.—General colour of the upper pelage, which is harsh
but not stiff, clove-brown on the median dorsal area, lightening
to mouse-grey on the cheeks, sides, and limbs; everywhere
grizzled with the pale tips of the under-fur, and on the sides
by the exposed pale portions of the spines also. When dis-
turbed the neutral-grey basal colour contrasts sharply with the
browner external tone. In certain lights a brilliant green sheen
is visible from nape to rump. Under surface of body and lmbs
and the upper surface of the hands and feet white to the bases of

Text-figure 1,

Skull of Hpimys berdmorei magnus.

the hairs; the white area extends to the extremities and the
upper lip, but does not include the bases of the vibrissze. Ears
rounded and almost naked, a small white patch below the ear-
opening, ‘lail bicoloured, only the basal three-fourths white
beneath, the distal fourth entirely dark; somewhat thickly clad
with short hairs, black on the dark, white on the white area, but
no pencil ; in the centre eleven rings of scutes to the centimetre.

Skull and Teeth.—\ have been privileged to examine the skull
of Blyth’s Mus berdmorei, all that remains of the type which
came from Mergui, Tenasserim; it lacks the bulle and the
posterior half of the cranium, while the teeth are only just
beginning to show signs of wear. The Klong Menao individual,
while otherwise resembling it, is considerably larger with ap-
parently a relatively longer rostrum. The zygomatic plate is,
however, actually narrower, as is also the anterior root of the
zygoma, while the posterior root is more robust; the fronto-

MAMMALS FROM SIAM, 59

parietal suture is less curved and the incisors are paler, being
ivory-white with white tips, though they project in the notable
manner of the type of Wus ber dmorei: while the molars are of
similar small size, and the rostrum is likewise elongated, with a
straight or only slightly curved upper profile. No information
18 available as to the bulle of the latter, but those of H. b. magnus
are perhaps larger than are to be found in any eastern rat of equal
size, being extremely dilated and kidney-shaped. Tliomas, in the
account of the skulls seen by him (see below), does not mention
this very notable feature.

Measurements.—I give, in the form of a comparative table, the
measurements of the present animal, of the type of Mus berdmorei,
and such others as have been published of animals which have
been allocated to the species *.

Collector's External Measurements, in millimetres.

| | Siam. | Type. | Thagata. Bhamo, | Manipur.
a | a : aw easel
| Head and body ...............) 210 | cirea 155 170 142 174

| UE A eect op ee lean SP | reco. {G3 oeetae

| Hind foot Poe eno arnl 38°5 | 35 32 35 36
Dptiangris oes et eae | O85 | u 20 29

Skull Measurements.

Siam. Type. | Thagata, | Manipur. |
Gueatestilemothnpnnee trates met estate
Condy lo-basilar length SARC EEA Geena kedeae ene
asallpl enc Ghee ses ee PN myntn cee ec 8674.
Palatalileno thew ease a | oleae) aiemae=2 21:4 29:4,
Plane HORT, sop cnosesbescedoeessn Sno ceo Py ||. teil Al 73
Drasheria mes eaves t enc sac ctides eect, ee 101 | 140 129 141
Upper molar series ...............-00.00 52] GO. 4, . OG 6-0 61
Length of nasals.. Beate one are ile One 160 140 16:0
Anterior breadth of ‘nasals. Be dane weaaare Al 48 | 42
Interorbital breadth ...........0..00.......4) ok |) OS 68 7-0
Ly comanicnonrerdtihinsesaeeeieee yee eens 240 | 215 218 PAs
(Cireyaveyl loreal 56 cos aaoesscocegacnucos coesen| WL(AO) 165
Interpanretalabneardtiwyeesseee es sene le lacy 137 10°3
a length ........-... Sh | aes 4:0
Extreme breadth between outer ‘edge
of infraorbital foramina ..............| 12:1 | 11:0 10°4 |
Zygomatic plate .. skeeneceel 4 408 50 - 4:0 Aneel
Extreme breadth between ~ auditory
meati .... ein al S38
| |
Breadth of basioccipital ‘at suture ...... 3°5
|

* Thomas, P. Z.S. 1886, p. 62, two unsexed specimens from Manipur; id., Ann,
Mus. Civ. Genova, ser, 2@, vol, x, (xxx.), 1892, two temales from Thagata, Tenass
serim aud Bhamo, Burma.

600 MR. 0. BODEN KLOSS ON

Remarks.—The above measurements show the much greater
size of the eastern animal as compared with the western indi-
viduals, while the colour of the latter, given by Thomas as ‘‘ clear
slaty grey,” is also very different from the brownish tone of the
other.

The type of Lpimys berdmoret was described as being of about
a foot in length, of which the tail was not quite half; hind foot
13 inches. Fur shortish, even, coarse and hispid, but not spinous,
of one quality only. Incisors white. Tail rather more copiously
clad than usual with short hairs. The upper side, originally given
as grizzled-grey, unmixed with rufous, was later stated by Blyth
(op. crt. Xxxil. p. 343) to be dull brown, which is in close agree-
ment with the colour of the present animal.

The species is in no way related to H. ferreocanus Miller, of
Peninsular Siam.

41. AGANTHION KLOsst Thos.

A single porcupine of the bengalensis type was obtained on the
mainland at Klong Yai.

It is remarkable how little information we have concerning
Hystric bengalensis. There is Blyth’s original description *
founded, I am able to state (thanks to authorities of the
Indian Museum, Calcutta, who have lent me the type skull
for examination), on a half-grown individual with incomplete
dentition, supposed to come from the Sunderbunds. There is
Jerdon, in the ‘Mammals of India,’ who borrowed from Blyth,
and there is Anderson, who, in his ‘ Zoological Researches,’ when
treating of H. yunnanensis, gives (passim) a few fresh details,
while Blanford, the latest author to deal with the species, had
no material for examination when writing for the ‘ Fauna of
British India,’ and simply repeated Blyth’s original description.
Beyond this unsatisfactory literature no other details of topo-
types seem available, and I am forced to supplement it by
measurements ef a skull from the Karen Hills given by Thomas
in his paper on the Mammalia collected by Signor Fea in
Burma and Tenasserim f.

The present example, while generally agreeing externally with
descriptions of Acanthion bengalensis (Blyth), differs in the fol-
lowing respects :—The longest bristles of the crest are only 4 to
5 inches long, but are tipped with white for more than half their
length; the white demi-collar is ill-defined on the middle of the
throat; the quills are white with a dark band at their centres,
rather than white and black with a more or less defined white
tip, and the few long flexible quills are white throughout, lacking
any dark middle band. Blyth’s description, however, is hardly

* Journal Asiatic Soc. Bengal, vol. xx, p. 170 (1851).
+ Ann. Mus. Civ. Genova, ser. 2 a, x. (xxx.), p. 87 (1892).

-MAMMALS FROM SIAM. 61

up to modern requirements for subspecific purposes, and is at
best that of a young animal only.

Measurements of the skull are as follows; those in parentheses
being of the Karen Hill animal referred to above :—Basal length,
119 (119) mm.; greatest breadth, 75:5 (68); mesial nasal length,
71 (64); anterior nasal breadth, 27 (29); posterior nasal breadth,
36°5 (40); length of naso-premaxillary suture, 43 (39); length of
frontal suture, 31 (29); bregma to back of occipital crest, 42 (34) ;
diastema, 36 (38); upper molar series, 31 (27); distance between
outer corners of the two infraorbital foramina, 57 (53); height
of nasion from centre of palate, 51 (51).

Thus the Siamese-~-Cambodian skull, while of the same length,
is broader than the other ; but the nasals are narrower through-
out; though longer; the tooth-row is longer, as is that portion of
the skull posterior to the bregma.

Other measurements that may be recorded are :-— Median dorsal
length" of skull, 139 mm.; median nasal length, 71; median
frontal length, 31; median parietal length, 18. Collector's
external measurements :— Head and body, 835; tail, 115; hind
foot, 933 ear, 45°5.

Having regard to these differences, together with geographical
derivation, S.E. Siam being more than a thousand miles distant
from the Sunderbunds, it seems possible that the eastern animal
may eventually prove distinct; but until the mammal survey of
India, now much curtailed, has been actively resumed again, and ©
topotypes of bengalensis are available, nothing can be done. For
the present, therefore, I content myself with the above remarks.

Pousargues, in Mission Pavie, Indo-Chine, Etudes Diverses, iii.
p- 533 (1904), states that H. bengalensis does not extend east-
ward beyond Burma, but Gyldenstolpe (Arkiv for Zoologi,
Band 8, No. 23, p. 20) has since recorded, under this name, a
poreupine obtained by him in Northern Siam which appears to
be intermediate in cranial dimensions between Fea’s Tenasserim
specimen and the present animal.

(At Mr. Kloss’s request I have examined this skull, and it is
referred to in my paper on Acanthion klossi, Ann. Mag. N.H.
(8) xvii. p. 136, Jan. 1916.—O. Thomas. |

42. MuUNTIACUS MUNTJAK, subsp.

An immature male, with the posterior molars not fully up,
was obtained on Koh Chang Id.

It is a very brightly coloured animal. Dorsal region and
upper side of tail fulvous-chestnut becoming ochraceous on the
under surface, limbs, base of ears, and sides of head. Forehead
and front of pedicels tawny,top of muzzle brown. On the nape
and the front of the lower limbs there is a scattering of blackish-
brown hairs, which are in excess near the hoofs, particularly on
the hind feet. A black line along the horn pedicels and the

62 MR. C. BODEN KLOSS ON

facial rib. Chin and throat, inner side and posterior outer side
of ears, axillary region, lower abdomen, inner side of thighs,
under side of tail, back of lower fore limbs, and a small patch in
front of the digits of each foot, white. Head and body, 980 mm. ;
tail, 185; hind foot, 291; ear, 105; height at shoulder, 610.
Skull, greatest length, 203; greatest breadth, 81.

The horns, which are not yet differentiated from the pedicels,
are tipped with velvet, and the distance in a straight line from
the tips to the base of the pedicels on the inner side is 146 mm.

The Barking Deer of Siam was described by Gray (P.Z.5S.
1861, p. 139) from a skull with deformed antlers as Cervulus
curvostylis. Recently Mr. R. lydekker, whose death all
interested in game animals will much regret, has defined this
race in the ‘ British Museum Catalogue of Ungulates,’ vol. iv., as
being of medium size (wpper row of cheek-teeth 2,3, inches),
general colour orange-tawny, fading to buftish on neck and
underparts.

The present specimen is so young (milk premolars still in place)
that, lacking other material from Indo-China for comparison, I
have not applied any subspecific name for the present. The skull
is remarkable for the reduced size and marked definition of the
lachrymal pit, which is far smaller than any other which I have
had the opportunity of mspecting, the upper edge being very
sharp and the pit immediately within and above this notably
concave. ‘The vertical portion of the lachrymal is, further, much
reduced in height.

43. CERVUS UNICOLOR, subsp.

Rusa peronti Gray, P. Z.S. 1861, p. 138 (2).

Cervulus cambojensis Gray, loc. cit. stupra.

Cervus unicolor Flower, P. Z.S. 1900, p. 372.

Cervus aristotelis de Pousargues, Mission Pavie, Indo-Chine,
Etudes Diverses, iii. p. 536 (1904).

Cervus unicolor equinus Gyldenstolpe, Arkiv for Zoologi,
Band 8, No. 23, p. 30 (1914).

A form of Sambhar is represented in my collection by an
immature female from Klong Yai, S.E. Siam. Height at
shoulder, 760 mm.

The hairy frontlet and antlers of a deer collected by Mouhot
in Cambodia was described by Gray (P.Z.S. 1861, p. 138), who
then considered it to be a Muntjac, as Cervulus CamPajne ts, but
was later identified by him as Rucervus schomburgki! (Brit. Mus.
Cat. Ruminants, p. 76 (1872); Brit. Mus. Hand-list Ruminants,
p. 145 (1873)). Lydekker, however, regards this specimen as
belonging to Cervus unicolor (Brit: Mus, C Cat. Ungulates, vol. iv.
p. 79 (1915)), and if he is correct, and the Indo Chinese Sambhar
prove to be distinct, it will have to be known as C. w. cambajensis

Gray.

(or)
es)

MAMMALS FROM SIAM.

44, TRAGULUS KANCHIL AFFINIS Gray.

Tragulus affinis Gray, P.Z.S. 1861, p. 138.

Tragulus javanicus Flower, P. Z.8. 1900, p. 374.

Tragulus kanchil pierrei Bonhote, Ann. & Mag. Nat. Hist.
ser. 7, vol. xi. p. 293 (1903); lLydekker, Brit. Mus. Cat.
Ungulates, vol. iv. p. 291 (1915).

_ Lragulus kanchit de Pousargues, Mission Pavie, Indo-Chine,
Etudes Diverses, i. p. 5385 (1904).

Tragulus kanchil affinis Bonhote, P. Z.8. 1907, p. 11 ; Gylden-
stolpe, Arkiv for Zoologi, Band 8, No. 23, p. 29 (1914) ; Lydekker
(partim).

A single female, with very worn teeth, from Ok Yam, Franco-
Siamese Boundary:

This race is differentiated from that of the Malay Peninsula
by the absence of any blackish nape-stripe ; otherwise the colora-
tion of the two is similar. The bony orbit appears to be a trifle
larger:

Measurements.—Head and boty, 438 mm; tail, 70; hind
foot, 110; ear, 35:5. Skull: greatest length, 89°5; greatest
breadth, 43;

This form was first erected by Gray (P.Z.8. 1861, p. 338)
upon material consisting of seven specimens from Cambodia
collected by Mouhot. In the course of his description, he states
that ‘“‘a specimen of the species has been in the Museum as above
named for many years: it is said to have come from Singapore ;
but that probably was only the port of transit.” This remark
ean hardly. be regarded as the citation of the type, so that the
name affinis must be confined to the Indo-Chinese form, for it
was further a nomen nidwm until rendered available for use
through being applied with description to the Cambodian
animals specified therein. .

In 1903, Mt: Bonhote (Ann. & Mag. Nat. Hist.) took the view
that affinis should be referred to the Malayan ahimal with the
specimen of dubious provenance for type, and redescribed a
Cochin-Chinese example under the name 7. &. pierrei. Later,
however (P. Z.S. 1907, p. 11), he found reason to change his
opinion, and accepted the name of afinis for the Indo-Chinese
form.

Even the strict systematist, who holds that the first locality
cited is that of the type-specimen, must, I think, accept this
view, for the title of the paper in which this species is dealt
with is ‘ List of Mammals, ete:, collected in Cambodia” and,
unless otherwise specially excepted in the text, this locality has
priority.

In the Brit. Mus. Cat. Ungulates, vol. iv., Lydekker appears
to have overlooked the above facts. The unfortunate term
“Lower Siam,” applied by several describers of species to the
northern half of the Malay Peninsula, 7. e., Peninsular Siam, has
been the cause of much confusion to others who do not use their

64 MR. C. BODEN KLOSS ON

atlases sufficiently *. Thus, Siracha in S.E. Siam, about 40 miles
S.E. of Bangkok, is regarded by Lydekker as practically the type-
locality of 7. ravus Miller (= 7". k. affinis of Lydekker), which
came from Trang in Peninsular Siam, about 400 miles south of
Bangkok. The outcome is that 7. k. affinis is given a distribu-
tion from Pahang, Malay States, north to Moulmein in Tenas-
serim, and thence east to Annam, while 7’. k. pierrei Bonhote
(= 7. k. affinis Gray), which name is accepted by Lydekker, is
supposed to extend from Lower Cochin-China west to Siam,
thus making two subspecies of the same species exist side by
sile; whereas inter se ravus and affinis are two well-defined
forms, the one spreading from the middle of the Malay Peninsula
and the other from Cambodia, the line where they intergrade
being still not clearly known.

In similar fashion Lydekker extends 7. javanicus napu F. Cuv.,
of Sumatra, up the Malay Peninsula, from Selangor to Southern
Tenasserim, and at the same time places in an intermediate
position, Trang ranging southward, 7. j. canescens Miller.
While it is highly probable that the latter name will have to
be regarded as synonym of the other, yet while the subspecies
are excepted the allocation of specimens to them results in an
impossible distribution. ‘

45. Sus CRISTATUS, subsp.
Sus (Q)) Grays ozs: 186 iii E39;

Sus cristatus de Pousargues, Mission Pavie, Indo-Chine,
Etudes Diverses, iii. p. 535 (1904).

An immature female, with posterior molars not up and one
milk-incisor still present, was shot on Koh Chang Id.

The strongly-narked crest is tipped throughout with light
isabelline, and there is a patch of clearly defined white bristles
at the angles of the mouth, a number are scattered over the
lower abdomen and in fewer quantity on the throat and chest.
The prevailing colour is black, clear on the cheeks and shoulders,
fore and lower hind limbs; but the forehead, sides, and thighs
are annuiated with buffy and white. The greater part of the
inner surface of the ears is covered with white hairs, and the
fringe along the edge is very short.

* To avoid similar confusion in future, I suggest the use of the following divisions
for Siam :—

i.) Northern Siam: the mountainous country north of the Thoungyin River-
mouth and the great bend of the Mekawng (about Lat. 18° N.)

(ii.) Central Siam: the great plain, south of (i.), watered by the Menam and its
tributaries and by the Bangpakong and the lower Mekawng and Petcha-
buri Rivers.

(iii.) Western Siam: the hill country between the Menam plain and the
Tenasserim Boundary, south to about Lat. 12° N.

(iv.) Peninsular Siam: the Malay Peninsula south of (iii.) to the Protected
Malay States. ;

(v.) Eastern Siam : the “ Korat Plateau ” east of (1i.), drained by the tributaries
of the Mekawng.

(vi.) South-eastern Siam: the coastal country south of the Bangpakong basin
and the Battambong-Cambodian frontier, drained by streams running
into the Gulf.

(For fuller details see Journ. Nat. Hist. Soc. Siam, vol. i. part 4, 1915.)

MAMMALS FROM SIAM, 65

The skull is remarkable for the antero-posterior length of the
bulle, which in this dimension are larger than those of full-
grown animals from Peninsular Siam and about twice the length
of those of Sus jubatulus, a small race occurring on Terutau
Island, off the west coast of that region and occupying a position
with regard to it very similar to that of Koh Chang in respect of
S.E. Siam. It is possible therefore that, when better material is
available, this pig may prove to be a representative of a local race.

Though Blyth, in 1875 (Cat. Mamm. & Birds of Burma, p. 43),
drew attention to differences in the Tenasserim animals, the
common wild pigs throughout Eastern Asia were all regarded
as typical cristatus until Miller separated the Peninsular Siamese
and Tenasserim animal under the name of S. jwbatus (Proc. U.S.
Nat. Mus. xxx. p. 745, 1906), and it is this, or some allied form,
that occurs in Southern Indo-China.

Head and body, 1110 mm.; tail, 190; height at shoulder, 610.
Skull: greatest median length, 265; greatest breadth, 116;
antero-posterior length of bulle, 26.

46. ORCELLA BREVIROSTRIS (Owen). (Text-fig. 2.)

_ Orcella brevirosiris de Pousargues, Mission Pavie, Indo-Chine,
Etudes Diverses, ui. p. 546 (1904).

Text-figure 2.

Photograph of Porpoise (Orcella brevirostris) at Klong Yai, S.E. Siam.

A male example of this cetacean was brought to me by fisher-
men at Klong Yai on 6th December, 1914, but my preservatives
being then nearly exhausted I only kept the skull.

Except that it had the profile of the head considerably less

Proc. Zoo, Soc.—1916, No. Y. 4)

66 MR. C. BODEN KLOSS ON

swollen and convex, while the anterior edge of the pectoral fins

was more curved and a neck more evident, it closely resembled

the figure given by Anderson (Zool. Res. pl. xxv. fig. 4).

“ Bon slaty-leaden throughout; greatest length, 3660 mm.
ive):

Skull: greatest length (condylo-basal), 286 mm.; basal length,
260; palatal length, 141; greatest breadth, 202; rostral breadth,
77. Length of mandible, 225. The skull is asymmetrical, par-
ticularly in the region of the nasal openings; but asymmetry
seems to be the rule with this species.

R.16 1.15
Teeth, R12 L123: all are worn down to mere flattened stumps

and there are no signs of premaxillary teeth.

De Pousargues (loc. cit. swpra) records a specimen from the
Mekong River.

This species is very common along the Chantabun coast.

While sailing from Klong Yai to Klong Menao we saw, late
in the afternoon of December 7th, numbers of white cetaceans
between ourselves and the shore. They presented a most bril-
liant appearance with the low sun shining on them, but none
came sutticiently near for details to be observed. They were,
however, of large size, as big as the present species, and I imagine
them to have been examples of Sotalia sinensis Flower.

EXPLANATION OF THE PLATE.

Map of the coast and islands of South-East Siam, showing the places at which
collections were made by Mr. C. Boden Kloss.

67

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1 He

ON DEATHS IN THE SOCIETY'S GARDENS. (Cl

3. Report on the Deaths which occurred in the Zoological
Gardens during 1915, 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 January 17, 1916: Read February 8, 1916. ]

INDEX. Page
IPatholocygemrer sec aietc aaah ocee aoe eee Hid
IB OCIERVASIES 5 355550dco00000 sce uboaScocononansons | tele

On January Ist, 1915, there were 733 mammals, 2073 birds,
and 3/1 reptiles in the Zoological Gardens ; and during the year
280 mammals, 1167 birds, and 237 reptiles were admitted, making
a total for the year of 1013 mammals, 3240 birds, and 608 reptiles.

During 1915, 280 mammals, 706 birds, and 179 reptiles have
died: that is, a percentage of 27°6 for mammals, 21-7 for birds,
and 29°4 for reptiles.

Out of the total deaths for the year, 1165 in all, 378 occurred
in animals which had not been six months in the Gardens: that
is, about one-third of the total number. It has been found that
after six months’ residence in the Gardens the death-rate of the
animals falls rapidly ; so it is assumed that by this time the
new arrivals have got over their journeys, or have died from any
diseases they may have brought with them, or have got quite
used to their new environment. Of these 378 animals 90 were
mammals, 200 were birds, and 88 were reptiles; and if these
be deducted from their respective totals, the death-rate per-
centage will come out as 18°6 for mammals, 15°6 for birds, and
14-9 for reptiles.

The following Tables show in outline the facts which have been
ascertained. Table I. summarizes the actual causes of death in the
three groups specified. Under Reptiles are included Amphibia.

Taste I.—Analysis of the Causes of Death.

Reference
Diseases. |Mammals.| Birds. |Reptiles.| to Notes
| | tollowing.
1. Microbie or Parasitic |
Diseases. | L 1
AUBOEODIIOSIE 56 cocoa scaenononuo oad 6 60 2 2
(Miy.COSIS# Pharr tok Presse 208s 10 | Be 6 3
JPMGWOTAOINNA, G50 con cenacconeoav ena cndl 48 70 34 4
SEOUGESIIVR cons soo coc obo onvasoscnenal| 5 Vin 1 5
JANSCESS! MAS). PR a em uals hig itt
JRCRGNIS — aa sccocsocenoscaccsuseo ae il
JEROME Soocboosd ocd son 40a suc aoo eel 1 2 6
Eimapyentai) .eeeeenseteeter sock 5 |
Bronchiectasis) 3. ssses seeds eee 1 |
By ClIGIS: cates. ene cree eee Rasa ae 1

78

PROF, H. G. PLIMMER ON

Tasie I,—Analysis of the Causes of Death (continwed).

Diseases.

1. Microbie or
Diseases con
Cystitis zi

Coccidiosis
Saccharomycosis .

Syngamus .........

bo

Organs.

Br oncho- -pneumoni

Pericarditis
Degeneration of he:

Hepatitis
Cirrhosis ..

Tract.
Gastritis .

Gastro-enteritis
Enteritis :
Intestinal obstr uctic
Intussusception

6. Diseases of Urinary and
Generative Organs.

Nephritis
Salpingitis

Atrophy of uterus
7. Various. —

Adenoma ,
JNO. so p00 cos00ac

Osteomalacia
Injuries
. mortem

Besides those tab

Parasitic
Hemoer evarinosis

MOOPRISTNOSMS. 529 o66 non coe aoccee oon)

. Diseases of esporetons

INTE CHABSTISS oon oncces coanaosco cco cao0ce
TBI ROVOVGLEUTMIS): joo coacen ogds0040 o0dea0.005|
Congestion of lungs ..

3. Diseases of the Heart.
4. Diseases ee Liver.

Fatty degeneration RN Ra
5. Diseases of the Alimentary

Gastric ulcer: ation . ‘

Gycutimi ce Ie SATA

Retained mice =e ae Oe

@arcinomaerec cee eee eee
SHINGO paapdocne pas nannesbeaonedou co:

Chloroform poisoning

Mammals.| Birds.

pig 130
gi, . Bet iec 34,
ee muscle : a: il

HOW sco sgo noe nes

He et oo bd

discovered post - } a 9

Reptiles.

roo:

ul

WD a:

Reference
to Notes
following.

coffe 0)

10

bn

hia

13

14

15
16

ulated above,

38 mammals, 113 birds, 4 reptiles, were killed by order or
by companions,

1 Pe] 6 9

Silage

Die gs

4] 9

died from malnutrition
or starvation,

were too decomposed
for examination,

DEATHS IN THE SOCIETY'S GARDENS, 79

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 II. 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 I1.—Secondary Diseases found in the animals tabulated
in Table I.

Reference
Diseases. Mammals.| Birds. |Reptiles.| to Notes
following.
MuberculOsiss eee. emo | sf 13 1
Mycosis Hy Foe Banca | Br 2 oe
IPFAATTNOWEY \ cncitnanncos cus sonosdaenono nse 5 1
Rericanditisieeassctcn tenner. 5
Reritomitisiee acute mee aes ee ac 2 a |
VAIDS CESS Me auscueremerrene au ie il 1 |
IR vormlioe aia roaster scat s esa eases 2 Ase |
La GIGh HDI). ..SacGaecacbanadnancenaeepeesnctor 2 a
AOTC a odoguonueeede acandoceneee HDC resEEE 1 2 2
Miata aise ase esa enaeeearnaye tenner: il 8 UY
Nill euerasmee rey cer sues tnee Sean ta era ete 8 2 6 1 18
IWOECOOVWOADE, doo sdnccascbecnab6cnn000 008 ve 3 19
ISGHTONOIS! 6 ct earns aeGereaascoauenadnoscan on: 2
SHUDOUGEENTOM — nongopcocane soo soncdoucnce: 2
INGOTS) arses ceriadone oseeenaadceeprbeee 3 ast fe
Annce bic IMfEChlOM) ..osss ssa eee eee see sce ape Bo 4 20
IGEN TNO ETAS ANE og prenos wooeoboaron: oe sa 17 21
Brom chit symecscecsss deste o-eossese eee 8
Broncho-pneumonia .................. 8 He
Congestion of lungs .................. 23 157 5
CCIE), OP MEATS sos 065 con boo coo nee cod Gor it 134 9
Ely dno thoralies cesses cree eee 1 oe
Dilatedeiearby ats. eee cree eee 2 Tl
Atheroma SNe eRe tA a 5 4,
PAMVEUTI SIN vss ses ohasiaes Aacseseceeeeseees| 2 tee 22,
Pericarditismcsesnsrascec se seaceeeeoes as 11
Degeneration of heart-muscle ...... 4 8 be
Ey dropericandiuimal see. eee eee ease: ies 2 Be dela
1h (Gioia tnt igepeeneneaogedaeneeeneae nba ocones ie 6 18)
Fatty degeneration of liver ......... iN 45 12 23
(CITA STEN aaesasceneonedben cageascaycnoeee 5 1 ae
GasiritisWeea seacoast eetc 2 1 |
Gastriemlcenationmer eters eta: 14 Id cee 24
Gastrorentenitismeeerte et ee serene 5 1
IDWS) ayaaecete cacdenoy soudenconeenees 41 109 6
ImbussuscepulOnierers ss seeeesera eee: 2 ty
Atony of intestine ..................... 1 A 95
Nephritis : 43 81 4
Shore ha hoes eee eee ees 1 | 26
Brean ancya ene ce reece Se esoe | it [hs rene aaa |
SEN OPOESTNE) coocencosnos conpsnononnansnboel ee ie |
SANCOMLAS AN caek ooh eecee emer aan 1 se | | 27
PAGO s Son EPA NE icahoane torte SenCoen ancy 3 4 Days
Hemorrhage 1 |
SOUT OUBPAWERTIONE) ala coe sco vee sao obaohe 1 | |
RICK EtG i ee. at cestoe remanent Sec 17 Bestian | |
Osteomalaciie eee ee eeee|| ae J
Injuries 4 9 |

80 PROF. H. G. PLIMMER ON

Table III. shows, in still further detail, the distribution of
the immediate causes of death amongst the various orders of
mammals.

TasLE III.—The Distribution of Diseases causing Death
amongst the principal Orders of Mammals.

|

| a | rs
| = Si eel 2 Hest
nS S od a a || co
P Boe | ee oe 2 Pee) ioe
Diseases =e @ | %& = =n es
. ao | > | = — a3 | 5
| 6 ti eS) 5 = D
Wise ieee || ses Je @ SI
| BS | a |S = eS Ss
| Ay | O | Fx ~ St
|

|
|

ulberculosishe. etre cesar Peer aes lb aa
| JETEMUTMONNE cocgnodsosconsedoseossnopeposoonal 1G
IB;TADVOIEY. | Saoveaagecaasah ssoosmmoccoantancl| elle du. Wo}
Peritonitis Be anes Enea vedorel| ue :
Sep ties mite meee etree aaspsncosesoncee nace 2 me
VEAVCSVUTAISS /Pisedyet AA a teare Aare dy arg iat eceael A erase a) | ee
IBYROTRCIVIECUAISIS: Gosecagsseascoccodensooseel! son ae a 1
Wocerdiosise pees ssen sso men sates ds. 5 oe. ilar 2)

GS UIU Siege eu meter abode te here MecteeslMmiee. ibe ole

Gs Gs ty
eo:

p13

wv
—
OU tt

IBronchiti Swen ees ccbe rece e aes eters
Broncho-pneumonia <0...) |.
| Congestion of lungs

| Atelectasis ORAS O 8 oe eke
(CHER NOEIS OE IWOP sooccsscesanncnacebocees|| cco ||
GRaIS HUGS piercer anc meron cette mecchseere eens |
Gastric ulceration 22)..-..........-.-5. 0.5]
(GAGHROSEMMUENGIS cos noes ceoonocoocnoeaeooe
WARMED GT Sies carton mrece Gactiascesse aceel|

Or g1@
(ou
2) Cos:
[a0 Cll a)
S)

$b Os
pave

ES

IOMALSTWRGADINOM asosscccscnoosocsecenecgees|| 2 ne
_ Intestinal obstruction .................. 1 rae
| LINDE yeoedss eecentanceacomeenceatcescolh 24 2 8 6 3

here bobo Oe:
Or
lon
Lo

CHSGU Suis aoereme dues deabetcentepeodeaes caer lal ros aaa 05
INTRO ONE OE WHATS 5 coosoocgs cos ccoss ace! sue i iiss 1
vetanmed@placental sssss-eeeeeeseeeeeeee 1 ms ae

ALCO MMA dasecracncaecensne te mnaeeeeetecemaeah meer ace 1 2
(CEURGTIAVONSTEY, enue she ce no peRaenea caceecn cutee! | ake 1 ae 1
UNCLE), Conerbous neue cantenmeen cccnetesaacaly et sje
PACT Od OrMaeetrrenigtiaee Becca eck ani cceeceae teen rntl Mame i

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

DEATHS IN THE SOCIETY S GARDENS. 81

TaBLe IV.
Mammals.

Mears it eetenace 1912. - 1913. 1914. 1915.

Total number of mammals.... 1391 1272 1261 1013
Muberculosise ..cesaensceeeees ae 14 31 12 6
IMiyCOSis) 2. qcectostuscne See 12 8 6 | 10
JEVEBISTONTUEY, co sognaseasnde oes acs noose50e0 45 34 53 48
TEROTAAEUATS). “Sundcéado sboseooieace ceuscnose 12 6 li 10
Broncho-pneumonia ............------ 30 25 24 34
Congestion of lungs ...............++- 14 14 14 13
Gastro-enteniiisme een eee eee: 11 7 16 11
Batenrtistie sce cence ener cee 38 33 33 25
Nephinitisp aise taeees tes teas: 89 90 66 40

Birds
Total number of birds ......... 3526 3518 3610 3240
(Rulberculosismmenes enue eee eee 79 104 113 60
IVI. COSISWMees ena desNte Run NEL eA S 72 5 88 32
JEMGUUACNE) Soeococosgsoessandapierococuaa|! AS 89 118 70
Coneestionyots Innes) esses peeeeeees| OS 98 133 130
IVINGERIEI Seen Naas cei an oe tenderer ae 154 148 169 150
INepliritisi sys, Ge wee fer ates ge Ba ky 104 135 129 81
Reptiles

Total number of reptiles ...... 1166 1169 1045 608
Muberculosts=ecsseseee eee see 11 6 4 2
My COSTS denne “eres prse cone eaisscie ars reece 2 i 10 6
JEFOITDORNEY, “age dadeseeahooods Sod cad ane 560 124 138 69 34
Congestion of lungs .................. 22 13 19 11
ISVOVE STATIS. A SseaemSeRERAEECer eeu auakicn 25 15 17 8

Notes on the foregoing Tables.

1. The total incidence of microbic and parasitic diseases in
the Gardens for 1915 is 7°8 per cent. for mammals, 5-3 per cent.
for birds, and 7°8 per cent. for reptiles. Practically this per-
centage is caused by the deaths of animals which had not been
six months in the Gardens; if these be excluded, the percentage
is under 1 per cent. for mammals and birds, and just over 1 per
cent. for reptiles.

2. The deaths from tuberculosis have never been so few since
accurate records have been kept as during this year. The
percentage of animals dying from tubercle is °5 for mammals,
1:8 for birds, and °-3 fer reptiles. Three ungulates died from
tubercle of human type, probably caused by spitting, which is
one of the vices of visitors to the Gardens. A Cercopitheque

Proc. Zoou. Soc.—1916, No. VI. 6

82 PROF. H. G. PLIMMER ON

and a Baboon were both pet animals, but neither of them was
allowed out of quarantine, which is the best way of keeping fresh
infections out of the Gardens. The birds show a remarkable
reduction of deaths from tubercle of nearly 50 per cent. ; and, if
the overcrowding of many of the bird-houses could be done away
with, there does not seem any reason why there should not be a
still further improvement. The greater number of the cases
have been amongst the Gallinaceous birds, but this year a
Flamingo succumbed to tubercle, for the first time since records
have been kept. In only 19 of the birds were the lesions gene-
ralized, which is a much smaller percentage than usual, and
indicates a less severe type of the disease. The 2 cases in the
Reptile-houses make the smallest number since records were
kept: there seems no reason why it should not be completely
driven out. One of these cases was in a Crocodile, in which it
was general; this is the first recorded case in this particular
animal.

3. All the mould diseases are grouped under Mycosis. 6 of
the mammals were Kangaroos and the disease was of the ordinary
type; it occurred in a Deer and a Squirrel, for the first time,
forming tumours in both—the moulds in each of these two cases
were of a species new to the Gardens. There is a very great
reduction in the number of cases occurring in birds, of considerably
over 50 per cent. A Duck 14 days old was filled with growth
of the organism, and, in a Pheasant, mycetomata (mycotic
tumours) occurred in the lungs, which is very unusual in birds.
Amongst the reptiles a Ceratophrys and an Anaconda died from
mycetomata, and a Ceratodus, which had been many years in the
Gardens, died from a mould disease of the skin, similar to that
from which several of the larger toads and frogs have died.

4. There is a more than relative decrease in the number of
cases of pneumonia, especially among the birds and reptiles. In
the mammals it seems to be particularly associated with pyorrheea
and rickets. In two of the reptiles it was due to worm eggs
embryos, in the rest it was pneumococcal.

5. The septicemias were due to dirty wounds in 8 cases, to
pneumococci in 2 cases, and to an abscess in the antrum in
1 case.

6. Due to worm-cysts in a Squirrel, and in both of the birds
due to injury, from, in the one case a nail, and in the other a
thorn.

7. This occurred in a Deer, associated with putrid bronchitis,
and was of the sacculated variety. It is the only case I have
seen In an animal,

8. This was an acute hemorrhagic cystitis ina Wolf, due to
infection with Bacillus coli.

9. In a Cobra in which over 60 per cent. of the erythrocytes
were infected.

10. Due to a rare parasite whose position is still uncertain,
and found here in a reptile for the first time.

and

DEATHS IN THE SOCIETY'S GARDENS. 83

11. There has been relatively a slight general increase in these
diseases of the respiratory organs. ‘They are, of course, largely
dependent upon weather.

12. There has been a slight decrease in the inflammatory affec-
tions of the alimentary tract. In 3 of the mammals and 50 of
the birds it was hemorrhagic, and in | of the reptiles it was
caused by worms. The remainder of the cases were apparently
due to the quantity or quality of the food not being suitable to
the animal.

13. There has been relatively a considerable decrease in the
number of cases of nephritis in the mammals and birds. 24 of
the cases in mammals and 7 of those in birds were acute. ‘The
remainder were chronic cases, of varying degrees, a number of
which were associated with other old-age changes. About 100
of the deaths have been due to old age, or to the ‘artificial old age
induced by captivity.

14. In a Kiang which died after delivery; the wall of the
uterus measured only one-sixteenth of an inch in thickness.

15. One of these cases was in a Wolf whose son died of the
same disease, in the same position, in July 1914.

16. Two of these cases were in Deer, one in the nose and the
other in the liver and intestine ; the third was a lymphosarcoma
of the mediastinal glands in a Hamster.

17. Under the term malaria are grouped 1 case due to Plas-
modium kochi, 7 cases due to Hamoproteus danilewskyi, and 1 due
to Plasmodium preecox: see also section on blood-parasites below.

18, 19, 21. See sections on blood-parasites below.

20. The blood of 3 Viperine Snakes contained a number of
large amcebee, probably from the intestine; they were also found
in numbers in the liver of a Mocassin Snake (comparable with
the flagellated organisms found in the blood of reptiles and
described in my Reports of 1912 and 1913, and in a paper on
Blood-parasites found in the Gardens during the years 1908—
1911: vide P. Z.S8. 1912, pp. 235, 406; 1913, p. 141; and 1914,

aS):

22. Of the aorta in a Seal and a oath

23. The large number of cases of fatty degeneration and
infiltration of the liver is probably due to too rich food and
insufficient exercise. The greater number of the birds are the
small, highly coloured birds from the Small Bird-house, whose
food-capacity is enormous.

24. By far the greater number of cases of gastric ulceration
occur in the Primates, generally in connection with diseases of
the kidneys or lungs.

25. In an Ibex with very considerable feecal accumulation and
symptoms of autointoxication.

26. Both kidneys of a Wallaby were filled with small stones :
none in the bladder.

27. This was a small sarcomatous growth in the kidnev of a
Coypu Rat, probably arising from an adrenal inclusion.
6? pS

84 PROF. H. G. PLIMMER ON

BuLoop-PARASITES.

_ During the year the blood of every animal that died has been
examined, with the result that parasites have been found in 46
cases, in 24 species for the first time.

They have been distributed as follows :—

Filarie. In 2 mammals; in | species for the first time.
In 6 birds; in 4 species for the first time.
In 1 reptile for the first time.
Trypanosomes. In 2 reptiles.
( Plasmodium kocht. In 1 mammal for the first
| time.
: Hemoproteus danilewskyi. In 7 birds; in 3 species
MN a Kp ‘ for the first ie
| Plasmodium precox. In 1 bird for the first
time.
Leucocytozoa. In 3 birds; in all for the first time.
Toxoplasma. In 1 reptile for the first time.
Hemogregarines. In 17 reptiles; in 7 species for the first
time.
Intestinal organisms. In 5 reptiles; in 2 species for the first
time,

The following list gives particulars of the blood-parasites in
detail :—
Embryo Filarie found in the blood of Mammals.

Hapitat, TYPE.
Lion Marmoset (Leontocebus rosalia) ... Brazil. Very long.

Found in the following for the first time:
Woolly Monkey (Lagothria infumata)... S. America. Long.

Embryo Filarice found in the blood of Birds.

Chilian Starling (Cureus aterrimus) ... Chili.

Found in the following for the first time :

White-eyebrowed Wood-Swallow (Arta- N.S. Wales. Long, thick.
mus superciliosus).
3 Red Birds of Paradise (Paradisea Waigiou Island. Short, stout; 2 of the

rubra). birds contained also
filarie of long thin
type.
Military Starling (Trupialis militaris) . Chili. Long, thin.

Embryo Filarie found in the blood of a Reptile : for
the first time.

Wolf-Snake (Coluber letus) ............... N. America, Long, thin.

Trypanosomes found in the blood of Reptiles.

2 Edible Frogs (Rana esculenta) ......... Europe. T. rotatorium type.

DEATHS IN THE SOCIETY’S GARDENS. 85

Plasmodium kochi found in the blood of the following Mammal

for the first time.

Mangabey (Cercocebus €thiopicus) ......ccrccrcncec cee eee tee ees vee

HAsrrat.
S. Nigeria.

Hemoproteus danilewskyi found in the blood of Birds.

Indian Dial Bird (Copsychus saularis) 20
Black-throated Lorikeet (Trichoglossus nigragenis)
2 Brown-necked Parrots (Peocephalus fuscicollis) .

Found in the following for the first time :

Yellow-fronted Barbet (Cyanops flavifrons) .....6..0...00. 00000
Larger Hill-Mynah (Gracula intermedia) ......0..000c00 cee cee eee

Red Bird of Paradise (Paradisea rubra)

India.
Tasmania.
Gambia.

Ceylon.
N. India.
Waigiou Island.

Plasmodium precox found in the blood of the following Bird

for the first time.

American Robin (Lurdus migratorius) .........s0c00e ese eee vee nes

N. America.

Leucocytozoa found in the blood of the following Birds

jor the first time.

White-eyebrowed Wood-Swallow (Artamus superciliosus) ...

Nonpareil Finch (Cyanospiza oid

NES INNO (CAGED CHEAPAD)) 630000000 sna05s son oshonbbeortnnaconaenpoic

N.S. Wales.
N. America.
S. Africa.

Toxoplasma found in the blood of the following Reptile

Sor the first time.

Say’s Snake (Coluber melanolewcus) ...... 20. .0ecencee eee cee cescevees

Mexico.

Hemogreyarines found in the blood of Reptiles.

Indian Python (Python molurus) ..

2 Indian Cobras (Naia MITES)

Hog-nosed Snake (Heterodon ‘aloban ee)
Blood-stained Terrapin (Cinosternum come)
fMsculapian Snake (Coluber re ae
Common Boa (Boa constrictor)........
Banded-tailed Tree-Snake (Leptophis 1 i iocer crews)
Anaconda (Hunectes marinus) . ane
Cooke’s Tree-Boa (Corallus aiseD)

Found in the following for the first time :
HaBirat.
Wolf Snake (Coluber letus) ...

Pennsylvanian Mud-Terrapin (Cinoster- N. America.

num pennsylvanicum).
Black-collared Cobra (Naia nigricollis). W. Africa.
2 West African Sand-Snakes (Psammo- W. Africa.
phis elegans).
West African Trionyx (Trionyatriunguis) W. Africa.
Rufescent Snake (Leptodira hotambeia). W. Africa.

Shielded Eryx (Hrya thebaicus) ......... Gold Coast.

N. America.

India.
India.

N. America.
N. America.
Europe.

S. America.
S. America.
S. America.
Trinidad.

TYPE.
Cells enlarged and de-
heemoglobinised.
Stout.

Long.

Cells enlarged and de-
hemoglobinised.

Stout.

Very small, like Lan-
kestrella.

Long, irregular: cells
dehzmoglobinised.

86 ON DEATHS IN THE. SOCIETY’S GARDENS.

Intestinal Organisms found in the blood of Reptiles.

HAsirat. TYPE.
Carolina Box-Tortoise (Cistudo carolina). N. America. Hexamitus.

Found in the following for the first time:

3 Viperine Snakes (TLvropidonotus Europe. Ameebz.
viperinus). :
Mocassin Snake (Tropidonotus fasciatus) N. America. Ameebee.

Addendum to Report on the Deaths for 1914, published
in Proc. Zool. Soc., March 1915.

Owing to the wrong figures having been sent to me from the
Office, the figures in paragraphs 1 and 2 of the 1914 Report,
p- 123, and in paragraphs 1 and 2 of the Notes, p. 126, are
incorrect: these paragraphs should read as follows :—

Paragraphs 1| and 2, p. 123.

On January Ist, 1914, there were 788 mammals, 2436 birds,
and 575 reptiles in the Zoological Gardens ; and during the year
373 mammals, 1174 birds, and 470 reptiles were admitted, making
a total for the year of 1261 mammals, 3610 birds, and 1045
reptiles.

During 1914, 309 mammals, 867 birds, and 301 reptiles have
died: that is, a percentage of 24-6 for mammals, 24 for birds,
and 28°8 for reptiles... ..).- . Of these 719 animals, 141 were
mammals, 375 were birds, and 203 were reptiles; and if these
be deducted from their respective totals, the death-rate percentage
will come out as 13°3 for mammals, 13°6 for birds, and 9°3 for
reptiles.

Paragraphs 1 and 2 of Notes on the foregoing Tables, p. 126,
will read as follows :—

1. The total incidence of infectious diseases in the Gardens is
about 7:5 per cent. for mammals, 9 per cent. for birds, and 8-1
per cent. for reptiles.

2. The following are the percentages of deaths from tubercle
during the year: mammals ‘9 per cent., birds 3-1 per cent., and
reptiles ‘38 per cent. on the total numbers for the year, etc., etc.

CO

I

ON AN ABNORMAL FROG.

4. A Frog with symmetrically Abnormal Hind Feet. By
R. W. Haromp Row, B.Sc., F.L.S., F.Z.S., Assistant
Lecturer and Demonstrator in Zoology, University of
London, King’s College.

[Received and Read November 23, 1915. ]
(Text-figure 1.)

Among the specimens used for the teaching of elementary
zoology at this College, there was recently found an example of
the Common Frog (Rana temporaria) in which both the hind
feet showed a curious and interesting variation. A photograph
of the external appearance of the two hind feet is given
(text-fig. 1, A), which clearly shows that on neither foot do five
functional digits exist; though in both cases a small calear, or
pre-hallux, as it is often called, is present in its normal position
in addition to four well-developed and normal toes.

The fact that certain of the toes always present definite
characteristics which enable them to be indubitably identified,
quite apart from their actual numerical position in the series,
renders it quite easy to determine the identity of the digits still
present in the four-toed specimen. Of these characteristics the
most obvious distinguishes the fourth toe, which is not only
longer than any of, the others, but also always possesses four
phalanges, which is one more than the number present in any
other digit. In the specimen under discussion four phalanges
can be distinguished on one toe on each hind foot, so that this toe *
is thus marked out as the fourth of the original series, and from
this it can be immediately determined that the missing digit is in
each case the first. This identification of the digits is shown on
the photograph. .

In view of the possibility, however unlikely, that symmetrical
mutilation of both hind feet might have occurred, one foot, the
left, has been completely dissected, and the dissection has shown
that the abnormality cannot be due to injury, for the first digit is
unrepresented by any tissue whatever.

During the dissection great care was exercised to determine
whether the muscles of the hallux were represented, and no trace
whatever of them was found, either in the form of small muscle-
masses or of fragments of muscular tissue or tendons, such as
would inevitably be left had the toe been bitten off or otherwise
amputated. The musculature of digit II (and of all the other
digits) was perfectly normal, and showed no signs of the inclusion
of muscles really belonging to digit I. Specially important is
the fact that from the aponewrosis plantaris only four tendines
superficiales arise, the one normally associated with the first digit
being totally absent. There is also no trace of any of the bones

88 MR. I. W. HAROLD ROW ON

of the hallux in the skeleton of the foot, nor any gap between
the calear and digit IL from which the hallux might have been
removed, and these facts taken together do away completely
with any possibility that the abnormality is due to injury. A
photograph (text-fig. 1, B) is given of the skeleton.

All the normal distal tarsal bones are present in the foot in
their normal position, though, unfortunately, they are not distin-
guishable in the figure ; the four digits present are also perfectly
normal in their anatomy.

Text-figure 1.

A. Photograph of the external appearance of the two hind feet, natural size.

B. Photograph of the skeleton of the left hind foot, natural size.

It was decided not to dissect the right foot, but as careful an
examination as possible was made from the outside, and revealed
no indication of the hallux in that foot either.

One question I cannot definitely settle. It is just possible
that the structure identified by me as the calcar is really the
remnant of the hallux, in which case the calcar is entirely absent
in both feet. I do not think that this can be the case, however,
for the following reasons. The position of this structure in the
abnormal specimen precisely corresponds to that of the calcar in
the normal frog, both as regards the foot generally and in its

AN ABNORMAL FROG. 89

relation to the distal tarsal bones: its size is just that of the
normal calear; and, in addition, in dissecting the foot I found
that its muscles corresponded exactly with the account given in
Gaupp’s Ecker-Wiedersheim’s ‘Anatomie des Frosches’ for the
musculature of the calcar.

One of the most interesting points in connexion with this
abnormality is the fact that in the manus of the frog there are
always only four digits, and the one usually accounted absent is
again the first of the series, though Emery (Anat. Anz. Bd. v.
1890, pp. 283-288; and elsewhere) has claimed that the missing
finger is really the fifth. But whether the absence of the first
digit from the hind feet of the abnormal specimen can be regarded
as additional evidence tending to disprove Emery’s theory, or not,
I cannot say. Another point of theoretical interest lies in the
presence of the calcar despite the absence of the hallux, which
seems to me to afford considerable support to the view that
the former does not belong in any way to the digital series.
Theoretical considerations based upon a single specimen are far
too doubtful, however, to render it worth while attempting to
discuss these questions.

Among the great number of structural abnormalities which
have been described for frogs of various species, a number of cases
of polymely and polydactyly occur ; but apparently this specimen
is the first in which a variation of this kind has been recorded.

i <
Fieve Wepre f

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Ns Tf, Ay!

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10

Horace Knight del.et lith. West, Newman chr

iS) OUMEAIL INE SIND) IE J PD) O12 TEs RUA.

PZ slole 2 OU Ewe Nees

Horace Knight del.et lith. West, Newman chr.

SOMALI IEVAINID) (BE) @ beh age

ON MOTHS FROM SOMALILAND. 91

5. On a Collection of Moths made in Somaliland by
Mr. W. Feather. By Professor E. B. Potton, M.A.,
F.R.S., F.Z.8. With Descriptions of New Species,
by Sir) Ga Hagkiawrsons | Barta, Da Baperoum Jive
Durrant, and Dr. Kart Jorpay.

| Received November 23, 1915: Read February 8, 1916.]

(Plates L& TILE)

INDEX.

GEOGRAPHICAL: Page
Localities at which collections were made ......... 92

SYSTEMATIC :
Odonionetiane ciate ese tee eae see eae ee enn Oss
(CPSSHMPAQTAS, FSD NS ces csoncsecocco sdoossooabeseeacannes » LOZ
“A CRO RESIS CMaaliol Ere neee eet een eat ea eee ere eee LO)
LEAT OUOTIOUES, AND, We socoogespaopdbscupescusossotascen 124!
IN FICODAROSGAIS., ROMS 186) pads ood soacdoeconspeeneonoondce LUN
ISASITOGHIGERE, (HENS TS, Gaqadocheochbdsssoconorsceanepnbocss LB?
J AGIAOCL ils (REDS ston nab os008 cob cenecobee coneegeenben anaes: Lly/
(COM GATOSOOCTHUG, ENS Wy cosedscusncenaceconsebcecooones — URE)
INOULONUG aa Clan Mowe dcctnecereemanie ea nteroesee Aee ae eee LAS
UNO OULU FASS 118 BO PR emia one octet saree any, 11743)

109 species and 2 subspecies are also described as new.

The moths described in the following pages were very carefully
collected by Mr. Walter Feather, who preserved excellent data.
Sir George Hampson has described the new species of all
the groups except the Sphingide, by Dr. Karl Jordan, the
Geometride, by Mr. L. B. Prout, and the Tineina, by Mr. J. H.
Durrant. The order of the species is from the more specialised to
the more generalised except in the Geometride, which Mr. Prout
has arranged in the opposite sequence.

Types of the species described by Sir George Hampson and
Mr. Durrant are in the collection of the British Museum, co-
types, when the series permits, in the Hope Department, Oxford
University Museum, and Mr. Feather’s collection. Dr. Jordan’s
and Mr. Prout’s types are in the Hope Collection, co-types,
when the series permits, in the British Museum, Mr. Feather’s
collection, and that of the describer. This statement renders
unnecessary any further reference to the disposition of types and
co-types in the descriptive part of this memoir.

* For explanation of the Plates, see pp. 181-2.

92 PROF. E. B. POULTON ON

The numbers prefixed to the names of species in the first part
of the memoir are those of Hampson’s ‘ Catalogue of Lepidoptera
Phalene in the British Museum,’ the position of new species
being indicated by letters added to the numbers of the most
closely allied species in the Catalogue. Thus species 2094a would
immediately follow 2094 in the Arctianz of the Catalogue.

The specimens were collected at the following localities, of
which the descriptions have been kindly furnished by Mr.
Feather :-—

MANDERA.

Forty-seven miles south-west of Berbera, alt. 3000 ft.

Rocks, grey and red granite.

Open and bush country, bush being mainly made up of thorny
trees, nearly all flat-topped Acacias or Mimosa. Big areas
covered with fibre-plants (Sanseveria ehrenbergi).

Gan Lippan.

(The ‘“ Lion’s Paw.”)

In the Golis Mts., seven miles east of Mandera, alt. 5800 ft.
Rock, limestone (Jurassic).

Good patches of big trees, mainly Juniper; also EKuphorbias.
Good grass land with scarcely any thorn-bush.

BuaGan.
Ten miles south of Mandera, alt. 3500 ft.
Rock, granite.
Thorn country, fairly open: few big wild Fig-trees.

HARGAISA.
One hundred and twenty miles south-west of Berbera, alt.
4000 ft.
Bush and trees same as Mandera, but a little denser. Bigger
areas of Sanseveria ehrenbergt.

BERBERA.

All the moths I captured at Berbera were taken in the
Bungalow at light. This was close to the sea; the maritime
plain is very thinly covered with a low-growing thorn-
bush.

Rock, coralline limestone.

DURBAR.
Nine miles south of Berbera, alt. 400 ft.
Rock, limestone.
Open land, few Palms with some rough grass.
Durbar is really the waterworks for Berbera, and at one time
there was a poor sort of Government garden there.

MOTHS FROM SOMALILAND. 93

With few exceptions the specimens were captured at Mandera,
and this place is to be understood when no locality is mentioned
in the text. Mandera is, however, always quoted for those com-
paratively few species which were also taken elsewhere. ‘The
specimens from Hargaisa dated Oct. 1908 were taken during
a period of about two or three weeks by Captain Jorkinson.

Mr, Feather writes concerning the method of capture :—
““T may say that all the moths were collected at light. I only
remember taking one species—I think a Geometer—otherwise,
and that I got in a porcupine-burrow along with a Skipper. I
sugared many times, but the only insects that came-were ants,
and they completely covered the sugar.”

Mr. L. B. Prout, in the introduction to the Geometridee
(p- 142), draws attention to the remarkable preponderance of
females; and the same unusual condition is to be found again
and again throughout the rest of the collection. Observations
made Sept. 12-15, 1915, in Bombay Harbour, on the return
from the visit of the British Association to Australia, have
led me to believe that nocturnal flights of female Lepidoptera
tend to occur during wet weather. The Bombay species in-
cluded the females of certain butterflies which flew at night
and came to light with the moths. The fact seems to be very
interesting and well worthy of a separate communication dealing
with these Somaliland moths and my own experience in Bombay.
In order to test the relationship of female preponderance to
wet weather, I have asked Mr. Feather to supply a record
of the rainfall and temperature at Mandera. He kindly replied
as follows :—

““T am enclosing the record of rainfall for Mandera, and
have used much the same words as I wrote in my diary. My
impressions of the rainfall were guided by what I had been used
to in England, as this was my first visit to Africa.

“IT cannot give you the amount of rain, as we had no rain-
gauge. I should say the annual rainfall at Mandera. is about
10-12 inches. Wet nights are a great rarity, the rain oftenest
coming in short heavy showers in the morning or afternoon.

‘The river I mention is, of course, a dry river-bed, except just
after rain. The longest time that I remember water running
down the channel was for about 8 hours. The river-bed is
about 120 yards wide, and in one part was a very small stream
of permanent water, which appeared between some rocks and
ran on the surface for 20-30 yards before disappearing in the
sand.”

94

PROF. E. B. POULTON ON

Recorp of Rain at Mandera, Somaliland, from Nov. 14th, 1907,
to June 23rd, 1909, and of Temperatures, taken in the shade,
from March 14th, 1908, to June 30th, 1909.

Morning and afternoon temperature :
Date. (Fahrenheit). Rainfall.
Noy. 14, 1907. en Rain in morning.

—= Ih, — eee Rain in afternoon.
Jan. 22, 1908 sunuilcedWiagleoaeaeise gee Fine drizzle all day.
Mar. 14, — 6.30 a.m. 65 deg. 1.30 p.m. 102 deg.

= ie =={_ B= 8 = ie =

= ig = | © = B= = = lie =

Si] |) = B= 10 ] =

== 13, = = = B= 1a —= Oe =

= i = | 6 = B= = = its

— 20, — 6.30 — 68 WSO = 100) ==

= Sie = = (= a — 109 = :

ooo) =SB= 2305 05 =

Sosa 0

= oh S|) 60 =] Gs osy = oe
April 5 = = =

=eoh SS es 1Ps0 SS oh S

Pees Ge ee a ae

ae Ae 2 a Go tes

a = OO 10 — 90 —

—- 88 — == 2 =

— 9 — — — 68 —

—= 10, = — — 68 — 10 — 78 — | Dull day: a few drops of rain.

= iil, = — — 68 — ==> & —

—= 1 = — — 64 — — — 84 —

5 ae =—- = © =

See — = Mas OOS AS

— 16, — — — 59 — 130 — 85 —

aR, = SES ile a hy ae 3

a= iy, = 20 — 96 —

= i — — — 66 — — — 99% —

= i = = = ii = == =

= 20, — — — 80 — — — 86 — | Slight shower in morning.

— 2, — 5.30 — 75 — 10 — 93 — | Heavy shower.

22, 71 La) 8B =

= B = — — 69 — 2.0 — 79 — | Heavy rain for 1% hrs.

— 24, — GO = 7 = 1.30 — 81 — | Rain for 2 hrs.

BEER OG Nk ee oe a Oo

= AW = — — 1723 — = = B=

eS he. HS = 8). Oreos eee

= 8, = a ee

— 99, — | 6380 — 77 —- — — 9% —

— 30, — GO = Wf = ==> & =
May 1, — — — 75 84

= = — =— 71 — — — 90 —

ese == &8=

= A 30 —= 73 = 12.30 — 93 — | Slight shower.

= & = GQ —= Ws = 2.0 — 95 — | Slight shower.

— 6 — — — 7 — — — 93 — |A short shower a little before
sunset. During the shower
hailstones { in. to #in. in

—- %- = = f= = = 8 — diameter fell.

a eaaee See es == =

=O) — — 10 —

= il = | = 75 = =

MOTHS FROM SOMALILAND

Morning and afternoon temperature

Rainfall.

Dee, (Fahrenheit).

May 12,1908.) 5.30 a.m. 73 deg. 1.30 P.M. 94 deg.
— 2B, — — — fA — —- — 94 —
— 4 — 6.0 — 7 — 20 — 94 —
— 16, — — — 7% — — — 3 —
— 16, — — — 7% — — — 8&4 —
— 17, — — = 0 — 130— 69 —
— 18, — — — 66 — — — 84 —
— 19, 65 — — 84 —
— 20, — — — 7% — — — gf —
— 21, — — — 7 — —- — gg —
— 2, — — — 68 — —- — 9 —
— 2, — — — 68 — — — 94 —
— a4 — — — 7% — —- — 94 —

25, 74 10 — 94 —
— 26, 73 —- — 94 —
— 27, — — — Ff — 1.30 — 93 —
28, 65 — 971 —
29, 77 —- — . 91 —
30, 79 — —- — 93 —
— 81 — — — % — 20 — 9 —
June 1, — — 76 — 9% —
2, 76 130— 94 —
3 78 94 —
— A ai — — 92 —
= & = — — 7 — — — 992—
—- 6 — — — 7 — 20 — 88 —
—- => = 74 91 —
— 8, 76 93 —
—- 9%, — 75 96 —
— 10, — SS = ff = —- — $$ —
11, 75 93 —
— 19, 74. —- — 92—
— 123, — = 73 93 —
— 165, — ee = = 9 —
= 1, 75 94, —
NY, i = = = '|S—s—*~—4, —
— 18, — = = = —- — 74 —
— 19*, — — — 7% — —- — 94.—
29, = — — 102 —
— 30, — = 73 101 —
July 1, — 79 — 102 —
2, 75 103 —
3) 78 98
4, 78 — 101 —
— 56 — — — 100 —
— 6, 5 100 —
7; 77 100 —
= 77 — — 103 —

Short shower.
Heavy shower in afternoon
for 15 hrs.; hailstones.

Shght shower.
Fanly heavy shower. Tempe-
rature fell considerably.

A good heavy shower, starting
at 3 P.M. and lasting till
5 P.M.; water coming down
the river in fair quantity.

Rain-storms all round, but
none at Mandera.

Water came down river last
night. Evidently heavy
rain in the hills, though
none at Mandera.

Heavy shower; water coming
down river.

* Here I have a note that I took the thermometer from the inside of the messhouse

and hung it on the veranda (well in the shade).
rose to 103 deg.

The temperature then immediately

PROF. E. B. POULTON ON

Morning and afternoon temperature

Date. (Fahrenheit). Rainfall.
July 9,1908.| 6.04.m.78deg. 1.30 p.m. 101 deg
10. = Sys 20 — tole
— es se 1G = = & io
ieee = im
— 13, — — — 1% — — — 102 —
es = Ss = 2 WA
= 16, = = = = = 103 =
— 16, — — — 123 — — — 102 —
= i = SS ie = = >=
— i138, — = = (= — — 102 —
— i, = — — 17 —
— 20, — — — WT— — — 99 —
= Fh, = = = = = 98 —
— 22 — == = == =
— 23, — = as (= BO = B=
— 245 = = 3 = 20 — 98 —
—= 2, — = = 3 = — — 98 —
— 26, — — — 100 —
= We = — — 73 — — — 9%
— 2, — — — 1% — — 998 —
— 29, — — 70 — == B=
— 30, — —= — 0) — — — 93 — | Had two rather dull days.
— 31, — — =< Sh — — — 97 — | A good shower this evening.
Awe, i, == = = = — — + 92 —
— B= = = 5 = = = yf =
—- 3 -— = S| B= — 9 —
= d= = = ff = = = =
— 5 —- — — 7% — = = it =
= Choc — = is = 2.30 — 101 —
fy ie = = = = => WO =
— fe = Oa 20 — 92 —
—=— th == Ss = B= 2.30 — 93 —
—. 10, — aes Gas OOD = Of —
— u,—}| —-- 71- = — 92 —
— 12, — = = 6 DEO — WT) —
= 1 = — — 7% — 2.0 — 998 —
ei — — 7% — 230— 99 —
SS == 76) — — == itl —
eG AO —= il =
ay ay — — 9 — 2.0 — 101 —
— 18, — = = a= = = 1 =
— 19, — ech: Wee, bo = = iol =
— O = | @) = © = 03 ==
ae eo 0h = = iil =
— TH = — — 79 — = = idl —
— PB sooudooee aqnsees A shower this evening.
— 24 — — — 7 — 2.30 — 98 —
= = ee = = 1D =
— 3 = — — 7 — = = 108 =
— 27, — = 78 — 2.0 — 100 —
Boss = | 630 = 77) 2801 100
= = ee 5 eo On oS
[ — 31, — | Berbera.—Temp. in Bungalow, 116 deg. |
Sept. 35 = 6.0 a.m. 76 deg O p.m. 98 deg.
= 4 = Ges Ws BeOS Be ==
= §& = | Oo Se =— = a)
=) 62S —- — 23 — — — 9% —
Bi == Sh ie ee
— 8 = 6.30 — 77 — — — 97
SG = | BO> ye 2) = 9 =
ai TIO) mes ee i AA) — 0 ==

MOTHS FROM SOMALILAND.

97

Morning and afternoon temperature “ip
Date. (useeennea) Raintall.
|
Sept. 11,1908. | 6.30 a.m. 75 deg. 2.30 p.m. 102 deg.
— 12, — 3.0 — 104 —
— iby 7.0 79 101
= ik = C0) —= 75) == a3) — > 8B) ==
Seis. 2 | 7.0% == ee 230K) A014
Be. == ||) 6:30 78 56) ==
i, — Bee Mer ly ae) ee
= ls — — — 7% — 2.30 — 96 —
iG) EEG) a0 i) a ey
— 20, — = = 72) — — — 9 — | Rain around, but only a few |
drops here: Mandera often
= Ml = = = — == © = seems to miss the rain.
2 2 a Gn) 2s80e——. 499)
SON yee Oh SE. QO ng
aoe i — — 74 — 2.00 — 96 — | A few drops of rain at noon.
— 2%, — = = 5 = — — 9 — |A good heavy shower, lasting
about 20 mins.
— 2, — 5.30 — 73 — 2.30 — 94 —
oy, BO. ae 1a => Of, 5
= SyRORe = ey ay LS Sa eee
SE O84 = Sh = Soe 5a
sei OS cat ie 1S — — 90 — |A very slight shower at night. |
A good shower lasting |
15 mins. this day. The last |
Oct. 1, — —= — (4°— — — 990 — few days have been cloudy
—- 2%.— = = (2 = 20) —= 8 = in early morning and _ late |
= 3 = = = i = 11,310) —= 2) == afternoon.
—- 4 — — — 7 — 20 — 92 —
— 5 — = = = 1.30— 90 —
— 6, — — — 72 — — — 90 — | Heavy shower for 10 mins.
— 7, — — 69 — — — 88 — | Shght shower in afternoon.
— = —| —~ 68 — 2:0) — 88 ——
= = = = B= = =— 0 =
ae 1 =< |) 630) —— eG en)
= i, = 7 CO = by = Seay)
= 1 = (lls —= (i — 1.30 — 94 —
— 123, — GO = 7 = 2.0 — 80 — | A few drops of rain.
= id = = = & = 1.30 — 86 — | Water down river in quantity,
but no rain at Mandera.
— 1, — — — 63 — — — 87 —
or TG Be Vez ote Foto 1 Wie ree
= if = BETES © (0 ee eee AUS) em eye
— 18, — — — 65 — — — 87 —
— i, = = = OS]. 8 = =
— 20, — — — 6 — — — 8 —
— O23 | ORD = Fe ee 5
=o 998 — Gl = ==) ES
ee | GO Go aie eae
sO, aes KS) es ie OS
= 8 = Ss= f= 8 = —
= 2 = = = @ = — — 78 — | A few drops of rain at night.
2 = = = — — 77 — | Slight shower in morning.
— 9, — | — — 68 — a
= WW = — — 58 — = = 87 —
— 30, — — — 61 — —- — 8 —
— 31 — — — 60 — —- — 8s —
Nove al, — eS Gi) ate) eee
—- 24— — — 68 — — — 80 — |A good shower.
—- 3—-— — — 66 — 1.30 — 76 —
Si = 68 oe Oy
[— 9, — | Gan Libbah.—Few drops of rain. |
Proc. Zoou, Soc.—1916, No. VII. i

98

PROF, E. B. POULTON ON

Morning and afternoon temperature

| Rainfall.

Dinike. (Fahrenheit).
(Noy. 10, 1908. Gan Libbah.—Rain for 2 hrs. |
— we — 6.0 a.m. 62 deg.
i SG 80 nim Was
ae =: DA a yey |
== 15, 59 20 — 80 — |
— 16, — 59 80
— 1% — — — 56 — 1.30 — 82 —
— 18, 60 80
— 19, — — — 59 — = = =
— 20, 59 == == fi) —
— 41, 57
ee — — 63 —
— 23, — — — 56 —
— 24, — = = & — 1.30 — 8 —
— 2, — — — 650 — 3.30 — 82 —
— 26, — — — i) —— 2.30 — 738 —
— 27, 48 2.30 — T4 —
= Heh 65
[— 29, — Berbera. | A few drops. |
Dec. 21, — — — §1 — 20 — F2 — |
— 25, 55 30 — 70 —
— 296, 53 5 |
— 27, — 6.30 — 48 — 20 — % — |
— 28, — 60 — 51 — 10 — 7 —
— 29, 5 LB — Ww =
oe 0: 53 (ie)
real 54 i)
Jan. 1, 1909 52 72, — |
— 2, 51 73 |
— 83, 49 72
— A, 53 70 |
= & = a |
== Hil, 60 72 — | Fairly heavy dews for the last |
== 12, 55 U8) oe [4 weeks.
— 413, 50 BO == Uy = -
550M 53 ei
Sa) eee une nee en s0e WSO
ace 60 |
— VW— = — 58 20 — 83 — |
Se 60 130 — 1 80;
== I — 60 80 —
== AY, 61 Sil ==
RO: = See eae eee |
oe 60 130 = 7 —
— 23, 63 130 — 73 — | A few drops.
— 24, 60 20 = We = |
== 28; 60 130)—— Sa
== 0, 60 Wa
— 27%, 58 76
== Be, 53 20 — WM —
—= By — — — 60 — LOM OF
— 30, 50 4.30 — 7 —
— 831, - 51 divorce
Wad, I, 52 — — 80 —
eo 55 a) =
ALS: 55 a |
Saris 60 ri |
a 60 = iy = |
= & = 61 76 — |
SAG 60 ri) a Ge =|
= 8&8 5 = => i} |

MOTHS FROM SOMALILAND. 99

|
| Morning and afternoon temperature | :
Date. (ubreaher): p Rainfall.
9, 1909. 6.0 A.M. 55 deg. 2.0P.mM. 73deg.
10, eS 56 (a) |
ante 58 1QO—= 85 = |
12, 57 = = 6 = |
13, — — — 58 — —- — Y9—
Ta ee Oe ee
ia Sf gs to] pS |
HGue Ss 10) == i ee |
17, — 54 85 —
18, — — — 59 — 1.30 — 81 — }
TG), ee = — FH = On |
20, 60 Sl |
he i SS Gil = DO = Gf = |
22, 61 — — 8 —
GB er eG) ee 120 = BR =
24, — — — 56 —
05. == ae NS = = =
9650 = 60 = m=
21 = == Bf — — — 83 —
Oe oe 57 81
i, = = = & = 1) — (O83 = |
— == i) = 1S) = Gi =
Sy 56 WO = iS =
4, — — 58 79 — i
B, 59 130— 81 —
ee 59 80
Us 61 WO = Fo —
8, | 61
9, 58 1.30 — 80 — |
10, — — — 60 78 |
11, 59 79 —
12, 58 = =
is = == 58 82 —
14, 57 81 |
15, 58 “hi |
1G, == ee aye 85
17, 53 20 — 87 —
18, 55 : 85
1, = SS Ne
20, — 58 I =
Oil, == 5G = ARO) =e sie
22, — — — 62 — = = % =
DE S53 8S FSS wy =
4, — — 63 — 130) — 88) ——
25, = se oO) = 90)
26, — — — B—- — — %—
a — Snes — — 75 — | Heavy rain.
23, — — — 68 — —= = 0) =
29, — => => (3 = — — 87 — | Good shower in afternoon.
30, — =] 0] 9] )—] fH —
31) — — 69 — = «=> '— '_ SB =
April 1, — — =< @) = 1.30 — 85 — | Fewdrops. Much rain in hills.
= of val — 37 — | Steady rain for 2 hours. |
3 — = =— = — — 82 — | No rain, but river came down
in strength. |
4 — — 7 — — — 72 — | Good shower.
5, — — — 71 — — — 82 — | Good shower.
6, — 70 Se) ==
ei _- 70 — 93 —
8, — —- — 1— — — 95 — | Rained in afternoon. Heavy
storms all round.

=e

100 PROF, E, B. POULTON ON
Morning and afternoon temperature :
Desi (Fahrenheit). Rainfall.
April 9, 1909. 6.0 a.m. 70 deg. 1.30P.m. 67 deg
— 10, — — — 68 — — — 82 —
= lh = — — 6 — Shower in early morning.
— 12; 68 — — 8 —
— 18, 72 —. 9 —
7 ba == — — 23 — — — 97 — | Shower in afternoon.
— 6b, — = 73 — 87 —
== AG = — — 7 — — — 85 — | Few short showers in night. |:
Little water coming down
river at daylight.
= My == = = & = — — 82 — | Very heavy shower: one could
only see for few yards. River
== Bey == = = — 1@ == & = in strong flood.
— 19, — — — 68 85
— 20, — — — 68 — —- — 9% —
— 2, — — — 5 — — — «s ——s"——?23) ——
= By = — §-—_— «13 — —. 85 — | Heavy rain-storm a few miles
BESO ies a 2 — — 82 — away.
— 4, — — — 67 — —- —. 7 —
—> 255, —— — — 67 — = = 8 =
— = 67 87
— 24%, — — — 1 — — — gf —
Tr Oh a 68 = 92 — | Few drops.
— il), 70 —- — 8 —
May i, — — 64 88
acne 70 — — 80 — | Slight shower.
—- 38 — — — 66 — — — 88 —
= 4, — — — 68 — — — 92 — |
pa AD a aso) — — 78 — | Rather dull. Raining all round,|
SG 72 82 [but not here.)
=a Uo 71 92 Slight shower.
== 72 90 Little rain in afternoon.
a) Byes = = (= — — 91 — | Shower in afternoon.
= 10, = = = 7 90 Little rain.
ety 73 95
ioe EG OR BES Cys
GE ie eo ee ee
— 4 — = == il = =. —. 92 —
— + tb, — — — 72 — Se
40 — 104 —
= 1G = — — 72 — 10) 95)
— W7%,— — — 73 - 92
= Ie == = — = — — 94 —
== is, = —= — = = = 8 =
= By = — — 72 — = = i =
= Bll = — — 7 — —- —. 92 —
— 2, — = — fl = —- — 98 —
Beo3e BENE oye) Aa ee OS |
— 94, — — — 7 — — — g9 — | Heavy rain for about 40 mins. |,
= BD = — — 69 — — —, 82 — | Fewdrops about 5 p.m. Much |
rain in immediate neigh- |,
— 26, — — — 67 — 20 — 90 — bourhood.
= Bi, = —- — 1 — 1.0 —. 89 —
— 2, — — — 71 — =. —. 90 —
= DM = — — 69 — — —-— +«§8 — .
Le s0) AO = 6 =
— $l, — —- — i — 0) == fet) ==
June 1, — — — 71 — —- — 90 —
— 2 — — 7 — — — 90 —
= 35° = = = Yt) = = = Se =
Riss STAG 2 SE Ba Ee Ds

|

MOTHS FROM SOMALILAND, © 101

Morning and afternoon temperature | aes
Date. = (Fahrenheit). Rainfall.
June 5, 1909. 6.0 A.M. 68 deg. 10pm. 90 deg.
= Ges Ss MS LS] B=
es So 5 GQ SS
Bane egs 1s eS 8 ae ie
—- 9% — — — 68 — — — 92 —
PEON e— BS (63) ays Teeter ee
I, ee eg i at uk OB =
= 1 = eB 94.
— 123, — — — 3 — — — 98 —
Sid Bee ae eo 05
eae ee ee) 3)
— 16 — — — 71 —
= We SE EE Sa ee eee
1 eR) ee Ee
== 16 eS ERS ate heme Goa
220s Se e903
ay | Ole ers yh ye 90. =
a3 YD, ee eo
Cue) Pili |
— 2, — —- — 1 — 10 — 95 — | Heavy shower about 6 P.M.
a) a ap es
5 ee |
2G. lie ae OO) |
= We = ee oo
=, (DB. = Bee iW eo) Sey Coe eee
a OT ee ASG ee | 0
a) Ee ae Ff ee ee eS

* | have a note here that the temperature usually rose to about 100 deg. about 3 p.m.

“ For a further period of a little over seven months there was
no rainfall at Mandera. This statement is from memory, but I
am confident of its accuracy, and am very sorry I cannot find my
diary to provide confirmation.” -

WALTER FEATHER.

HETEROCERA.

Fam. AMATID 4.

273. APISA CANESCENS Wlk.

Mandera.—1908: June 18,—1 ¢; Sept. 15,—1 ¢. 1909
Jan. 8,—1 ¢; Jan. 12,—1 ¢d; Jan. 16,—1 GC.
Gan Libbah.—1908: June 24,—1 dG.

283. MeErarctTiA BURRA Schaus.
1909: Apr. 20,—1 ¢.

In this and all succeeding species where no locality is men-
tioned, Mandera is to be understood.

102 PROF, E. B, POULTON ON

Fam. ARCTIAD 2A.

Subfam. Noiina.

63c, NOLA CHIONEA Hmpsn.
1908: Mar. 22,—1 @.

Subfam. LirHosiAn2&.

843. SrccrA SORDIDA Butl.
1908: Oct. 25,—1 °@.

Subfam. ARCTIANA.

1677. MAENAS ARBORIFERA Butl.

1908: Apr. 30,—1¢; Oct. 18,—1 9. 1909: Mar. 28,—1 3;
Apr. 8,—1 6; Apr. 11,—1 &6; Oct. 14,—1 ¢. 1910: Mar. 6,
—2 6; Mar. 12,—1 ¢; Mar. 14,—3 ¢.

1730 a. DiAcrIsIA DIVERSATA Hmpsn.
1909: Sept.—1 9.

Dracrisia var. near 1812. tinzata, W1Ik.
1909: May 10,—1 ¢.

1858 6. HEsTIGMENE GRISEATA, sp.n. (PI. I. fig. 1, 2.)

Q. Head and thorax brownish grey, the back of head and
tips of tegule orange-yellow, the patagia with small black spots
near base; palpi black at tips; abdomen fulvous orange with
lateral series of small black spots. Fore wing brownish grey; a
small black spot at base of cell; black points in the angles of
cell and two beyond lower angle. Hind wing white tinged with
reddish brown. Underside brownish white, the costal area of
both wings tinged with red-brown; hind wing with black
discoidal spot.

1909: May 21,—1 9 (type). Hap. 40 millim.

2068. TERACOTONA SUBMACULA WIlk.
1909: Oct. 22.—1 ¢.

2088. UTETHEISA PULCHELLA L.
1909: May 11,—1 9; May 21,—1 5. 1910: Jan.—1 ¢.

2094 a. SECUSIO SOMALIENSIS, sp. n. (PI. I. fig. 2, 9.)

@. Head and thorax pale reddish brown tinged with grey ;
the vertex of head with minute black streak; the tegule,
shoulders, and patagia near base and tips with black spots ringed
with whitish; the metathorax with minute black spot; palpi
brown at sides; pectus and legs whitish tinged with brown, the

NG)

MOTHS FROM SOMALILAND. 103

former with black spot at side; abdomen brownish ochreous
with dorsal and sublateral series of black spots, the ventral
surface whitish tinged with brown. Fore wing pale reddish
brown; a subbasal black point on costa ringed with white ;
obliquely placed antemedial black spots on and below costa and
in cell and spots nearer the base below median nervure and
above vein 1, all ringed with white; two diffused waved white
medial lines, rather oblique to below the cell, then incurved ;
obliquely placed postmedial black spots ringed with white below
veins 8 and 7, then a series of diffused white spots with minute
black points on the spots below veins 5 and 4; a subterminal
series of diffused white spots in the interspaces. Hind wing
pale grey-brown. Underside of both wings uniform pale grey-
brown.

1908: Nov. 13,

1 Q (type). Hap. 36 millim.

2098. Sucusto srricgATA WIk.

Mandera.—1908: Sept. 25,—1 9.

Gan Libbah.—1908: June 25,—1 6; Nov. 6,—1 2. 1909:
Nov. 4,—2 Q.

Fam. AGARISTID 4,

84. Rorura aisHa Kirby,
1909: Apr. 8,—1 ¢.

122. AicocurA BREvivirraA Hmpsn.

1909: May 6,—1 9; May 10-9 @. 1 2 specimen without
data.

162. TuERTA TRIMENT Feld.
1909: Apr. 5,—1 ¢; Apr. 14,—1 3, 1 Q; Apr. 20 or 21,—
WG,

Fam. Nocrurpa,
Subfam. AGROTIN&.

47 a, CHLORIDEA ALBIVENATA, sp. n. (PI. I. fig. 3, 2.)

Q. Head and thorax rufous mixed with ochreous ; antenne
brownish, white towards base; palpi, pectus, legs, and abdomen
ochreous irrorated with brown, the dorsum of abdomen thickly
irrorated. Fore wing ochreous tinged with rufous and slightly
irrorated with blackish, a stronger rufous shade along median
nervure expanding towards the postmedial line; a diffused blackish
streak below base of cell; a faint diffused oblique blackish ante-
medial Jine from costa to median nervure; reniform a diffused
blackish spot ; the veins beyond the cell slightly streaked with
white to the postmedial line, which is whitish slightly defined
on each side by blackish, bent outwards below costa, then
minutely dentate, excurved to vein 5, then oblique, a fuscous

104 PROF, E, B. POULTON ON

and rufous shade beyond it; a terminal series of black points;
cilia whitish tinged with brown. Hind wing ochreous suffused
with brown, the terminal area broadly suffused with blackish ;
a large blackish discoidal spot ; cilia white, tinged with brown at
base. Underside ochreous, the costal areas irrorated with brown;
fore wing with some fuscous along median nervure; both wings
with large black discoidal spots and black subterminal shade from
below costa to above inner margin.

1909: Oct. 20,—1 2 (type). Hap. 24 millim.

56. CHLORIDEA OBSOLETA Fabr.
1909: Mar. 2,—1 9.

304. EuxoA SPINIFERA Hibn.
1908: Nov. 20,—1 @.

Subfam. HADENINA.

1799. Dramas EUMELA Stoll.
1909: Feb. 28,—1 9; Apr. 8,—1 ¢; Apr. 14,—1 9.

1850. CirPHIS LOREYI Dup.
1909: Jan. 11,—1 @.

Subfam. AGRONYCTIN A.

3139. PrRIGEA CAPENSIS Guen.
1908: Nov. 24,—1 9°.

$552. TAMBIODES INCERTA Rothsch.
1908: June 7,—1-2 (in B.M.).

3623 a. THALATHA MELANOSTROTA, sp.n. (PI. I. fig. 4, 3.)

o. Head and thorax white irrorated with black scales, the
latter strongly tinged with rufous except the tegule; antenn
fulvous; palpi white, reddish brown above; pectus white; legs
white and brown; abdomen red-brown mixed with some white
and irrorated with black, the basal crest rufous, the anal tuft
and ventral surface white. Fore wing grey, tinged with red-
brown except on terminal area and irrorated with large black
scales ; faint traces of a medial line, oblique towards costa, then
sinuous; an indistinct double dark postmedial line, very oblique
towards costa, then sinuous and incurved below vein 3; a series
of black points before termen. Hind wing white tinged with
red-brown, the costal area and termen more strongly tinged ;
cilia white. Underside of fore wing suffused with brown ;
hind wing white, the costal area and termen to vein 2 irrorated
with brown.

1909: Apr. 8,—1 ¢ (type). Hp. 26 millim.

MOTHS FROM SOMALILAND. 105

3786. CEroLA PULCHRA B.-Baker.
1909: Apr. 6,—1 9; Apr. 9,—1 9; Apr. 14,—1 ¢.
3792 a. Maropo HETEROCHROA, sp. n. (PI. I. fig. 5, d.)

Antenne of male bipectinate with rather long branches to
apex, of female ciliated.

3. Head and tegule ochreous white, the latter with slight
brown lines at middle and tips; thorax bluish white slightly
mixed with pale brown; palpi with the 2nd joint, except at tip,
and the 3rd joint brown; frons with lateral brown bars; pectus,
legs, and abdomen creamy white, the fore tibie and the tarsi
banded with blackish. Fore wing bluish white tinged in parts
with brown, especially on costal and terminal areas, the veins of
terminal half with slight dark streaks ; a subbasal brown point
below costa ; antemedial line slight, dark brown, angled outwards
below costa and strongly in submedian fold and above inner
margin; claviform defined by dark brown, minute; reniform faint,
yellowish with slight brown centre; postmedial line slight, dark
brown, defined on outer side by yellowish except towards costa,
strongly bent outwards below costa, then waved, incurved below
vein 4, and with a slight brown shade before it towards inner
margin, some white points beyond it on eosta, and slight black-
brown streaks above and below vein 6 and between veins 4 and 2;
cilia intersected by slight white streaks. Hind wing pure white,
the terminal area slightly tinged with brown. Underside white.

2. More strongly tinged with reddish brown ; fore wing with
round whitish orbicular stigma and some fiery red on outer edge
of reniform and on the yellowish beyond the postmedial line ;
hind wing suffused with reddish brown ; underside tinged with
red-brown.

1908: Oct. 13,—1 9 (type); Nov. 24,—1 ¢ (type). 1909:
Mar. 12,—1 9; Apr. 14,—1 9; Apr. 20,—1 9; Apr. 22,—1 9;
Apr. 26,—1 9; Sept. 30,—1 2; Oct. 22,—1 9; Nov.6—1 Q.
Hep. 32-36 millim.

3878. LAPHYGMA ExIGUA Hiubn.
1909: Jan. 15,—1 5,1 @.

Genus QDONTORETHA, nov.

Type, O. featheri.

Proboscis fully developed ; palpi porrect, short, slender ; frons
with large, conical, truncate prominence with raised edges pro-
duced to two minute teeth below and two at each side; eyes large,
round; antennz of male almost simple; thorax clothed almost
entirely with scales, the metathorax with depressed crest; build
slender; tibie slightly fringed with hair; abdomen clothed with
rather rough hair, but without crests. Fore wing long and very
narrow ; the apex rectangular, the termen evenly curved and
not crenulate; veins 3, 4 stalked; 5 from just above angle ;

106 PROF. E. B. POULTON ON

6 from well below upper angle; 7, 8, 9, 10 stalked; 11 from ceil.
Hind wing with the cell long; veins 3, 4 stalked; 5 obsolescent
from just below middle of discocellulars ; 6, 7 shortly stalked ;
8 anastomosing with the cell near base only.

In key differs from Prometopus in the frontal prominence being
toothed at edges and the fore wing having veins 3, 4 stalked.

3880 a. ODONTORETHA FEATHERI, sp.n. (PI. I. fig. 7, 3.)

3d. Head white; antenne tinged with fuscous; frons with
black bars at sides; palpi mostly black; thorax and abdomen
grey-white mixed with some blackish ; pectus, legs, and ventral
surface of abdomen white, the tarsi black ringed with white.
Fore wing grey-white, the terminal half with black scales mixed
except a patch in and just beyond the cell from costa to vein 2;
the darker area defined on inner side by a faint oblique medial
line angled outwards just below the cell, with a black streak in
the cell from it to the pale patch, which is somewhat constricted
at discal fold. Hind wing white; a brown discoidal striga and
some faint strive on termen except towards tornus; the underside
with some black on costa towards base, a rather diffused black
mark on vein 8 just beyond the cell, and the costal area slightly
irrorated with black.towards apex.

1909: Mar. 12,—1 ¢ (type). Hap. 24 millim.

3989. ATHETIS LEUCONEPHRA Hmpsn.
1908: Sept. 24,—1 9; Sept. 27,—1 9; Oct. 13,—1 Q.

3998 a. ATHETIS DISCOPUNCTA, sp.n. (Pl. I. fig. 8, 9.)

Q@. Head and thorax creamy white irrorated with rufous and
a few black-brown scales ; antennae brown except at base ; palpi
tinged with red-brown towards tips; abdomen whitish suffused
with red-brown. Fore wing white irrorated with pale red-brown
and a few black-brown scales; small subbasal, antemedial, and
postmedial black spots on costa; a black point just beyond the
cell; traces of a postmedial line formed by red-brown and black
scales arising from the costal spot, excurved from below costa to
vein 4, then incurved; some minute blackish streaks on post-
medial part of costa; subterminal line represented by slight
blackish streaks and spots except towards costa; the terminal
area tinged with rufous except at apex; a series of small black
spots just before termen ; cilia rufous at base, chequered rufous
and white at tips. Hind wing white, the termen tinged with
rufous except towards apex. Underside white, the costal and
terminal areas of fore wing and apex of hind wing irrorated with
rufous.

1909: Sept. 11,—1 2 (type). Hap. 28 millim.
4020 a. ATHETIS ECTOMELENA, sp.n. (PI. I. fig. 9, 3.)

3. Head and thorax ochreous; antenne brownish: palpi
blackish at sides; tibiae irrorated with blackish, the tarsi blackish

MOTHS FROM SOMALILAND. 107

with pale rings; abdomen ochreous white with diffused fuscous
dorsal bands. Fore wing ochreous ; a minute black subbasal spot
on costa and slight point below the cell; a small black ante-
medial spot on costa, and traces of a sinuous line with slight
black marks on it below the cell and above inner margin; two
small black spots at middle of costa; a black subterminal band,
broad at costa and narrowing to a point at inner margin, ex-
tending, except towards apex and tornus, to beyond the slight
pale subterminal line, which is slightly angled outwards at vein 7
and excurved at middle; the termen ochreous with a series of
minute black lunules; cilia whitish, tinged with brown at base.
Hind wing white, with a slight brown terminal line except
towards tornus; cilia ochreous at base, white at tips, and with a
brown line through them towards apex. Underside white, the
fore wing and costa of hind wing tinged with ochreous; fore
wing with the terminal area suffused with fuscous except towards
tornus ; the cilia ochreous at base followed by a brown shade and
the tips white; hind wing with some brown on apical part of
termen.

1908: Oct. 20,—1 ¢ (type). Haxp. 30 millim.

Genus CoNSTANTIODES, nov.

Type, C. pyralina.

Proboscis absent; palpi upturned, the 2nd joint reaching to
vertex of head, slenderly scaled, the 3rd moderate, thickly sealed ;
frons smooth, with ridge of hair above; eyes large, round ;
antenne of male bipectinate with moderate branches, the apex
ciliated ; thorax clothed almost entirely with scales, the meta-
thorax with depressed crest; tibie slightly fringed with hair ;
abdomen with dorsal crest at base only. Fore wing narrow, the
apex rectangular, the termen evenly curved, crenulate; veins
3 and 5 from near angle of cell; 6 from upper angle; 9 from
10 anastomosing with 8 to form a narrow areole; 11 from cell.
Hind wing with veins 3, 4 from angle of cell; 5 obsolescent
from below middle of. discocellulars; 6, 7 from upper angle;
8 anastomosing with the cell near base only.

In key differs from Plusilla in the fore wing being narrow
with the termen crenulate.

4030 a. CONSTANTIODES PYRALINA, sp. n. (PI. I. fig. 35, ¢.)

3 2. Head and thorax white mixed with some red-brown ;
palpi with some dark brown towards extremity of 2nd joint ; abdo-
men creamy white, dorsally tinged with brown. Fore wing creamy
white tinged in parts with brown and slightly irrorated with
black, the termen yellowish tinged with rufous ; a slight curved
blackish subbasal line from costa to vein 4; antemedial line
reddish brown defined on inner side by white, oblique to sub-
median fold, then almost obsolete; some white in end of cell ;
reniform slightly defined by red-brown, large, somewhat angled

108 PROF. E. B. POULTON ON

inwards on median nervure, a red-brown shade beyond it from
costa beyond the postmedial line followed by some white; post-
medial line blackish, oblique towards costa, then slightly waved,
at vein 3 retracted to inner edge of reniform, then oblique to
inner margin, the veins beyond it with slight black streaks except
towards costa; some oblique white and dark striz on costa
towards apex ; subterminal line white, slightly waved from below
costa to vein 4, then oblique; some rufous at apex; a waved
blackish terminal line. Hind wing creamy white; a slight waved
brown terminal line; the underside with the apical area irrorated
with a few red-brown scales.

1908: June 1,—1 9 (in B.M.); Sept. 21,—1 ¢ (type). 1909:
Mar. 11,—1 2; Apr. 7,—1 g. Eap. 22 millim.

4103a@. Evuiopica 1GNECoLoRA, sp.n. (PI. I. fig. 10, 2.)

Antenne of female bipectinate.

@. Head and thorax fiery rufous; antenne black; pectus
and legs rufous; tarsi dark brown ringed with white; abdomen
ochreous brown, the ventral surface whitish tinged with rufous.
Fore wing fiery rufous; traces of .a curved deeper red ante-
medial line; a whitish point in middle of cell; reniform defined
by whitish points ; postmedial line indistinct, deep red, oblique
towards costa, then slightly waved, excurved to vein 4, then
incurved ; some slight whitish points beyond it on costa; sub-
terminal line represented by a slight whitish striga from costa
and whitish points above and below vein 6 further from termen ;
a terminal series of slight whitish points. Hind wing white,
the costal area, and terminal area to vein 2, tinged with pale
brown. Underside of fore wing brownish white, the costal area
red ; hind wing with the costal edge red.

1909: Dec. 15,—1 9° (type). Hap. 26 millim.

41036. Eratopica PpHmocausTA, sp.n. (PI. I. fig. 11, 2.)

2. Head, thorax, and abdomen deep purplish red tinged with
brown; antenne black; palpi black-brown except at tips; tarsi
black-brown with slight pale rings. Fore wing deep purplish
red tinged with brown ; a very indistinct sinuous brownish ante-
medial line; reniform red incompletely defined by ochreous,
narrow ; postmedial line indistinct, dark, oblique to vein 6, then
dentate and incurved below vein 4, some minute pale points
beyond it on costa, a terminal series of ochreous points. Hind
wing white tinged with brown, the cilia pure white at tips.
Underside of fore wing pale brown; hind wing white, the costal
half suffused with brown.

1909: May 9,—1 9 (type); May 10,1 9. Hap. 26 millim,

4524, KLYDNA BISIGNATA Hmpsn.
1908: May 12,—1 9.

MOTHS FROM SOMALILAND. 109

4676 a. RABILA ALBIVIRIDIS, sp.n. (Pl. I. fig. 13, ¢.)

Antenne of male laminate and minutely ciliated.

3, 2. Head, thorax, and abdomen white slightly mixed with
brownish; antenne tinged with ochreous. Fore wing pale
yellow-green irrorated with white, the costal area whiter to
beyond middle. Hind wing white tinged with brown. Under-
side white; fore wing suffused with brown, except the costa
and inner area which are irrorated with brown ; hind wing with
the costal and terminal areas irrorated with brown.

Ab. 1. 9. Fore wing with deeper green patch with a golden
tinge and defined by whitish on inner basal area, its outer edge
rounded and a similar small round spot distinctly defined by
white before tornus.

1908: May 28,—1 9; June 2,—1 ¢; June 21,—1 ¢ (type).
IBO a Ape, Ol Qs zyoR hall Ge eyo AA al is Wig Wah.
—1 92; May 8—1 9 (B.M.); May 10,—2 ¢; Sept. 16,—
1 9 ab. (B.M.). Year?: May,—1 $. Hap. 20-24 millim.

4742 a. ACRAPEX ALBICOSTATA, sp.n. (PI. I. fig. 14, 3.)

6. Head whitish mixed with dark brown, the antenne
ringed with’ brown towards base, thorax white tinged with
red-brown, the tegule with slight brown medial line; pectus,
legs, and abdomen white, the fore legs brown in front. Fore
wing white tinged and irrorated with red-brown, the costal
edge brown, the inner half dark brown to the postmedial line,
extending except at base to discal fold and leaving some yellow
on inner margin, met at the postmedial line by an oblique
brown fascia from termen below apex; subbasal and ante-
medial slight double oblique brown strize from costa; a black
point in middle of cell and slight striga on discocellulars with
point beyond it; postmedial line slight, brown, strongly bent
outwards below costa, then slightly waved, excurved to vein 4,
then incurved and double towards inner margin, the area beyond
it with black streaks between veins 8 and 4; an oblique slightly
waved brown subterminal line below the oblique fascia; a
terminal series of black points. Hind wing pure white. Under-
side white, the costal area of fore wing tinged with ochreous and
irvorated with red-brown.

1908 : Sept. 26,—1 ¢ (type). Hap. 22 millim.

4755. Sesam1a conrtota Hmpsn.
1909: Jan. 12,—1 9.

Genus PACHYCOA.

Type, P. olivacea.

Proboscis fully developed ; palpi obliquely upturned, slender,
the 2nd joint reaching to about vertex of head and slightly
fringed with hair behind at extremity, the 8rd short and
thickly scaled; frons with flattened corneous plate at middle
covered by a tuft of hair above and corneous plate below; eyes

110 PROF, E. B. POULTON ON

rather small, round; antenne of female somewhat laminate and
almost simple; thorax thickly clothed with rough scales and
hair, the metathorax with spreading crest ; tibie slightly fringed
with hair; abdomen without crests. Fore wing thickly clothed
with rough scales, the apex rounded, the termen evenly curved
and not crenulate; veins 3 and 5 from near angle of cell; 6 from
below upper angle; 7 from angle; 8,9, 10 stalked ; 11 from cell.
Hind wing with veins 3, 4 very shortly stalked; 5 somewhat
obsolescent from well below middle of discocellulars ; 6,7 from
upper angle; 8 anastomosing with the cell near base only.

In key differs from Yantholepis in the abdomen being without
crests,

4824 a. PACHYCOA OLIVACEA, sp.n. (PI. I. fig. 12, 9.)

2. Head whitish tinged with olive-brown and the frontal tuft
with rufous ; antenn and palpi brown; thorax olive-brown, the
metathoracic crest darker brown; abdomen olive-brown : pectus,
legs, and ventral surface of abdomen whitish suffused with brown.
Fore wing olive-brown with a reddish tinge except on terminal
area; the Ist line almost medial, slight, whitish, oblique to
subcostal nervure, then erect; postmedial line slight, whitish,
excurved to vein 4, then oblique; cilia whitish tinged with
brown and chequered with chocolate-brown at tips. Hind wing
dark brown, the cilia silvery white at tips. Underside brown,
the costal and.terminal areas of fore wing and the hind wing
irrorated with white.

1909: Sept. 16,—1 @ (type). Hap. 22 millim.

Genus ACRORIESIS, nov.

Type, 4. cgnifusa.

Proboscis fully developed; palpi obliquely upturned, slender,
the 2nd joint reaching to above vertex of head, the 8rd short,
thickly scaled; frons with flattened corneous plate at middle
covered by a tuft of hair above and corneous plate below; eyes
large, round ; antenne of female somewhat laminate and almost
simple; thorax clothed with scales and hair mixed, the meta-
thorax with spreading crest; tibie slightly fringed with hair;
abdomen without crests. Fore wing with the apex rounded,
the termen evenly curved and not crenulate; veins 3 and 5 from
near angle of cell; 6 from upper angle; 7, 8 and 9, 10 stalked;
11 from cell. Hind wing with veins 3, 4 from angle of cell;
5 somewhat obsolescent from well above angle; 6, 7 from upper
angle ; 8 anastomosing with the cell near base only.

In key differs from the other genera without an areole in the
fore wing having veins 7, 8 and 9, 10 stalked.

48246. ACRORIESIS IGNIFUSA, sp.n. (PI. I. fig. 6, 2.).

@. Head and thorax whitish suffused with cupreous red ;
pectus and legs white, the latter tinged with brown ; abdomen
brown, the ventral surface white tinged with rufous towards
extremity. Fore wing pale grey-brown suffused with cupreous

MOTHS FROM SOMALILAND, Wa

red to the postmedial line except towards base; a subbasal
chocolate-brown spot on inner margin and streak in end of cell ;

postmedial line double, brown filled in with white, very oblique
from costa to vein 6 towards termen, then excurved to vein 4,
then very inwardly oblique, with fiery red beyond it, except
between veins 6 and 4, followed by a white line; cilia with fine
whitish line at base: and white tips. Hind wing white tinged
with brown, the cilia pure white. Fore wing grey-brown,
the costa white towards apex; hind wing white with small
brown discoidal spot and curved postmedial line.

1909: Sept. 16,—1 9 (type). Hxp. 18 millim.
4857 a. KUTERPIODES PICTIMARGO, sp.n. (PI. I. fig. 15, 3.)

Antenne of male laminate and almost simple.

3. Head and tegule bright rufous; thorax ochreous ; fore
and mid tibie and the tarsi red-brown ringed with white ;
abdomen ochreous white, the 2nd to 4th segments dorsally
tinged with red-brown. Fore wing ochreous Siti, the area
beyond the antemedial line from costa to below the cell and
vein 3 suffused with bright rufous to termen; subbasal line
black with. some rufous before it on costa, sinuous, from costa
to submedian fold; antemecial line black, oblique, sinuous,
incurved above vein 1; claviform defined by red-brown at
extremity ; orbicular defined by red-brown, round; reniform
with whitish centre and annulus defined by red-brown; a
sinuous red-brown medial line; postmedial line black, slightly
defined on outer side by white on the rufous area, strongly
bent outwards below costa, slightly ineurved at igen fold,
incurved below vein 4 to below end of cell and excurved above
vein 1, some white points beyond it on costa ; subterminal line
slight, white, defined on inner side by small rather dentate
black marks from costa to vein 3, angled outwards at veins 7, 6
and inwards at discal fold, then minutely dentate, a crimson
patch beyond it at apex with oblique black striga from apex; a
terminal series of minute black lunules defined on inner side by
white, more strongly towards apex; cilia pale rufous with a
reddish-brown line near base. Hind wing silky white; the
underside with the costal area irrorated with red-brown and
with faint red-brown postmedial shade from costa.

2. Thorax, abdomen, and the basal and inner areas of fore
wing tinged with rufous; hind wing red-brown, the cilia
whitish.

1908: Aug. 15,—1 9 ; Sept. 26,—1 3 (type). 1909:
Mar. 15,—I 3 Apr. 8,—1 2 Xtype). Hap. S 20, 9 22

millim.

4857 6. KUTERPIODES CROCEISTICTA, sp. n. (Pl. I. fig. 16, 3.)

dg. Head and thorax creamy white; frons and palpi tinged
with orange, the latter with some black at side of 2nd joint ;
patagia with orange patches; fore and mid tibie tinged with

112 PROF, E. B. POULTON ON

orange, the tarsi orange ringed with white; abdomen white,
suffused with dark brown except at base and extremity. Fore
wing creamy white; antemedial line represented by orange
strie from costa and inner margin, a black point above sub-
median fold and orange point below it, inwardly oblique;
postmedial line represented by an orange striga from. costa,
black points above and below vein 5, and below the end of cell
by a black point above submedian fold, black and orange point
below it, and orange striga from inner margin. Hind wing
silky white with a very faint brown tinge. Underside of fore
wing suffused with brown.

1909: Mar. 26,—1 ¢ (type); May 8,—1 9; May 10,—1 oc.
Hap. 14 millim.

4885 a. PARATUERTA NANA, sp.n. (PI. I. fig. 17, 3.)

©. Head and thorax white with some brown scales; antennze
ringed with brown towards base ; abdomen white dorsally irro-
rated with brown, the double basal crest with some blackish
scales with a metallic gloss. Fore wing white irrorated with
brown, the terminal area more thickly irrorated; a sinuous
black-brown streak in submedian fold to the postmedial line,
with the area below it and also the area from just before the
postmedial line to the subterminal line chocolate-brown mixed
with grey ; antemedial line hardly traceable to submedian fold,
then blackish and strongly angled outwards above inner margin ;
a faint diffused reddish-brown spot in end of cell almost conjoined
toa similar discoidal spot; postmedial line black-brown, obliquely
curved and slightly waved from costa to the streak in submedian
fold where it terminates, the brown before it angled inwards at
diseal fold ; subterminal line black-brown, obliquely curved and
slightly waved, angled inwards at vein 1; a fine dark terminal
line. Hind wing ochreous yellow, the inner area tinged with
reddish brown; the postmedial area reddish brown to near
termen, which is yellowish irrorated with brown; a terminal
series of brown strive. Underside of both wings white, the
terminal areas broadly suffused with brown, the costal area of
fore wing irrorated with brown.

g. Fore wing with the costal area and disk grey irrorated
with brown and hardly paler than the inner and terminal areas,
the orbicular and reniform defined by dark brown, the latter
faintly on outer side, the former round; hind wing brownish
ochreous.

1909: Apr. 11,—1 ¢ (type); Apr. 30,—1 @ (type). Hmup.
Cie One ee O aman time

Subfam. HRASTRIANA.

5068 6. ENIsPA FLAVIPARS, sp.n. (PI. I. fig. 18, 3.)

¢. Head and thorax rufous; pectus, legs, and abdomen
whitish tinged with brown, the last with some rufous at base of

MOTHS FROM SOMALILAND. 113

dorsum. Fore wing irrorated with silvery scales, the costal half
rufous to beyond the cell, the rest of wing pale. olive-green
banded with pale yellow; an indistinct interrupted antemedial
band; a small brown spot in middle of cell and curved discoidal
striga ; the postmedial line dark and bent outwards below costa
with a yellow spot before it at costa, a yellow spot at discal fold
and incurved band from vein 4 to inner margin, some yellowish
points beyond it on costa; an interrupted maculate subterminal
yellowish band. Hind wing irrorated with silvery scales, pale
olive-green with the terminal area pale yellow; the underside
pale yellow.

1908: Oct. 31,—1 ¢g (type). 1909: Mar. 12,1 S$. Hap.
16 millim.

5142. KupuemMaA Apmova Feld.
1909: Oct. 11,—1 9.

5144, KuBLEMMA REDUCTA Butl.

1908: June 1,—1 g ; Oct. 13,—3 ¢ ; Oct. 23,—1 $; Nov. 17,
—1 3. 1909: May 8—1@2; May 10,—1 ¢g,1 9.

5149, HuBLEMMA NicRivitrA Hmpsn.

Mandera.—1908: Sept. 20,—1 $6; Oct. 11,-1 6. 1909:
Mar. 12,—1 6; Mar. 26,—1 o.
Hargaisa.—1908: Oct..—l ¢. ~

5158 @. KUBLEMMA EREMOCHROA, Sp. n. (PI. I. fig. 19, °¢.)

3. Head, thorax, and abdomen ochreous slightly tinged
with rufous; antenn tinged with fuscous ; palpi and fore legs
blackish. Fore wing ochreous tinged and irrorated with rufous
and with a few blackish scales; the costal edge blackish towards
base; traces of a waved rufous antemedial line; minute black
points in middle of cell and on discocellulars sometimes present ;
traces of a rufous medial line, oblique to the discocellulars, then
inwardly oblique; postmedial line indistinct, rufous, oblique
towards costa, then inwardly oblique, very slightly waved and
sometimes with some blackish scales on it; some faint pale and
rufous marks on costa towards apex; an oblique rufous subter-
minal shade with a series of minute white points on if, sometimes
with some black scales on their outer edges and with one to three
black points towards costa; a terminal series of black points with
more prominent spot at submedian fold. Hind wing white with
a faint rufous tinge; traces of a sinuous rufous postmedial line ;
a punctiform blackish terminal line. Underside of fore wing
suffused with red-brown except the marginal areas; hind wing
with the costal area irrorated with rufous.

@. Fore wing more strongly suffused with rufous, the white
points on the subterminal line usually obsolete; hind wing more
strongly tinged and irrorated with rufous.

1908: July 19,—1 9 ; July 31,2 g. 1909: Jan. 9,—i 9 ;
Proc, Zoot. Soc.—1916, No. VIII. 8

114 PROF, E. B. POULTON ON

Jan. 14,—1 ¢ ; Jan. 15,—1 @ ; Jan. 17,—1 2 ; Jan. 18,—1 ¢
(type); Jan. 19,—1¢,19 (type); Mar. 30,—1 gS. Hap. 18-22

millim.

5214. EKuBLEMMA scrruLA Rmbr.

1908: June 29,—1 2; Sept. 16,—1 9; Sept. 17,—19; Oct. 28,
—1 @; Nov. 17,—1¢; Nov. 19,—1¢. 1909: Jan. 19,—1 9 ;
Feb. 22,—1 9; Mar. 13,—1 6.

5282.c¢. KuBLEMMA ocHRICosTA, sp.n. (PI. I. fig. 20, 9.)

@. Head white, the antennz tinged with ochreous, the palpi
with grey-brown; thorax whitish tinged with grey-brown ; pectus
and legs white, the fore legs tinged with grey-brown, the mid and
hind legs with ochreous; abdomen ochreous white. Fore wing
ochreous white suffused and irrorated with grey-brown, the costal
area broadly ochreous; black pomts in cell towards extremity and
on discocellulars with a slight white streak between them ; some
very slight white streaks in the interspaces of terminal area, the
streak im discal fold extending to near end of cell. Hind wing
white with an ochreous tinge.

1909: Feb. 23,—1 9° (type). Hcp. 18 millnn.

52826. KUBLEMMA ARENOSTROTA, sp.n. (PI. I. fig. 21, ¢.)

g. Head white, the antenne and palpi tinged with ochreous ;
thorax whitish mixed with grey-brown ; pectus, legs, and abdomen
white tinged with ochreous. Fore wing ochreous irrorated with
white and grey-brown except on terminal area, the ochreous
forming diffused fascize on median nervure and above vein 2 to
the oblique grey-brown subterminal shade; the costal edge white ;
minute brown spots on each side of discocellulars; some slight
brown points on termen; cilia white and grey-brown with a
fine white line at base. Hind wing white slightly tinged with
ochreous. Underside of both wings almost pure white.

1909: Jan. 20,—1 ¢ (type). Hap. 20 millim.

5296. EKUBLEMMA CoNISTROTA Hmpsn.
1908: Aug. 24,—I 9.

5320 a. ToANA NIGRILINEATA, sp.n. (PI. I. fig. 22, ¢.)

S 9. Head, thorax, and abdomen pale grey-brown ; antenne
ringed with black; palpi, frons, and fore legs black-brown. Fore
wing pale brownish grey slightly mrerated with dark brown ;
a small subbasal black spot on costa; antemedial line strong,
black, oblique to submedian fold, then incurved to inner margin ;
a slight brownish medial line, excurved beyond lower angle of cell
and above inner margin; postmedial lne strong, black, arising
from the same pomt on costa as the medial line, oblique and
sinuous to vein 4, then inwardly oblique to submedian fold and
excurved above inner margin; traces of a brownish subterminal

MOTHS FROM SOMALILAND. 1 iL)

line; a strong slightly waved black terminal line; cilia with
fine brown lines through them. Hind wing whitish tinged and
irrorated with brown; postmedial line almost obsolete on costal
half, then black, oblique to vein 4, then inwardly oblique to sub-
median fold and oblique to inner margin ; a black terminal line.
Underside whitish tinged with red-brown ; hind wing with slight
brown discoidal striga.

1909: Mar. 22,—14 (type); Mar. 26,—19. Hap. 18 millim.
5576 a. CHIONOXANTHIA LEUCOPHAA, sp.n. (PI. I. fig. 23, 3.)

S$ 9. Head, thorax, and abdomen grey-white mixed with
brown ; palpi black-brown ringed with white. lore wing grey-
white suffused witn brown; a slight smuous blackish subbasal line
from costa to submedian fold; antemedial line double, blackish
filled in with white and defined on inner side by white, sinuous, a
black streak beyond it in submedian fold; orbicular white defined
by black, round, some black in the cell between it and the white
discoidal bar; postmedial line brown defined on each side by
white, obliquely excurved from costa to vein 4, then incurved ;
subterminal line whitish defined on inner side by diffused brown
forming a dark patch on costal area, angled inwards at discal
fold, excurved at middle, then incurved and slightly waved; a
terminal series of blackish striz. Hind wing whitish suffused
with brown; the underside white irrorated with brown, a small
brownish discoidal spot, curved postmedial line, a diffused sub-
terminal line.

1908: May 28,—2 9 (1 in B.M.); June 1,—1 ¢ (type);
June 2,—1 2. zap. 16 millim.

5589 a. CHDICODIA STRIGIPENNIS, sp.n. (PI. I. fig. 24, ¢.)

¢. Head and thorax red-brown slightly mixed with whitish ;
antenne dark brown; palpi at base and the base of 3rd joint
white; abdomen grey irrorated with brown ; pectus and ventral
surface of abdomen white tinged with brown. Fore wing red-
brown tinged with grey and irrorated with blackish forming
obscure streaks on the veins and above and below submedian fold,
except on the terminal area which is slightly paler except at
middle and tornus ; an indistinct waved brown antemedial line,
double at costa; a black discoidal striga with point above it on
costa; postmedial line brown, defined on inner side by whitish
towards costa, oblique to vein 6, then slightly waved and incurved
below vein 4; the postmedial area rather darker brown with some
whitish points on costa ; subterminal line white defined on inner
side by brown, very slightly excurved below vein 7 and at middle ;
a terminal series of minute black lunules. Hind wing pale red-
brown, the cilia white tinged with red-brown at base; the under-
side white, the costal and terminal areas irrorated with brown,
the apex suffused with brown, traces of sinuous postmiedial and
subterminal lines.

1908: Oct. 15,—1 3 (type). Hrp. 20 millim,

Q#

116 PROF, E. B. POULTON ON

5589 6. CHpicopIA LIMBATA Butt.

1908: Apr.28,—1 3; May2,—19; May4,—29; July16,—1 9
(B.M.); July 17,—1 4; July 24,—19; Aug.15,—1 4,19; Sept.
13,—1 9; Sept. 15,—1 9; Sept. 16,—1 9; Sept. 23,—1 9; Sept.
25,—192; Sept. 26,19; Sept. 27,135,199; Sept. 29,—1 9;
Sept. 30,—1 2; Oct. 1,—19; Oct. 3,—1 9; Oct. 4,—1 9 (B.M.);
Oct. 11,—1 2; Oct. 13,—1 6; Oct. 14,—1 9 ; Oct. 18,—1 2 (BM);
Nov. 17,—1@. 1909: Jan. 17,—19; Jan.19,—1 ¢ (B.M.); Feb.
22 1G; Mar. 10,19; Mar 11,1 ¢; Apr. 8—1lg, 19);
Apr. 10,—16,19; May 8,—1¢ (B.M.); Nov. 23,—1 ¢. 1910:
Jan. 12,—1I @?.

5589 c. CHDICODIA MELANOGRAPHA, sp. n. (PI. I. fig. 25, 9.)

9. Head, thorax, and abdomen pale red-brown; antenne
blackish ; palpi except at tips, pectus, legs, and ventral surface of
abdomen white, the fore and mid tibie tinged with brown, the
tars’ brown ringed with white. Fore wing pale red-brown; a
black point on costa near base, some scales m base of submedian
fold and a slight patch of scales on the costa before the antemedial
line, which is black, waved; a black discoidal striga, its lower
extremity touching the sinuous blackish medial line, which is
excurved to lower angle of cell; postmedial line blackish,
approximated to the medial line, shghtly waved, oblique to vein 5,
then inwardly oblique; subterminal Hne rather diffused, black,
very slightly excurved below vein 7 and at middle; some black
scales on termen. Hind wing pale red-brown, the termen rather
darker red-brown te vein 2; cilia white shghtly tinged with
rufous. Underside white tinged with rufous.

1909: Apr. 10,—1 @ (type). Hap. 24 millim.

5633 a. OZARBA SEMITORRIDA, sp.n. (Pl. I. fig. 26, g.)

3. Head ochreous brown; thorax red mixed with leaden grey-
brown; abdomen pale ochreous; palpi, pectus, legs, and ventral
surface of abdomen white, the fore and mid tibiz and tarsi banded
with brown. Fore wing deep red suffused with dark leaden grey,
especially towards. costa, to the reniform and postmedial line, the
rest of wing white tinged with red-brown and with a red patch
on postmedial part of costa; mmute subbasal white points on
costa, in and below the cell; traces of a waved antemedial line
defined on inner side by a whitish striga from costa; a white
point in middle of cell; reniform white with pale brownish centre,
narrow and oblique; postmedial line treble, red-brown filled in
with white, obliquely excurved from costa to vein 4, then ineurved,
touching the upper and lower extremities of the reniform, three
white points beyond it on the costal patch ; subtermmal line
white defined on inner side by brown, excurved below vein 7 and
at middle; a dark brown terminal line; cilia with brown shades
at discal and submedian folds. Hind wing whitish suffused with
reddish brown, the cilia whiter. Underside whitish tmged with
red-brown except on inner area of hind wing.

MOTHS FROM SOMALILAND. eee leley

@. Head, thorax, and basal half of fore wing much redder,
sometimes crimson-red and with the markings of outer half of
fore wing crimson-red.

1908 : Sept. 22,—1 $ ; Sept. 30,—1 ? (type); Oct. 18,—1 9;
Oct. 22,—1 S (type). 1909: Mar. 24,1 9. Hxp. 3 18, 2 20

millim.

5635 a. OZARBA ENDOSOOTA, sp.n. (PI. I. fig. 27, 2.)

¢ 2. Head and thorax ochreous, the head between antennz
and patagia with deep red patches, the patagia with black-brown
stripes above ; antenne dark brown; pectus and legs white, the
fore legs dark brown in front, the tarsi dark brown ringed with
white ; abdomen white, dorsally suffused with brown. Fore wing
with the basal half ochreous tinged with red-brown, the area
below the cell suffused with dark brown except at base, the rest
of wing grey-white irrorated with dark brown; slight brown
marks on costa towards base; an oblique antemedial brown striga
from costa and sinuous line from cell to inner margin defined on
inner side by whitish and with short brown streaks before it in
submedian fold and above inner margin ; a brown spot in end of
cell and whitish discoidal striga ; a small black spot on costa above
end of cell; postmedial line hardly traceable, excurved to vein 4,
then incurved, some black suffusion beyond it on costa; sub-
terminal line white, curved, a blackish patch beyond it at discal
fold; a terminal series ef minute blackish spots. Hind wing
whitish strongly suffused with brown; the underside white
irrorated with brown, the terminal area suffused with brown, a
small blackish discoidal spot and curved postmedial line.

1908: Oct. 11,—19 (type); Nov. 22,—1 ¢. Hep. 20 millim.

5637. OzARBA CONSANGUIS Hmpsn.
1908: Oct. 17,—1 9; Oct. 25,1 9. 1909: Apr. 7,—1 Q;
Apr. 8,—1 @.

5637 a. OZARBA HEMIPYRA, sp. n. (PI. I. fig. 28, 2.)

@. Head whitish mixed with blackish, the upper part of frons,
antenne, and palpi blackish, the last with whitish ring at ex-
tremity of 2nd joint; thorax black with some reddish scales ;
pectus and legs ochreous white, the fore legs with some black in
front, the tarsi banded with blackish ; abdomen reddish ochreous
irrorated with black, the basal crest and a bar before the anal tuft
black, the ventral surface ochreous. Fore wing black slightly
mixed with red to the medial line, the rest of wing fiery rufous
with a slight greyish tinge on terminal area; subbasal line black
slightly defined by red, waved, from costa to submedian fold ;
antemedial line black slightly defined on inner side by red at
costa and inner margin, waved; medial line closely approximated
to the antemedial line, black slightly defined on outer side by
white, incurved just below median nervure, a black point beyond

DSi PROF. E. B. POULTON ON

it on costa ; postmedial line only defined by a deeper rufous shade
on its outer side, excurved to vein 4, then incurved, some whitish
points beyond it on costa; subterminal line whitish defined on
inner side by a deep rufous shade, excurved below vein 7 and at
middle, then waved, some deep rufous beyond it at discal and
submedian folds ; a terminal series of minute deep rufous lunules.
Hind wing whitish suffused with brown and with a fine brown
terminal line; cilia paler. Underside of fore wing whitish suf-
fused with brown and with some reddish ochreous at middle of
costa ; hind wing whitish irrorated with brown and with a small
dark discoidal spot.
1908: Oct. 2,—1 2 (type). Hap. 20 millim.

5638. OZARBA HEMIMELZNA Hmpsn.

1909: Mar. 21,—1 9; Mar. 28-2 9; Apr. 7,—1 ¢;
Octn5 —a110

5639 a. OZARBA HEMISARCA, sp. n. (PI. I. fig. 29, 3.)

3d. Head, thorax, and abdomen ochreous with a faint rufous
tinge; palpi except at tips, pectus, legs, and ventral surface of
abdomen white, the fore and mid tibiz ochreous, the tarsi
ochreous ringed with white. Fore wing with the basal half
ochreous slightly tinged with rufous, the terminal half white
slightly tinged with olive-brown ; two slight rufous marks on
costa towards base; traces of a waved rufous antemedial line with
a small deep rufous spot at costa; a red-brown medial line, oblique
and sinuous to lower angle of cell, then slightly incurved, with
minute blackish spots on it at costa and upper angle of cell and
with the area between it and the closely approximated white
postmedial line rufous, this line oblique to vein 4, then incurved ;
subterminal line whitish, defined on inner side by brown towards
costa, slightly angled inwards below costa and incurved below
vein 3; a black-brown spot at apex and terminal series of points ;
cilia tinged with red except at apex. Hind wing ochreous white,
the area beyond lower angle of cell with a reddish tinge, the
termen tinged with brown towards apex; cilia white. Underside
ochreous white.

1908: Nov. 19,—1 ¢ (type). Hap. 18 millim.

5639 5. OZARBA EXOLIVACEA, sp. n. (PI. I. fig. 30, 3.)

3. Head and tegule pale reddish ochreous ; antenne brown ;
thorax white slightly tinged with brown; pectus and legs
ochreous white, the fore Falbies and the save brown ringed
with white; abdomen ochreous tinged with brown. Fore wing
white irrorated with blackish scales, the terminal half faintly
tinged with olive-green except at apex, the costa suffused with
brown towards base; an indistinet double waved brownish ante-
medial line; an oblique dark brown medial shade diffused to the

postmedial line and on postmedial costal area; reniform with

MOTHS FROM SOMALILAND. 119

rufous centre and white annulus, narrow, oblique, and con-
stricted at middle, a whitish patch above it on costa; postmedial
line double, dark, oblique towards costa, then excurved to vein 4,
then incurved, nee white points beyond it on costa; sub-
terminal line whitish defined on inner side by brown, slightly
excurved below vein 7 and at middle; a slightly waved brown
terminal line; cilia with series of brown marks except at apex.
Hind wing ochreous suffused with reddish brown especially
towards termen. Underside white irrorated with brown; fore
wing tinged with ochreous except the inner area.

2. Head, thorax, and abdomen ochreous ; fore wing with the
basal half tinged with ochreous, the terminal half suffused with
pale olive-green, the medial shade narrower and not diffused to
the postmedial line except below the cell or on the postmedial
costal area.

1908: Sept. 22,—1 9 (type); Oct. 15,—1 ¢ (type). Hap.
22 millim.

5639 c. OzARBA MESOZONATA, sp. n. (PI. I. fig. 31, d.)

3 9. Head, thorax, and abdomen white faintly tinged with
brown ; antenne brown; palpi brown at sides except at ex-
tremities of 2nd and 3rd joints; tarsi black-brown ringed with
white. Fore wing white, the basal area faintly tinged with
brown, the terminal area suffused with rufous ; two slight dark
marks on eosta near base; a broad chocolate-brown medial band
edged by black lines defined by white, narrower towards costa
and slightly constricted in the cell; some whitish points on
costa and a brown patch on costal ar ea before the faint brownish
subterminal line, which is slightly excurved at middle; a
terminal series of black striz slightly defined on inner side by
white; cilia dark brown irrorated with grey, white at apex.
Hind wing ochreous white tinged with brown; a fine brown
terminal line. Underside white tinged with reddish ochreous.

1908: Sept. (919; ‘Sept. 21, g 5 Sept. 221 9;
Octawis. Ing. (uype)e LOO Apr Oe ape ho
20 millim.

3639 d. OzaRBA ENDOPLAGA, sp.n. (PI. I. fig. 32, 3.)

$ 9. Head and thorax ochreous white; antennz brown ;
abdomen ochreous tinged with brown; palpi, pectus, legs, and
ventral surface of abdomen white, the palpi tinged with brown
towards base, the fore and mid tibie suffused with brown, the
tarsi brown ringed with white. Fore wing creamy white suffused
with rufous especially on terminal half; a large conical chocolate-
brown patch defined by white on medial area from below costa
to inner margin, with slight black streak above it on costa in the
male; some whitish points on postmedial part of costa and some
prove on costa before apex; a black-brown terminal line defined
on inner side by pale yellow which expands towards costa ; cilia
dark brown with a greyish tinge. Hind wing white tinged with

120 PROF. E. B. POULTON ON

reddish brown especially in female, the cilia whiter. Underside
ochreous white tinged with brown.

1908: Sept. 27,19; Oct. 11,—1 2 (B.M.). 1909: Apr. 19,
—l ¢ (type); Nov. 7,—1 9. Hap. 20 millim.

5656. OZARBA SANCTA Staud.
1908: June 29,—1 ¢.

5685. OzARBA PH#A Hmpsn.
1908: Feb. 11,—1 ¢.

5713. AmMyYNA ocro Guen.
1908: Jan. 30,—1 g. 1909: Apr. 7,—1 $; Apr. 30,—1 @.
5718. AmynaA PuNCTUM Fabr.

1908: May 29,1 ¢. 1909: Mar. 14,—2 9; Apr. 6,—1 ¢,
1 2; June 25,—1 ¢.

5891. Husrroria MIANOIDES Hmpsn.
1909: Apr. 7,—3 @.

5942 a. HULOCASTRA ARGYROSTROTA, sp.n. (PI. I. fig. 33, 7

3 @. Head and thorax ochreous slightly tinged with rufous;
antenne reddish brown; palpi brown except at tips; tibize
and tarsi banded brown and white; abdomen brown with
white segmental lines, the ventral surface ochreous white
irrorated with brown. Fore wing with the basal half ochreous
slightly tinged with rufous, the terminal half suffused with
red-brown and black-brown and with patches of silver scales ;
slight subbasal and antemedial brown marks on costa and traces
of a sinuous antemedial line with some silvery scales beyond it ;
a sinuous black medial line defining the inner edge of the dark
area; an ochreous discoidal striga with some black before it;
postmedial line black defined on outer side by ochreous, more
strongly at costa, oblique from below costa to vein 6, slightly
incurved at discal fold, oblique and slightly waved below vein 4,
an oblique brown line beyond it towards costa ; subterminal line
represented by silver scales defined on inner side by blackish,
forming diffused marks below costa and at middle, excurved
below vein 7 and at middle; a terminal series of black striz
slightly defined on inner side by white ; cilia with white patches
at apex and discal fold. Hind wing whitish suffused with
fuscous brown; the underside bluish white irrorated with fuscous
brown, the terminal area more suffused with fuscous, a small
blackish discoidal spot and curved postmedial line.

1908: Sept. 21,—1 2 ; Oct. 15,—1 9 5 Oct. 24—1 ¢ (type).
Year ?: Oct. 25,—1 gd. Hep. 16 millim.

5984. LopHorACHE FULYIRUFA Hmpsn.
1909: Apr. 5,—1 9.

MOTHS FROM SOMALILAND. NDA

6081. HopLorARAcHE NUBILA Hmpsn.
1908: Sept. 21,—-1 @.

6081 a. HopLoraRACHE ECTORRIDA, sp.n. (PI. I. fig. 36, 3.)

Hoplotarache nubila, ab. 1, Hmpsn. Cat. Lep. Phal. B.M.
ky JOS Ds

3d. Head and thorax white, the dorsum of thorax with black
scales mixed except in front; antenne fuscous; palpi black
at tips, the frons with lateral black bars; tarsi black ringed
with white; abdomen creamy white with dorsal fuscous segmental
bands, the ventral surface white. Fore wing white; subbasal
line defined on each side by grey, sinuous, from. costa to median
nervure; grey streaks on costa and above vem | before the
antemedial line, which is defined on each side by grey, waved,
some grey beyond it below median nervure; small dark grey
annuli in middle of cell and on ahecouell lars: an oblique dark
grey striga from middle of costa, spot above median nervure
and waved black line from cell to inner margin; an oblique
dark grey postmedial striga from costa, two black strive beyond
the cell with some grey before them and a waved black line
from lower angle of cell to inner margin; the terminal area
chocolate-brown, leaving an oblique wedge-shaped white patch
on costal area beyond the postmedial line and below the cell
extending to the medial line; subterminal line white with
two small wedge-shaped black marks before it below costa,
excurved below costa and at middle, then incurved and slightly
waved, and with black marks beyond it above and below vein 2;
a terminal sertes of small black ‘spots defined on inner side by
white; cilia wholly white at middle, red-brown at base, with
white tips towards apex and dark leaden-grey tips at discal
fold and towards tornus and with slight blackish line through
them. Hind wing white; the underside with brown spots at
middle of costa and apex and postmedial bar from costa.

2. Dorsum of thorax grey and black ; abdomen red-brown ;
fore wing with more grey suffusion on the white area; hind
wing red-brown, the cilia white at tips; the underside white
suffused with brown, a brown discoidal bar and postmedial line
excurved beyond the cell.

S08 Noy So 1909 Apr 6, eo Apr. 8d.
1 @ (types) ; Apr. 9,—1 9 ; Apr. 10,—2 Ove” ANoir. vee OF
May 7,—1 9; May 9,—1 o..

Also in the British Museum from Br. E. Africa, Athi Valley

(Crawshay), 1 3. Hxp. 20 millim.

6081 6. HopLoTaARACHE C#RULEOPICTA, sp.n. (PI. I. fig. 37,3.)

3. Head, thorax, and abdomen ochreous tinged with rufous;
pectus, legs, and ventral surface of abdomen ochreous white, the
fore and mid tibia and the tarsi brown ringed with white. Fore
wing with the basal half creamy white, the terminal half olive-

122 PROF, E. B. POULTON ON

brown ; subbasal line double, olive-brown, sinuous, from costa to
vein 1; antemedial line double, olive-brown; a black point in
middle of cell and incomplete black discoidal annulus slightly
defined by white; medial line dark, defining the pale area,
oblique to lower angle of cell, then incurved; an oblique wedge-
shaped postmedial creamy-white patch from costa, then a diffused
dark line, strongly imcurved and with patches of silvery-blue
scales beyond it; subterminal line interrupted in places, cupreous
red defined on each side by creamy white, excurved to near termen
below vein 7 and at middle, then slightly waved; a terminal
series of black striz defined by creamy white; cilia creamy white
from vein 4 to submedian fold. Hind wing creamy white, the
veins and terminal area tinged with brown; the underside
creamy white, the terminal area tinged with brown, a small
blackish diseoidal spot and faint brownish medial and postmedial
bars from costa.

@. Fore wing with the basal half tinged with red-brown ;
hind wing cupreous red-brown, the cilia white at tips, the renter
side creamy white tinged with red-brow n, the terminal area
suffused with red-brown, a small dark discoidal spot and curved
postmedial line.

1909: Apr. 9,—1 ¢; Apr. 15,—1 @ (type); Apr. 24,—i ¢
(type). Hup. 22 millim.

6089. MervrarropLAstTA INSsociA Wk.
1908: May 4,—1 3.

6091 a. AULOTARACHE PLUMBEOGRISEA, sp.n. (PI. J. fig. 34, 2 .)

Q. Head, thorax, and abdomen reddish ochreous, the patagia
suffused with leaden grey; pectus, legs, and ventral surface of
abdomen ochreous white, the fore legs tinged with red-brown.
Fore wing brownish suffused with leaden grey; some ochreous
and rufous on inner margin towards base; traces of a sinuous
dark antemedial line from cell to inner margin, faintly defined
on inner side by ochreeus; claviform a very narrow ochreous
mark defined by some black scales; orbicular on outer side
and reniform on inner side very faintly defined by black; post-
medial line dentate, indistinct and brown from below costa to
vein 6, then blackish and defined on outer side by yellow and red
patches in the interspaces, oblique below vein 4; a terminal
series of minute black points defined on inner side by white
points. Hind wing white, the costal and terminal areas tinged
with brown. Underside of fore wing and costal area of hind
wing suffused with reddish brown.

1908: Sept. 23,—1 3; Sept. 25,—1 9; Oct. 17,—1 Q (type);
Oct. 24,—1 9. 1909: Apr. 15,—1 9; Apr. 20,—2 2; May 9,
—1 9 (B.M.); May 10.—1 9; May 12—1 9. Hap. 22-

26 millim.

MOTHS FROM SOMALILAND, 12s}

6122. Taracun zeLLERL Wllern.
1908: Sept. 18,—1 ¢.

6155. TARACHE UMBRIGERA Feld.
1908: May 24,—1 ¢. 1909: Apr. 18,—2 9.

6167. TARACHE OPALINOIDES Guen.

Mandera.—1908: July 17,—1 9. 1909: Apr. 9,—1 9 ;
Apr. 10,—1 ¢; Apr. 27,—1 9.
Hargaisa.—1908: Oct..—1 d,1 2.

6175, TARACHE CARNESCENS Himpsn.
1909: Oct. 22,—1 2.

6182. TaRACHE HORTENSIS Swinh.

1908: Aug. 16,—1 @; Sept. 28,—1 2; Sept. 29,—1 ©.
1909: Apr. 7,—1 2; Apr. 8—1 2; Nov. 24,—1 ¢.

6187 a. TARACHE MESOLEUCA, sp.n. (PI. I. fig. 38, 3.)

3. Head, thorax, and abdomen white, the dorsum of thorax
behind the tegule with grey mixed. Fore wing pale leaden
grey, the basal area with some white mixed; a subbasal white
point below costa and streak above base of vein 1; a waved
white antemedial line ; a white medial band; a black annulus in
the cell towards extremity and rather elliptical discoidal annulus;
a triangular white patch on postmedial part of costa with the
faint diffused red-brown postmedial line arising from it, strongly
incurved below vein 4; a very indistinct inact eabeen sain
line with some white on it at costa, incurved and with white
seales on its outer edge below vein 3; a terminal series of black
strie defined on inner side by white on inner half; cilia with
white mixed from vein 3 to submedian fold. Hind wing white,
the costal area and termen, except towards tornus, tinged with
brown ; the underside white with small brown discoidal spot.

1908: Aug. 15,—1 ¢ (type). Hap. 18 millim.

6191 a. TaRACcHE MroGoNA, sp.n. (PI. I. fig. 39, 9.)

2. Head grey-brown, the palpi white except at tips; thorax
brownish white with fuscous scales mixed; pectus and legs
white, the fore and mid tibie banded with fuscous, the tarsi
black ringed with white; abdomen red-brown, the ventral
surface white. Fore wing with the basal area brownish white
with some red-brown scales towards costa and a grey tinge
at base of inner margin, its outer edge rather oblique and
diffused ; a black point in the cell near base; the rest of wing
chocolate-brown tinged with purplish grey; a conical brownish-
white postmedial patch on costa with the blackish postmedial
line arising from it, incurved below vein 4 to below end of cell
and slightly angled outwards at vein 1; subterminal line formed

124 PROF. E. B. POULTON ON

by brownish-white scales, slightly waved, ineurved below vem 3
and ending at tornus ; a terminal series of black strize ; cilia with
a slight dark line through them. Hind wing pale red-brown,
the terminal area darker; cilia fuscous with a white line at base
and white tips except towards tornus ; the underside with white
patch on costa towards apex.

1909: Apr. 10,—1 @ (type). Hap. 24 millim.,

Subfam. HureLian”.
6258. EuTELIA DiscistrigA Wlk.
1909: Feb. 27,—1 9; Apr. 20,—-1 9; June 12,—1 9.

6258 a. HUTELIA GRISESCENS, sp.n. (Pl. I. fig. 40, 3.)

3 2. Head, thorax, and abdomen grey, the thorax tinged with
rufous ; palpi with the base of 2nd and 3rd joints brown ; tarsi
brownish with pale rings ; abdomen with some rufous on dorsum,
the crests and anal tuft blackish. Fore wing with the basal
area rufous defined by the deeper rufous antemedial line, which
is angled outwards below costa, then incurved ; the rest of wing
grey ; a faint reddish-brown medial line, bent outwards to the
discocellulars and incurved below the cell; postmedial line black
with some fuscous beyond it towards costa, oblique and slightly
sinuous to vein 6, then almost obsolete and much interrupted,
excurved at middie then incurved, some rufous beyond it at discal
fold and in submedian interspace; a browntsh patch on costal
area with two white points at costa before the subterminal line,
some yellowish rufous below and beyond it; subterminal line
slight, whitish and somewhat waved, incurved below costa and
vein 3; a terminal ‘series of black strie; cilia dark brown,
chequered with white at base. Hind wing grey-white, the
terminal area tinged with brown and with brownish streaks
on the veins; a fine black terminal live ; cilia chequered blackish
and white; the underside with the costal area and terminal area
to vein 2 tinged with rufous, a blackish discoidal point and
punctiform postmedial line.

1909: Mar. 13,—1.d¢. 1910: Mar.14,—1 9; Mar. 16,—1 ¢
(type). Hap. 5 22, 2 26 millim.

Subfam. STICTOPTERIN ZA.

6458. STENOSDICTA GRISEA Hmpsn.

1908: Aug. 15—1 ¢,2 9; Aug. 24,—1 9; Sept. 13,—1 2;
Sept. 21,—1 3g; Oct. 15,—1 3; Oct. 28,—1 9. 1909: Feb. 18,
—l1 g; Mar. 14,—1 9; Mar. 22,—1 3; Mar. 28,—1 ¢.

Subfam. AConTIANE,

6863. EARIAS INSULANA Boisd.
1908: Oct. 31,—1 ¢.

Or

MOTHS FROM SOMALILAND. 2

6980. MAuRILIA ARCUATA WIlk.
1908: Oct. 25,—1 3.

7068. NeGera LuMINOsSA WIlk.
1908: July 11,—1 9°.

7116. AcontrIA ALBAGO F,
1909: Apr. 24,—1 9;. Apr. 26,—1 9.

7117. AcontrA GEPHYRIAS Meyr.
1909: May’8,—1 ¢.

Subfam. CAaTrocaLina,

7353, ULOTHRICHOPUS TINCTIPENNIS. Hmpsn..
1909: July 6,—1 ¢.

7362. CHELECALA TREFOLIATA Butl.
1910: Jan. 10,—2 9°.

7367. HyporacHa INDECISA Wlk.

1908: July 24,—1 ¢ ; Aug. 26,--1 9°.

7423. CYLIGRAMMA LATONA Cram.

1909: May 20.—1 9; May 21,—1 9; May 23,—I ¢;
May 29,—1 6; May 30,—I 2; May 3l-—3 g,2 9; June 1,
8 ¢, 69; June 2,—2 9; June 4-2 6; June 5—1 9 ;
June 6,—6 g, 4 9; June 7,2 g,1 9; June 9,1 6, 8 Q;
June 14,—1 ¢; Sept. 8,—1 d. 1910: June 4,1 9°.

7493 a, ACANTHONYX SERIOPUNCTA, sp.n. (Pl. I. fig. 41, 3.)

3. Head and thorax ochreous yellow, the tegule tinged with
rufous ; antenne whitish tinged with rufous; abdomen ochreous
white with dorsal rufous segmental lines, the ventral surface
ochreous. Fore wing ochreous yellow sparsely irrorated with
red-brown scales; subbasal red-brown points below costa and
cell ; a minute antemedial red-brown spot below costa and points
on median nervure and vein 1; an oblique chocolate-brown
discoidal bar tmged with grey, rather rounded above; a minute
postmedial red-brown spot below costa, then a curved series
of pots on the veins; fine brownish lines on termen and
through the cilia. Hind wing white. Underside white; fore
wing with the costal area ochreous, the terminal area tinged
with ochreous except towards tornus ; hind wing with the costal
area and the termen narrowly tinged with ochreous.

Hargaisa.—1908: Oct.,—1 ¢ (type). Hap. 40 millim.
7667. ACH&A CATELLA Guen.

Mandera.—1908: Dec. 17,—1 9. 1909: Jan. 9,—1 g;
May 24,—1 9; May 25,—2 9; June 7,—1 ¢; July 9,—1 9.
Durbar.—1908: Dec. 6,—3 ¢.

126 PROF. E. B. POULTON ON

7747. PARALLELIA ALGIRA L.
1909: Apr. 8,—1 6.

7764 a. PARALLELIA RECTIFASCIA Laweett.
1909; Apr. 22;—1 GC.

7786. GRAMMODES STOLIDA Fabr.

1908: May 4,—19; Aug. 28,—1 6; Sept. 11,--1 dg; Sept. 12,
—l1 dg; Sept. 23,—1 3; Sept. 25,—1 9; Sept. 80,—2 9; Oct. 1,
—1 ¢. 1909: Mar. 14,—1 ¢; Apr. 15,—1 9; Apr. 24,—1 9;
May 9,—-1 3, 49; May 10,235,592; May 12,—29; May 14,
13g; May 21,—1 g., 1910: Feb. 9,—1 9.

7792. CHALCIOPE HYPPASIA Cram,
1908: Nov. 19,—1 ¢.

7855.: Mocis REPANDA F,
1909: June 4,—1 GC.

8075. CERocALA ILLUSTRATA Holl.

Mandera.—1908: Feb. 11,—1 9; Apr. 28,—1 9; May 28,—
1 2; June 30,—1 9; July 27,19; July 31,16; Aug. 23,—
1 g; Aug. 26,—1 9; Sept. 22,—1 9; Oct. 22,—1 6; Nov. 13,—
19; Nov. 25,—1 2. 1909: Jan. 9,—2 9; Jan. 15,—1 9;
Jan. 17,—3 9; Feb. 14,—2 9; Feb. 16,—1 9; Feb. 17,—2 9;
Feb. 21,—1 ¢; Mar. 10,—1 9; Mar. 13,—2 9; Mar.17,—3 92;
Mar. 20,—1¢9,192; Mar. 21,—1 9; Mar. 22,—2 9; Mar. 24,—
13,19; Mar. 26,—49; Mar. 28,—19; Mar. 29,—19; Apr. 7,
—l 9; Apr. 8,—2 $,29; Apr. 10,—1 36,29; Apr. 11,—3 6,
192; Apr. 14,—1 6, 19; Apr. 19,—19; May 8,19; May 10,
—1d9,29; June 9,—1 d; Oct. 11,—1 6; Nov. 11,—2 9; Nov.
12,2 2. 1910: Mar. 14,-—-1 9.

Gan Libbah.—-1908: June 24,—1 9.

Berbera.—1908: Mar. 4,—2 9.

8077 a. CEROCALA ALBIMACULA, sp.n. (PI. I. fig. 42, 3.)

3d. Head, thorax, and abdomen brown mixed with white, the
thorax mostly brown, the tegule dark brown, white at base and
tips; antenne ringed black and white; tarsi brown ringed with
white; ventral surface of abdomen white. Fore wing whitish
almost wholly suffused with grey-brown and reddish brown,
leaving a rather quadrate white patch beyond the reniform ;
antemedial line obsolete on costal area, then double, black,
slightly sinuous, with diffused silvery and black scales before it,
the outer line slightly defined on outer side by white: orbicular
and reniform with silvery and brown centres incompletely defined
by black; the former small, round, the latter with dark streak
before it in lower part of cell; postmedial line black slightly
defined on inner side by whitish, excurved below costa and

MOTHS FROM SOMALILAND. War

between veins 3 and 2 to the subterminal line, then retracted
upwards to lower angle of cell, waved to vein 1 and oblique to
inner margin, some black and silvery seales beyond it in its
sinus; subterminal line whitish defined on inner side by black
marks and some silvery scales between veins 7 and 3, angled
outwards below vein 7 and excurved at middle; a waved dark
terminal line; cilia chequered brown and white. Hind wing
white suffused with reddish brown ; a dark discoidal spot with
some white beyond it; an indistinet dark subterminal shade
with dark patches beyond it below apex and at middle with white
above them; cilia white chequered with brown. Underside
white; fore wing with round black discoidal spot, some brown
suffusion from below end of cell and fuscous subterminal and
terminal marks towards apex; hind wing with black discoidal
spot, some brown suffusion from below end of cell to the sinuous
brown postmedial shade, and blackish patches on termen below
apex and at middle.

@. Fore wing with more white, especially at base and on
terminal area except at apex.

1908: Sept. 29,—1 2 (type); Oct. 11,—1 ¢ (type). 1909:
Apr. 7,—1 9; Apr. 20,—1 $; Oct. 5,—1 9; Oct. 14,—1 9.
up. & 28, 2 34 millim.

8078. CrrodALA opp1A Druce.

1908: Feb. 1,—1 g; Feb. 7,—1 9; Feb. 11,—1 ¢; Sept. 17,
---19; Sept. 18,—1 9; Sept. 2 3 ie Sept. 22,—1 9; Sept. 23,
—1 9; Sept. 27,—1 3d; Oct. 3,—1 9. 1909: Feb. CLs QO;
Mar. | ae @; Mar. 11,—1 9; Mar. 14,—1 9; Mar. 19,--1 9;
Mar. 22,— 19; Mar. 26,—1 9; Apr. 6,—1 Q; Apr. 7,—1 3;
Apr. 8,—1-9; Sept..— 1 ¢; Oct.5,—1 9; Nov. 6,—1 92. 1910:
Jan. 5,—1 @.

8092. GNAMPTONYX INNEXA WIk.

ie abe Bites July 17,—1 @; een A Q. yee
Mar. 14,— oN Oke Meng, 28, —I @; Mar. 22; Apr. 3,—
19. 10 ae ar. 14,—1 9; Mar. 20,—1 a

Hargaisa.—1908: Oct.,--1 9. -

8117. PericyMA MeraALEucA Hmpsn.
1908: Aug. 24,—1 g. 1909: May 10,—1 9

+ .

8125, CorryrA LEUCOPTERA Hmpsn,

The ser Hee, besides the ty ypical form, ineludes specimens agree-
ing with C. dispar Piing., 0. ‘fascioluta Warr., C. balnearia Dist.,
Ca impar Wmpsn.,and C. eremochroa Himpsn., White h are evide ntly
forms of one variable species; they also occur together in the
Hoggar Mts., 8. Sahara, vide Rothschild, A. M. N. H. (8) xvi.
p. 255 (1915).—G. F. H.

1908: Mar. 22,—1 g; July 8.—2 9; Aug. 20,—1 9 (B.M.);

ef

Sept. 16,—1 3 ; Sept. 19,—1 6,193 Oct. 29, G(r vias

128 PROF. E. B. POULTON ON

1909: Feb. 17,—1 2 (B.M.); Mar. 11,—1 9° (B.M.); Apr. 7,
Ons Acre 0-1 12) (BAMe) SaiSept. 821 ase vi) aor
Sept.,—4 g; Oct. 14,—2 ¢ (1 in B.M.),19; Oct. 22,—1 ¢; Dee.
28,—1 S. 1910: Feb. 10,—1 9; Mar. 20,—1 3.

8132. Cortryra RosACEA Rebel.

1908: July 17,—-1 2; Oct. 2,—1 9; Nov. 25,—17¢ (B.M.).
1909: Mar. 14,—1 ¢; May 10,—1 9; Oct. 14,—1 9.

8135. CorryTa CANESCENS W1Ik.

Mandera.—1909: Apr. 14,—1 o.
Hargaisa.—1908: Oct.,—1 92.

Subfam. PHyToMETRIN2.

8292. PHytromerra NI Hiibn.
1908: Oct. 1,—1 9. 1909: Apr. 6,—1 2; Sept.-—l 9.

8295. PHYTOMETRA LIMBIRENA Guen.
Hargaisa.—1908: Oct..—1 ¢.

8330. PHytomEeTRA AcUTA WIlk.

1909: Mar. 30,—1 9; Apr. 6,—1 3; Apr. 29 —1 OC” Miaye3.
—l1 3; May 21,—1 ¢.

Subfam. Nocrumnaz.

PANDESMA ANYSA Guen.
1909: Apr. 6,—1 ¢.
POLYDESMA COLUTRIX Geyer.
1908: June 18,—3 2; July 2,19; July 5,—2 ¢ (LinB.M.);
July 25,—1 9; July 26,—1 9; Aug. 1,—1 9. 1909: July 13,
¢

PROCONIS ABROSTOLOIDES Hmpsn.

1909: Sept. 17,—1 9; Sept..—Il 9; Oct. 11,—1 92 (B.M.).
1910: Feb. 10,—1 9.

AUTHADISTIS CAMPTOGRAMMA, sp. n. (PI. I. fig. 44, ¢.)

Antenne of male with fasciculate cilia.

3 9. Head and thorax pale red-brown mixed with blackish
and some whitish; palpi with some black towards base; pectus
whitish; fore tibiz with a black band, the tarsi black ringed with
white; abdomen white tinged with reddish brown. Fore wing
pale red-brown mixed with some whitish and irrorated with dark
brown; subbasal line black, excurved below costa and. ending at
submedian fold; antemedial line black, oblique towards costa,
then erect and very slightly angled inwards at submedian fold ;
a double sinuous blackish medial line, oblique to discal fold, then

MOTHS FROM SOMALILAND. 129

erect; a curved black discoidal striga; postmedial line black,
strongly bent outwards below costa, then exeurved to vein 3 with
a slight inward curve at discal fold, at vein 3 retracted with a
downward curve to lower angle of cell, then erect and sinuous,
some slight blackish marks beyond it on costa; subterminal line
blackish, slightly waved and interrupted, somewhat angled out-
wards elon veins 7 and 4, then incurved; a waved soekicn
terminal line. Hind wing pure white. Underside white, the
costa of both wings slightly irrorated with brown; fore wing
with some black points on terminal part of costa, a minutely
waved black terminal line, the cilia brown at tips; hind wing
with minutely waved black terminal line from apex to vein 2.

Ab. 1. Fore wing with the postmedial line not retracted to
lower angle of cel], but curved downwards between veins 3 and 2,
then erect.

1908: Sept. 3,—1 2; Sept. 17,—1 9; Sept. 18,—1 9 ab. 1
(B.M.); Sept. 19,—1 9; Sept. 23,—1d; Sept. 24,19; Oct. 2,
—1 9; Oct. 3,—1 ¢ (type); Oct. 24-2 g (lin B.M.). 1909:
Mar 0 Oo (BM) Marl); Mar. 29>) 9; Apr 8;
—192. xp. 22 millim.

AUCHENISA CERURODES, sp.n. (PI. I. fig. 43, ¢.)

¢. Head and thorax white mixed with some rufous and black;
antenne rufous ; palpi black towards base; tarsi black ; abdomen
white, dorsally mixed with rufous and black. Fore wing white
slightly irrorated with fuscous brown; black streaks on “sein iL
and inner margin to near middle; an antemedial black patch
on costa with slight sinuous line from its outer edge to submedian
fold; an oblique blackish shade just beyond the cell between
veins 6 and 2; a blackish discoidal spot defined at sides by
white; postmediai line double, the outer line black, the inner
line black at costa, then slight brown and minutely waved,
curved, from costa to vein 2, an oblique wedge-shaped black-
brown patch beyond it on costal area; traces of an oblique
slightly waved brownish subterminal line; a terminal series of
black strie. Hind wing semihyaline white, a black discoidal
lunule and terminal series of small black spots, minute towards
apex. Underside white; both wings with black discoidal spots
and terminal series of striz; fore wing with the costa suffused
with brown expanding towards apex and with white postmedial
mark on it.

1909: Oct. 22,—3 ¢ (including type). Hap. 32 millim.

CATEPHIA PYRAMIDALIS, sp. n. (PI. I. fig. 45, 3.)

3 9. Head and thorax brown mixed with grey-white; palpi
white in front; pectus white; tarsi black ringed with white;
abdomen whitish, dorsally suffused with brown, the crests blackish.
Fore wing grey suffused and irrorated with dark brown; subbasal
line black, sinuous, from costa to vein 1; antemedial line black,
waved; claviform defined by black; orbicular defined by black

Proc. Zoou. Soc.—1916, No. TX. g

130 PROF. E. B. POULTON ON

and with blackish point in centre, round ; reniform with blackish
centre defined on inner side by white and black lines and on
outer side by white, narrow and somewhat produced at lower
extremity ; medial line blackish, oblique to the reniform, oblique
and sinuous below the cell; a triangular whitish shade from post-
medial part of costa to beyond the reniform; postmedial ine
black, strongly bent outwards below costa, then excurved with
a curve inwards at discal] fold, oblique and sinuous below vein 4,
a sinuous dark line beyond it, and some white points on costa ;
subterminal line dark brown, waved, angled outwards at vein 7
and excurved at middle, then incurved; the veins of terminal
area with slight dark streaks ; a terminal series of small black
lunules. Hind wing white, the veins and inner area tinged with
brown, the terminal area broadly fuscous brown; cilia white,
tinged with brown at apex, middle, and tornus. Underside
white, the costal areas irrorated with brown, the terminal area
suffused with brown ; both wings with blackish discoidal lunules
and postmedial line, excurved below costa of fore wing.

Ab. 1. Fore wing with the postmedial triangular patch whiter
and more distinct.

Ab. 2.. Fore wing with black-brown fascia above vein 1 between
the ante- and postmedial lines.

1908: Sept. 21,—1 9; Oct. 30,—1 2. 1909: Mar. 9,—1 9 ab. 2
(B.M.); Mar. 13,—1 9; Mar. 26.—1¢: Mar. 30,—19; Apr. 6,
—1 9: Apr. 8—1 g, 1 2 (¢ is ab. 2, in B.M.); Apr. 10,—1 9:
Ayr. 112 9; Apr: 15,—1 9) Apr. le, Oct. 14 lice
1 2 (ep, wayne, IWS Oem 224 Os Wor OI @ (BNL) 3
Nov. 7,—1 @, ab. 1 (B.M.). 1910: Mar. 6,—192; Mar. 16,
—2 2 (lab.1in B.M.). ap. 24-30 millim.

CAYEPHIA POLIOCHROA, sp.n. (PI. I. fig. 47, 2.)

2. Head, thorax, and abdomen white mixed with brown ;
frons with lateral black bars ; pectus white; legs white tinged
with rufous. Fore wing whitish suffused with brownish grey ;
subbasal line black, from costa to submedian fold; antemedial
line black, oblique and sinuous to submedian fold, then angled
inwards at vein 1, a slight oblique black streak before it above
inner margin; claviform red-brown defined by black and with
black streak from it to the postmedial line, oblique, with its
upper edge extending to median nervure ; orbicular and reniform
defined by blackish except above, rather elliptical, the latter with
some fuscous in its lower part; an oblique blackish shade from
costa to the reniform, and waved Jine from submedian fold to
inner margin; postmedial line black, strongly bent outwards
below costa, incurved at discal fold, angled outwards at veins 4, 3,
then incurved and sinuous; a faint waved whitish subterminal
line with slight blackish streaks before it in the interspaces; a
fine waved black termina] line; cilia brown with a whitish line
at base. Hind wing white, the terminal area fuscous brown,
the inner area tinged with brown; cilia white, the tips brownish

MOTHS FROM SOMALILAND. WIL

at middle. Underside white, the costal and terminal areas
irrorated with brown; both wings with brown discoidal spot and
subterminal shade.

1909: May 12,—1 9 (type). Hap. 40 millim.
CATEPHIA PERICYMA, sp.n. (PI. I. fig. 46, 3.)

3. Head and thorax pale grey mixed with brown, the tegule
with elliptical black-defined annuli; frons with lateral black
bars; tarsi black ringed with white; abdomen brown mixed
with whitish, the ventral surface whitish. Fore wing pale grey
thickly irrorated with brown and black, the veins beyond the
cell with slight dark streaks; a black-brown fascia below base of
submedian fold ; antemedial line black, excurved below the costa
and cell, incurved in the cell and below submedian fold, an
oblique black-brown shade before it on inner area, and a shade
beyond it in submedian interspace to the postmedial line, filling
in the claviform, which is large, defined by blaek, extending to
the cell and acute at extremity; orbicular and reniform large,
defined by black, the former round, the latter elliptical; a slight
oblique brown shade from middle of costa extending into the
reniform; postmedial line black, strongly bent outwards below
costa, then oblique to vein 6, oblique from vein 5 to below 4,
then strongly incurved, dark brown streaks beyond it on veins
3 and 2, and a black streak just below vein 2 with a slight white
mark below it beyond the postmedial line; a waved black
terminal line. Hind wing white, the inner area tinged with
red-brown, the terminal area dark cupreous brown, broad at
costa; narrowing to tornus; its inner edge sinuous; cilia white,
with brown line through them between veins 4 and 2. Underside
white ; fore wing with the costa slightly tinged with purple, a
brown discoidal spot, a subterminal brown shade except towards
costa and inner margin, the area beyond it irrorated with pur-
plish ; hind wing with the costa slightly irrorated with purplish,
a brown subterminal shade except towards tornus with some
brown aud purplish irroration beyond it:

2. Abdomen whiter; fore wing with the brown shade on
basal and median areas more diffused to inner margin; and with
slight brown shade before the postmedial line except towards
éosta:

1909: Mar. 14,—1 ¢,1 9 (types); xp. g 34, 2 40 millim.
CaTEPHIA MESONEPHELE, sp.n. (PI: I. fig: 48, ¢:)

dg. Head and thorax whitish mixed with dark brown; the
tegule except at tips tinged with rufous; frons with black lateral
bars ; palpi with some dark brown at sides of 2nd and 3rd joints;
pectus white; legs tinged with rufous, the tarsi dark brown
ringed with white; abdomen white, dorsally suffused with brown,
the crests dark brown. Fore wing grey-white tinged with brown,
the basal area suffused in parts with dark brown; subbasal line
. black, sinuous, from costa to submedian fold; antemedial line
Q*

132 PROF. E. B. POULTON ON

black defined on inner side by white, excurved below costa and at
middle and more strongly to inner margin, where there is an
oblique black bar before it; the medial area with oblique bright
red-brown fascia from median nervure through the claviform to
the postmedial line at inner margin ; claviform defined by black,
extending to median nervure, oblique and acute at extremity ;
orbicular and reniform with white annuli rather imcompletely
defined by brown, the former with brownish centre, round, the
latter with its centre faintly defined by brown and with small
brown spot in lower part, large, elliptical ; postmedial line black,
slightly bent outwards below costa then slightly sinuous, rather
oblique to vein 4 then incurved, a faint brown line beyond it to
vein 4; an oblique red-brown shade from apex and faint post-
medial line, angled outwards at vein 3; a waved black terminal
line forming points at the interspaces; a fine white line at base
of cilia. Hind wing pure white, the terminal area fuscous brown
from apex to vein 2. Underside of fore wing white, the terminal
area broadly suffused with fuscous.

1908 : Oct. 3,—1 ¢ (type). Hap. 24 millim.

CaTEPHIA EURYMELAS, sp.n. (Pl. I. fig. 49, 3.)

3 2. Head and thorax grey-white mixed with brown, the
tegule with black lines ; frons with lateral black bars ; palpi white
with some black at sides of 2nd and 3rd joints; pectus white ;
legs white tinged with rufous, the tarsi rufous ringed with white ;
abdomen rufous, the crests blackish, the ventral surface white.
Fore wing grey suffused in parts with reddish brown, an oblique
whitish shade from costa towards apex to end of cell ; a subbasal
black striga from costa and oblique streak above vein | ; ante-
medial line double, the outer line black, the inner indistinct,
waved, angled inwards above inner margin ; claviform slightly
defined by black, narrow; orbicular and reniform defined by
black, the former round, the latter incompletely defined on outer
side and with blackish mark in lower part; postmedial line
blackish, bent outwards below costa, then sinuous, oblique to
vein 3, then incurved, some white points beyond it on costa ;
subterminal line reddish brown, diffused on inner side, oblique,
excurved below vein 7 and at middle; a terminal series of black
points. Hind wing white, the terminal area broadly black-brown
from apex to submedian fold, then narrowly black-brown, the
inner margin tinged with brown; cilia white with some black-
brown at veins 2 and 1. Underside white, the terminal areas of
both wings broadly blackish to submedian fold leaving some
whitish on costa and termen of fore wing and at apex of hind
wing.

Ab. 1. Patagia and basal half of fore wing strongly suffused
with black-brown, the latter with the terminal half whiter
slightly tinged with brown and with blackish marks at apex, at
discal fold beyond the postmedial line and at termen and between
terminal parts of veins 3 and 2.

MOTHS FROM SOMALILAND. 133

1908: Sept. 25,192. 1909: Mar. 14,29; Mar. 15,—19;
Mar. 20,—1 9; Apr. 8—1 6 (type), 1 9 (B.M.); Nov.6,—1 9,
ab. 1(B.M.). Hay. 22-26 millim.

Lyncestis ptascora, sp. n. (PI. I. fig. 50, 3.)

3. Head, thorax, and abdomen white mixed with some grey ;
the tegule with black band near tips, the abdomen dorsally
suffused with fuscous from near base to beyond middle; tarsi
black ringed with white. Fore wing white slightly tinged with
grey, a broad oblique fuscous grey shade from costa towards apex
to inner margin beyond middle, a slight black streak below basal
half of costa, and the veins of terminal area streaked with black;
the basal area with grey shades along median nervure and vein 1;
a slight oblique dark antemedial line from cell to inner margin ;
a slight black streak in end of cell; cilia tinged with brown.
Hind wing white, the veins towards termen streaked with black,
the apex tinged with brown; a blackish terminal line. Underside
of fore wing suffused with reddish brown; hind wing with the
costal area irrorated with reddish brown, a subterminal shade
from costa to vein 2.

2. Greyer; hind wing with the terminal area broadly suffused
with black.

1908: Sept. 16,—1 d (type); Sept. 26.—1 $. 1909: Mar. 19,
—1 3; Sept. 20,—1 9 (type). Hap. 28 millim.

LYNCESTIS AMPHIX Cram,
Year ?: Mar. 20,—1 6d.

SPHINGOMORPHA CHLOREA Cram.

Mandera.—1908: Nov. 17,—1 ¢. 1909: Mar. 28,—1 9;
Apr. 5,—1 5; June 6,—1¢,29. 1910: Feb. 9,—1 2.

Gan Libbah.—1908: June 24,14; June 26,—1 9.

Hargaisa.—1909: Nov.,—1<¢.

PASIPEDA SAMBESITA WIk.
1909: Apr. 24.—1d; May 21,—19; July 6.—1<¢.

OcuLAsa CoRNUTA Hmpsn.
1908: Nov. 17,—1¢@. 1909: Mar. 14,—19; Mar. 28,—1 6.

ASPLENIA RUBRESCENS, sp. n. (PI. II. fig. 1, 3.)

3 9. Head and thorax red-brown mixed with some whitish and
a few dark brown scales; pectus and legs whitish, the fore tarsi
ringed with fuscous; abdomen red-brown, the ventral surface
whitish. Fore wing bright rufous with slight dark irroration, a
whitish shade tinged with rufous just beyond the cell; slight sub-
basal blackish points on costa and in the cell; antemedial line
black, waved, double at costa; a small black annulus filled in with
white in the cell towards extremity, and a slight discoidal lunule

134 PROF. FE. B. POULTON ON

defined by blackish; a diffused blackish medial line, excurved
‘beyond the cell, then incurved and slightly waved : postmedial
line indistinctly double, red-brown and blackish filled in with
whitish, slightly bent outwards below costa, then slightly waved
and produced to black and white points on the veins, excurved to
vein 4, then incurved, some pale points beyond it on costa ; sub-
terminal -line whitish defined on inner side by dentate black
marks in the interspaces, excurved below vein 7 and at middle; a
terminal series of black striz; cilia with same white at tips.
Hind wing pale red-brown with a dark subterminal shade; some
dark suffusion on termen towards apex and a dark terminal line ;
cilia white at tips; the underside white tinged with rufous, a
dark discoidal point, eurved red-brown postmedial line with dark
points on the veins and slight subterminal shade.

1909: Mar. 26,—1 3g; Apr. 7,—19; Apr: 8}— 1g); Aprs 9
Weg Ayo WO 1oe Noe Wall Gy (suis) Joes yl ej 9
Apr. 20,—1 9; Apr. 22,—1¢; Apr. 23,—1 9; May 7,—1¢6
(B.M.),12; May 10,—1 4,19; May 12—1¢ (type); May 21,—
19. Hap. 24-28 millim,

TEPHRIAS TRIGONOSEMA, sp.n. (PI. I. fig, 2, @.)

Q. Head and tegule yellow tinged with rufous, the tegule
with a ryfous band behind them; thorax creamy white; antennze
red-byown ; legs slightly tinged with brown; abdomen whitish,
dorsally tinged with brown. Fore wing creamy white slightly
irrorated with red-brown, the cgsta red-brown to beyond middle ;
a conical antemedial chogolate-brown patch from just above
median nervure, its bage extending to the scale-tooth on inner
margin and ouftiyardly resting on vein 1; faint oblique rufous
antemedial and medial strie from costa; postmedial line chogo-
late-brown, arising below gosta, straight to vein 4, then retracted
upwards to ppper angle of cell, then running downwards with a
slight inwards curve to submedian fold jpst beyond the antemedial
patch, and with a trjangular chogolate-brown patch below it
between vein 4 and submedian fold; the area beyond the post-
medial line and below the outer part of the antemedial patch
suffused with red-brown shading to ochreops at termen; sub-
terminal line indistinct, dark brown, oblique, dentate, angled
outwards below veins 7 and 4, Hind wing creamy white suffused
with red-brown especially towards termen; the underside creamy
white irrorated with red-brown, the apical part of terminal area
suffused with brown, a slight discoidal spot and faint curved post-
medial line,

1908; Apr, 27,—1 9 (type). Hap. 22 millim,

PLECOPTERA POLYMORPHA, sp. n. (gi INES tie Sh sy)

3. Head and thorax brownish white slightly irrorated with
fuscous ; antenne brownish; pectus and abdomen white. Fore
wing white, tinged in parts with pale red-brown and irrorated

9

MOTHS FROM SOMALILAND. ES

with black scales ; a slight red-brown subbasal line from costa to
submedian fold; antemedial line red-brown, sinuous; two small
almost conjoined black discoidal spots; postmedial line red-brown
with some diffused blackish at costa, straight and almost erect to
vein 4, then slightly incurved, a series of black points beyond it
from costa to vein 4; traces of a whitish subterminal line, ex-
curved below vein 7 and at middle; the termen and cilia suffused
with red-brown; a fine waved red brown terminal line with minute
dark points at the interspaces. Hind wing white tinged with
red-brown and irrorated with black, the termen and cilia more
strongly tinged with red-brown; a red-brown postmedial line,
axonal bey ond lower angle of gelll ; a waved red-brown ter mdm
line. Underside white faintly tinged with brown and irrorated
with a few black scales; hind wing with black bar on upper
discocellular.

2. Varying from whitish tinged with rufous to purplish grey
suffused with reddish brown, the lines of both wings and discoidal
spots of fore wing often Sind iseinagtt.

Ab. 1. Both wings with strong red- brown shade before the
postmedial line which is defined on outer side by white.

1903: Sept. 12,—2 9 (1 ab. 1 in B.M.); Sept. 22
Sept. 24,—1 ¢ (type). 1909: Mar. 14,—1 9; Apr. 18,—1 9;
Ain 245 OB EME). vAspr a6) 16 OB IM») Apr 27)
Sept.,.—2 5; Oct. 4,—19; Nov. 12,—1 9 (type).

Also in B.M. from Abyssinia, Tamasso (Degen), 1 2. Hap.
30-32 millim.

PLEecoprerRA HYPOXANTHA Hinpsn.
1909: July 6,—2 @.

ACANTHOLIPES CIRCUMDATA Wk.
1909: Oct. 8,—1 9.
ACANTHOLIPES TRIMENI Feld.

1909: May 12,—1@. ,

ANTARCHM#A SUBFLAVALIS W1lk.
1908: Oct. 25,

ANTARCH#A FRAGILIS Butl.
1908: Sept. 14,—-1 9; Nov. 18,—1 2. 1909: Oct. 14,—1¢9.

TATHORHYNCHUS EXSICCATA Led.
1909: May 10,—1 ¢.

ANOMIS FIMBRIAGO Steph., or EROosA Hiibn.

1909: Apr. 6,—1 9. ‘The females of fimbriago and erosa
aE Ure
cannot be distinguished with certamty.

136 PROF. E. B. POULTON ON

CALPE VAGABUNDA Swinh.

Mandera.—1908: May 29,—1 9; Sept. 16—19. 1909:
Mar. 12,—1 6; Apr. 15,—192; Sept. 9,19.

Hargaisa.—1908: Oct.,—1¢ (B.M.).

ARGADESA MATERNA L.

1909: Apr. 6,—1¢3,19; May 21,—1¢.

OPHIDERES FULLONICA L.

Hargaisa,—1909: July,—1d,19.
This species and the last were often seen at light, but were not
sufficiently attracted to be easily captured.

Subfam. Hypunina.
SARMATIA INTERITALIS Guen.
1908: Sept, 26,—1¢. 1909: Apr, 22-19; May 10,—19.
SIMPLICIA CAPALIS Wlk.
1908: June 1,—1 9,

NopaRIA EXTERNALIS F,
1908: June 2,—1 ¢. 1909: Jan. 16,—1 ¢.

HYPENA sTRIGATA F, (ABYSSINIALIS Guen. ),

1909: Apr. 6;—1¢; May 21—1¢4; July 6—16.
Hypena gussauis Wlk.

1909: May 26,—19. 1910: Mar. 16,—1 9.
HYPENA MASURIALIS Guen.

1909: Apr. 6,—1 9; Nov. 10,—1¢.

RAYNCHINA ANTIQUALIS Hibn,

1909: Qot, 22,—1 9.

RHAYNCHINA PERANGULATA, sp. n. (PI. II. fig. 7, 2.)

3 2. Head, thorax,and abdomen grey-white mixed with reddish
brown; palpi tinged with red-brown and irrorated with black ;
ventral surface of abdomen white irrorated with brown. Fore
wing grey tinged with red-brown and irrorated with black; ante-
medial line white, very oblique from costa to submedian fold,
where there is a small fan of raised scales below its extremity ; an
elliptical red-brown spot in end of cell with white streak below it
on median nervure and small white patch beyond its lower ex-
tremity ; postmedial line fine, blackish, defined on inner side by
white and on outer side also towards costa, very oblique to discal
fold where it is acutely angled, then oblique to inner margin
below the antemedial line, some white points beyond it on costa
and an oblique white shade from apex to its angle; a slight

MOTHS FROM SOMALILAND. 137

dentate white subterminal line with oblique chocolate-brown shade
beyond it from just below apex, then a series of dentate chocolate-
brown marks on its outer edge ; a fine chocolate-brown terminal
line and white line at base of cilia. Hind wing reddish brown ;
a fine dark terminal line; cilia paler; the underside whitish
tinged and irrorated with brown, a brown discoidal point and
curved postmedial line.

1909: Apr. 7,—1 @ (type).

Also in B.M. from Br. KE. Africa, Taveta ing 23; Mosam-
bique, 1 ¢; Transvaal, Kranspr uit (Janse), 1 2. Exp. 20-26
millim.

RHYNCHINA REVOLUTALIS Zell.
1908: Aug. 24,—1 9.

RHYNCHINA ALBISCRIPTA, Sp.n. (PI. II. fig. 8, 3.)

Antenne of male minutely serrate, with fasciculate cilia.

3S. Head and thorax white irrorated with brown ; palpi with
the 2nd joint fuscous brown except below, the 38rd with fuscous
band ; abdomen white, dorsally tinged with brown. Fore wing
white suffused with brown, the inner area to the postmedial line
and the termen whiter; antemedial line dark brown defined on
each side by white, oblique to submedian fold, then inwardly
oblique ; a minute blackish annulus in the cell towards extremity ;
a slight white discoidal lunule defined by dark brown; postmedial
line black-brown defined on each side by white, oblique to vein 6
and below vein 4; some white points beyond it on costa; a sub-
terminal series of minute white spots in the interspaces, defined
on inner side by slight somewhat dentate black marks with den-
tate white marks before them towards costa; a fine blackish
terminal line; cilia with white lines at base and near tips. Hind
wing white tinged with reddish brown; a fine brown terminal
line; cilia white at tips; the underside white, the costal area and
terminal area to vein 3 irrorated with red-brown, a faint curved
postmedial line.

1908: Sept. 19,—1 ¢ (type); Oct. 1,—1 ¢.

Also in B.M. from Sudan, Port Sudan (W/rs. Waterfield), 3 3.
Hep. 20 millim.

RHYNCHINA ENDOLEUCA, sp. n. (FI. II. fig. 6, 3.)

Antenne of male bipectinate with short fasciculate branches,
the apical part ciliated.

3. Head, thorax, and abdomen grey-white tinged with brown ;
the crest at base of abdomen fuscous. Fore wing grey-white
tinged with brown; a slight white streak in basal half of sub-
median fold, the area below it paler and tinged with red-brown ;
antemedial line represented by a striga of raised blackish scales
from costa, some black scales on inner area and a small spot
further from the base below the cell; a point of raised black
scales in the cell towards extremity and a bar from origin of

138 PROF. E. B. POULTON ON

vein 2 to inner margin with a slight rufous line before it; a slight
brown line from above end of cell to vein 6, dentate at veins 7
and 6 ; some minute blackish streaks on costa towaids apex and
an oblique whitish shade from apex to end of cell with a rufous
tinge on its outer side and short black streaks in the interspaces,
ending in an oblique black bar above vein 2 just beyond the bar
below the cell ; a subterminal series of minute black strie, oblique
from below vein 3 to submedian fold, then erect, with a slight
streik beyond it in submedian fold; a slight sinuous blackish
terminal line and fine white line at base of cilia which are inter-
sected with black at the veins. Hind wing whitish tinged with
red-brown; a fine brown terminal line; cilia whiter, slightly
intersected with brown at the veins; the underside white tinged
with rufous and irrorated with brown except on basal and inner
areas, a brown discoidal point, traces of a postmedial line from
costa to discal fold and a punctiform black terminal line.

Q. Fore wing with the basal half suffused with fuscous brown
to subimedian fold in which the white streak is stronger and the
inner area more contrasting, the white shade from apex more
prominent.

Ab. 1 much darker.

1908: Sept. 20,—1 9 (type); Sept. 27,1 ¢ (type).

Also in B.M. from Sudan, Port Sudan (Mrs. Waterfield), 1 3,
1°; Br. HE, Africa, Sabaki R. (Gregory), 1 9 ; Kitu (Crawshay),
19; Takaunga (7. Thomas), 19; Munisu (Lord Delamere), 1°.
Eaup. 3 20, 9 24 millim.

MAGULABA GRISEA, sp.n. (PI. II. fig. 4, 3.)

3 9. Head and thorax black-brown mixed with reddish brown ;
ectus, legs, and abdomen grey, irrorated with dark brown, the
palpi and fore legs suffused with black, the tarsi ringed with
whitish. Fore wing grey-white suffused with reddish brown and
irrorated with blackish ; a sinuous blackish antemedial line; a
white point in middle of cell and slight whitish discoidal striga
placed on a sinuous blackish medial shade, incurved below the
cell; an indistinct blackish postmedial line, excurved below costa
and at middle, incurved at discal fold and below vein 4; a brown
subterminal shade with series of more or less prominent black
marks on it, excurved below vein 7 and at middle; a terminal
series of black points. Hind wing grey suffused with brown and
irrorated with dark brown; traces of two postmedial iines with
the area between them rather paler; a terminal series of fuscous
striz ; the underside white irrorated with brown, a dark discoidal
striga and rather diffused brown postmedial and subterminal
lines.

1908: Feb. 24,—1 ¢ (B.M.); Sept. 13,—1 6; Sept. 22,16;
Oct. 3,—1 3 ; Oct. 29,—1 6; Nov. 13,—1 ¢ (type). 1909:
Apr. 20,—1 3 3 Apr. 22,—1 9 (B.M.).

Also in B.M. from 8. Nigeria, Sapele (Sampson), 13. Lup.

22 millim.

MOTHS FROM SOMALILAND. 139

NAARDA NIGRIPALPIS, sp.n. (Pl. II. fig. 5, 3.)

3S. Head and thorax brown mixed with grey; antenne blackish
slightly ringed with grey; palpi blackish; fore legs black, the
tarsi slightly ringed with white ; abdomen grey suffused with
brown. Fore wing grey thickly wrorated with brown; antemedial
line dark brown and sinuous; a minute ochreous spot in middle
of cell and ochreous discoidal bar defined at sides by dark brown ;
a rather diffused erect brown medial line ; postmedial line dark
brown, sinuous, slightly incurved below vein 4; subtermimal line
whitish defined on inuer side by brown, shghtly sinuous ; a ter-
minal series of dark brown strie. Hind wing grey suffused with
brown; a slight brown discoidal lar and some dark scales at
middle of inner margin; a curved waved brown postmedial line ;
a rather diffused waved subterminal line; a terminal series of
dark brown striz; the underside whitish irrorated with brown,
the discoidal bar and postmedial and subterminal lines more
distinct.

1908: Oct. 25,—1 ¢ (type). Hap, 18 millim.

Fam. LYMANTRID&.

Euproctis FAScIATA W1k.
1909: Mar. 14,—1 g¢. 1910: Mar, 10,—1! ¢.

L2&LIA TESTACEA W1k.

Mandera.—1909: Oct. 6,—1 9.

Hargaisa.—1908: Oct.,—2 ¢.

CasaMA Vitis Wlk.

1908: May 2,—1¢,19; May 4,—1¢,19@; Aug. 15,—l1¢,
19; Aug. 24,1 ¢; Aug. 25,—19; Aug. 26,—1 4,1 9; Sept. 3,
13; Sept. 15,—1 ¢; Sept. 18.— 29; Sept. 19,—19; Sept. 21,—
i eo Septs 22, ge LO Ochy se OCU Elis
Oct. 15,—19. 1909: Jan. 13,—19; Jan. 18,—19; Feb. 17,—
ae Heb. 25 "1O- Mar. 12-135 5) Mar 13) 2) Mare 1)
19; Mar. 24,—1¢; Apr. 6,—1 ¢; Apr. 7,—36; Apr. 8,—_l1¢;
Apr. 11,—19; Apr. 14,16; Apr. 24,—1¢ ; Apr. 26,—1¢ ;
Apr. 30,—1¢; Aug. 17,—1 ¢; Oct. 5,—1 ¢; Oct, 7,-1 3;
Nov. 25,—1 92. 1910: Jan. 8,—1¢.

DASYCHIRA MISERATA Holl,

1903: Aug. 26,—1 <.

Dasycuira REMOTA Druce,

Year?: Mar. 24,—] 9.

ACLONOPHLEBIA INconspPicya, sp. H. (PI. II. fig. 9, 3.)

¢. Head, thorax, and abdomen dark red-brown mixed with
some whitish. Fore wing whitish tinged with red-brown and
thickly irrorated with dark brown, the veins with dark streaks ;

140 PROF. E. B. POULTON ON

a dark brown subbasal patch from costa to below the cell; ante-
medial line blackish, excurved from below costa to submedian
fold where it is slightly angled inwards ; traces of a sinuous dark
medial line; a curved black discoidal striga; postmedial line
blackish slightly defined on outer side by white, somewhat
dentate and produced to slight streaks at veins 7 to 2, bent
outwards between veins 5 and 3, retracted below vein 2, and
slightly angled outwards at submedian fold and vein 1; some
slight whitish marks on costa towards apex; cilia brown, inter-
sected with white at veins 7 to 2. Hind wing white, somewhat
semihyaline, the veins shghtly tinged with brown; the cilia
with slight brown spots from apex to vein 2. Underside of
fore wing with the terminal area white with a subterminal
brown shade rather diffused on inner side and dentate on outer
between veins 7 and 3.

Hargaisa.—1$08: Oct.,—2 ¢ (including type). Hap. 30 millim.

Fam. SPHINGIDA.
By Dr. Karu Jorpan.

Herse convonvut L.
Sphinx convolvuli Linné, Syst. Nat. ed. 10, p. 490. n. 6 (1758).
1909: July 6,—1 9; Nov.6,—1¢.

Powiana Micra R. & J. (1903).

Poliana micra Rothschild & Jordan, Nov. Zool. ix. Suppl.
p- 809, no. 766, text-fig. 6 (1903 : Somaliland).

These two males from Mandera are in a better state of
preservation than the type, and therefore appear purer grey
on the fore wing. The only difference I notice is in the ante-
and postmedial double lines being less filled-in with fuscous and
a little further apart below the apex of the cell than in the type
specimen, the only example hitherto known to us of this species.
The genitalia of one of Mr. Feather’s specimens have been
examined ; they are identical with those of the type.

1908: Oct. 31,—1 ¢. 1909: May 13,—1 ¢.

Hipporion cELERIO L.
Sphinx celerio Linné, Syst. Nat. ed. 10, p. 491. n. 10 (1758).

Mandera.—1909: Apr. 5,—1 ¢.
Berbera.—1908: Dec. 2,—1 9.

HiPPOTION ROSEIPENNIS SOMALICUM, subsp. n.

3 Q. Ab H. ros. roseipenni ala antica in disco lineis quinque
fuscis notata distinguendum.

In true roseipennis Butl. (1882), which is known to us from
Delagoa Bay northward to British Hast Africa and Unyoro, the
fore wing bears two distinct lines in the outer half, the proximal

MOTHS FROM SOMALILAND. 14]

line crossing the fuscous patch which is placed at the apex of the
cell, and the outer line being nearly continuous with the oblique
apical streak. In between these two lines there are at most
faint traces of two or three other lines paraliel with them. In
somalicum, on the other hand, the three additional lines are
quite distinct, the first and second additional lines being in
the male nearly, and in the female fully, as well marked as the
proximal line. In this character H. r. somalicum represents
an ancestral stage.

The genitalia do not appear to differ from those of HZ. r.
roseipenies.

Mandera.—1909: Oct. 14,—1 9.

Hargaisa.—1909: Nov.,—1 ¢ (type).

Hiprorton ros# Butl.
Darapsa rose Butl. A. M.N. H. (5) x. p. 433. n. 5 (1882).
1909: Oct. 6,—I1 ¢.

Fam. NoTODONTID &.

xARGETTA XYLOCHROA Hmpsn.

Mandera.—1908: May 29,
Feb. 17,—1 ¢.
Hargaisa.—_1908 : Oct..—1 ¢.

SCRANCIA DISCoMMA, sp.n. (PI. II. fig. 10, 2.)

@. Head and thorax white mixed with reddish brown and
blackish, the patagia white slightly pencilled with brown; pectus
and legs white with a few brown scales; abdomen white tinged
with brown and with slight lateral blackish spots except towards
extremity. Fore wing white irrorated with a few black scales,
the inner half tinged with red-brown, the veins streaked with
black except on basal and inner areas and at costa ; a small round
black-brown discoidal spot surrounded by white. Hind wing
white, the terminal area tinged with brown, broadly at costa,
narrowing to tornus; cilia white, faintly tinged with brown.
Underside of fore wing suffused with brown, the terminal area
whiter; hind wing with the costal area suffused with biown.:

1908: Oct. 15,—1 9 (type). xp. 35 millim.

192; Nov. 22,—1 6. 1909:

STENOSTAURA IMPEDITUS WIk.
1808: Feb. 24,—1 ©.

Fam. GEOMETRID &.
By Louis B. Prout.

These form, on account of the number of new and interesting
species, an extremely important part of Mr. Feathev’s collection.
Their general affinities, as might be expected, are with the fauna
of Abyssinia, British East Africa, and to some extent Socotra

142 PROF. E. B. POULTON ON

and Southern Arabia, and desért forms are strongly in evidence.
Nearly all the species are of small sizé, the majority belong to a
few groups (espécially Acidaliids and the Macaria group), and one
extensive subfamily (the Larentiine) is almost absent, being re-
presented by only two specimens, while even of these one is the
somewhat anomalous Pséudosterrha phileéaria. The presence, in
the groups named, of a number of closely allied and inconspicuous
Species (In some cases also strongly variable) has rendered a
satisfactory working-out of the material a matter of no small
difficulty ; and this” difficulty has been increased by a curious
and unexplained circumstance which deserves mention—the
very marked preponderance of females, this sex alone being
represented in not a few cases where ‘there is quite a good
series of examples. We are not unaccustomed to meeting wiih
Geometrid collections in which the niales alone of many species
are present, and are able readily to attribute this to the greater
activity of the sex, the fact that the collection was made chiefly
at light, and so on; but it is less easy to suggest what difference
in The tis or what particular method of collecting, has resulted in
the capture of the females only of so many spécies. That the
phenomenon is not confined to a sitgle genus or group will be
seen by referring to the details given “yellow, under Mierochthonia
featheri, Acidatiastis subbr “UNnNESCENS, fephrina, nearly the whole
of the Acidaline’, etc.*

Subfam: HemiriEer ».

Ten species are tepresented, most of them more or less highly
specialised forms, and including two additions to the handful of
known species in which the Gharacter istic green colour of the
subfamily has given place to some shade of brown or sand-
colour.

VICTORIA SEMATOPERAS, sp. n. (PI. II. fig. 26, ¢.)
3 2, 32-33 mm. Face and upper side of palpus dark red;

crown of head, base of antenna, and basal one-third or more of
costa red mixed with lustrous blue-blackish scales. Abdomen
dorsally slightly reddish, crests lustrous, pale on summit, then
reddish, a deep black spot (dot) near base of each. Fore wing
with termen almost smooth; green (in all three discoloured by
relaxing) ; discal dot white, eneieled with a black-dusted red
ring; distal margin with similarly coloured dark spots, namely a
small one in front of R',a much larger one from R' to beyond R?,
and a small or moderately large one at tornus. Hind wing with

(=)
the excision between the “aca not deep, discal dot as on fore

* (After the above paragraph was written a number of additional specimens of
Geometridge were set and added to the collection. Mr. Prout wrote (Feb. 19, 1915),
concerning these additions:—‘*They do not upset my geueralisation as to the
preponderance of females; indeed, they rather strenethen it, being almost ex-
clusively of that sex except in one species (Heterostegane indularia) whose males
were already well in evidence.” On this subject see also p. 93.—E. B. P. |

MOTHS FROM SOMALILAND: 143

wing or less clearly ringed, tornus with indications of dark
markings. Fore wing beneath with the terminal markings
weakly indicated, hind wing quite unmarked.
1909: Mar. 14,—1 92 ; Dec. 30;—1 G (type): 1910: Jan. 2,
i on
Unfortunately all have lost the hind legs. It is evidently
a near ally of V. triplaga Prout, from German Hast Africa.

PRASINOCYMA PERPULVERATA, sp: n. (PI. II. fig. 25, 3.)

3 9, 16-21 mm. Palpus in male rather slight for the genus,
in female about 13 times diameter of eye, with 3rd joimt exposed
buat rather short for the genus. Antenna of male with the outer
pectinations longish, the inner much shorter. Hind tibia of male
not dilated, the four spurs closely approximated. Head, body;
and legs concolorous with wings, the vertex appreciably paler
than the face. Wings rather narrower than in typical Prasino-
cyma, hind wing not at all bent at R’, cells relatively long (fully
one-half); fore wing with SC’ generally free, R' not or very
shortly stalked, M* connate or very shortly stalked, hind wing
with two stalkings, White-grey with a slight tinge of brown;
coarsely irrorated with fuscous, the irroration under a lens
inclining to resolve itself into minute longitudinal strigule ;
cell-spots strong, elongate. Under surface less strongly irrorated:

Apparently variable, the name-typical. form, with uniform
irroration, the commonest; here I refer the following :—

Mandera.—1908: June 1,--1 2; July 17,—1 2; Sept. 27;
—1 5,1 92. 1909: May 10,—2 9,1 ¢ (type).

Three females show on the fore wing an ill-defined dark basal
patch, dark median band from hind margin about to cell, and
dark terminal dashes between the veins: ab. subfasciata, ab. n.

Mandera.—-1909: May 10,—1 2. 1910: Mar.,—1 @ (type):

Hargaisa.—1908: Oct.,—1 9.

One male is smaller (16 mm.) and with still more markings;
the median area being broadly dark-mixed in anterior half,
a dark proximal shading (forming a large, strong spot at costa)
indicating the position of the obsolete subterminal line; the hind
wing somewhat shorter, with distal area somewhat darkened,
the antennal pectinations apparently continuing slightly less
far down the shaft : ab. perScripta, ab. n. (? sp: div.):

Mandera.—1908: July 17,—1 ¢ (type).

By the length of the cells and the female palpus, as well as by the

facies (which recalls Veromia pulvereisparsa Hmpsn.), perpulverata
should probably be made the type of a new genus.

CHLORISSA STIBOLEPIDA (Btlr.) *.
Comibena stibolepida Btlr. Cist. Ent. ii. p. 394 (1879).

* {The parentheses around the names of authors placed after scientific names in
this paper are used in accordance with Article 23 of the International Rules of
Nomenclature (Proc. 7th Int. Cong., Boston 1907, p. 44 (1912))—Eprror. |

144 PROF. E. B. POULTON ON

Hemithea albistrigulata Warr. Nov. Zool. iv. p. 39 (1897).

Hemithea vermiculata Warr. ibid. p. 41 (1897) (n. syn.).

1909: Apr. 20,—1 @; Apr. 24,—1 9.

A very widely distributed African species, and perhaps not
structurally differentiable from C. faustinata Mill. (S. Palearctic)
and ©. solidaria Guen. (Indian).

NEROMIA MALESCRIPTA (Wartr.).

Hemithea malescripia Warr. Nov. Zool. iv. p. 40 (1897).

1908 :- Nov. 13,—1 ¢. 1909: Mar. 1,—1 ¢.

Both examples small, with the crests red.

Distributed in Abyssinia, British East Africa, Transvaal, and
Natal. I have a note, dealing with the difficult group to which
this species belongs, in the press for the ‘ Annals of the Transvaal
Museum.’

N®ROMIA MANDERENSIS, sp. n. (PI. II. fig. 24, Q.)

, 20-22 mm. Face and palpus red. Vertex and antenna
whitish, the latter with minute ciliation ; occiput green. Thorax
and abdomen concolorous with wings. Wings above green with
whitish strigulation, quite like the greenest forms of Chlorissa
stibolepida Btlr., costal edge of fore wing ochreous, otherwise
markingless ; fringes green, lighter distally. Under surface paler
green, costal edge of fore wing as above.

Mandera.—1908: Sept. 25-1 @ (type); Nov. 13,—1 9.
1909: May 8,—1 9.

Hargaisa.—1908 : Oct.,—1 2 (a worn example).

Hind wing less elongate than in Chlorissa stibolepida, termen
smoothly rounded; but best distinguished by the structure.
* Palpus reaching beyond frons and shortly rough-scaled, but
with 3rd joint small; hind tibia with terminal spurs only.
If the male antenna should prove to be pectinate, the species
should be considered a Microlovia, aberrant in the rather short
3rd joint of the palpus.

Genus Hemipromopes, nov.

Palpus slender, in male rather short, in female moderate,
in both sexes with 3rd joint short or shortish. Tongue absent.
Antenna short and rather thick, in male with moderate, in
female with short pectinations. Pectus somewhat hairy ; hind
tibia in male short, greatly dilated (recalling that of Synclysmus),
all the spurs present, terminal very short, the outer almost
obsolete ; in female with terminal spurs well developed, median
short, sometimes entirely absent. Abdomen not crested, in
female robust. Frenulum in male slight, in female absent.
Fore wing with SC! from cell, free, R' about connate with SC. a,
R* rather far forward, M' about connate or very shortly stalked
with R*. Hind wing with termen smooth, C anastomosing to

MOTHS FROM SOMALILAND. 145

scarcely one-half cell, DC not very oblique, SC° well stalked,
R? little before middle of discocellulars, M! well stalled.

Type of the genus: Hemidromodes robusta Prout (Hiero-
chthonia).

A connecting link between Hierochthonia and Syndromodes ;
the absence of the male, and the curious fact that the female
examined had no trace of median spurs (though both legs are in
good condition), led me to refer the species to the former genus.
From Syndromodes it differs in antenna, male hind leg, robust
female abdomen, etc.

HEMIDROMODES ROBUSTA (Prout).
Hierochthonia robusta Prout, Nov. Zool. xx. p. 435 (19138).

6g, 14-18 mm.; 6 9, 18-24 mm.

1908: June l,—2 3g; July 17,—1 3g; July 31,—1 @; Aug. 24,
—1$. 1909: Jan. 16,—1 9*; Mar.19,—1 9; Mar. 24,—1 9;
Apr. 11,—1 3; May 10,—1 6; May 21,—1 9; Nov. 7,—1 9*.

Excepting the two females marked *, all are smaller—mostly
considerably smaller—than the type specimens from Port Sudan.

In this species the fringes (which in neither of the originals
were quite perfect) are long, proximally green, distally ochreous
whitish.

HiIEROCHTHONIA FEATHER, sp. n. (Pl. IT. fig. 23, 9.)

2, 24-26mm. Face green. Palpus minute, whitish. Tongue
vestigial. Antennal shaft white, pectinations long. Vertex
white; occiput green. ‘Thorax above green, beneath white.
Abdomen robust, dorsally green, becoming white posteriorly
and ventrally. Fore wing moderately broad, SC’ from cell,
anastomosing with C, SC* from shortly after R’, anastomosing
with SC’, R* well stalked, R® rather extremely placed, M* shortly
stalked ; uniform bright green, nearly as the genus Hwchloris
or slightly more bluish, distal one-third of fringe white. Hind
wing moderately broad, costal margin rather long, apex rounded,
C anastomosing to near end of cell, R* rather extremely placed,
M' short-stalked ; green, rather paler than fore wing, especially
towards base and costal margin. Under surface pale green.

1908: Nov. 20,—1 @. 1909: Jan. 15,—1 9; Jan. 16,—1 9;
ei, Zeal 2s egos, UO I a oso Il Os Aare sy) Oe
Mipr 22) 2 Mia ol OV (Gy pe); Decwli = S=ior

Probably related to petitaria Chr., notwithstanding the broader
wings and strongly pectinate antenna. It is unfortunate that
the males in this group are still unknown.

CoMOSTOLOPSIS STILLATA (Held.).

Nemoria stillata Feld. Reise Novara, Lep. Het. t. 127. lies, 17)
(1875).

Eucrostes rubristicta Warr. Nov. Zool. vi. p. 23 (1899).
Proc. Zoon. Soc.—1916, No. X. 10

146 PROF. E. B. POULTON ON

Eucrostis rufostellata Mab. Ann. Soc. Ent. Fr. Ixvii, p. 740
(1900).

1909: May 21,—1 @.

A quite typical example of this widely distributed African
species, extending its known range.

EKUCROSTES ASTIGMATICA, sp.n. (PI. IT. fig. 22, 9.)

S, 15-16 mm.; 2,19-20 mm. Superficially very like pygmea
Rbl.* (=inselaris Prout), but larger, of a still more vivid green,
and without the discal dots, the pale postmedian line not
discernible, or only suggested in certain hghts. “‘Snow-white,”
in my description (Gen. Ins. exxix. p. 246) of the costal edge,
was not absolutely accurate, as there is, in a good light, a very
delicate tinge of pinkish or violet in the white in both species.
Structurally like the African members of the genus, the male
palpus being less minute than in pygmea, the 3rd joint in the
female rather less long and slender, and the female antenna
bipectinate, the longest branches about twice as long as the
diameter of the shaft.

1908: Sept. 21—1 g. 1909: Apr. 21,—l1 ¢ (type);
Apr. 22,1 23 May 10;—2 2-

ACIDALIASTIS SUBBRUNNESCENS, Sp. 0.

2, 14-16 mm. Much more strongly and uniformly dusted
with sand-colour (sometimes more yellowish, sometimes more
brownish) than miera Hmpsn., the dark lines only weakly (in
the darkest example scarcely at all) indicated, but with very
conspicuous white lines proximally to the first and distally
to the second; the former of these is more oblique than in
micra, not reaching costa, the latter forms a rather more
appreciable curve than in that species; discal dot absent.
Hind wing white, becoming more or less tinged with sand-
colour towards termen, in the darkest-marked example showing
a curved white postmedian line. Fore wing beneath with white
outer line and white hind margin, hind wing all white.

1908: June 1,—-1 9; June 29,—1 9; Sept. 14,1 92 (type);
Sept. 17,—1 @; Sept. 23,—1 9. 1909: Mar. 24,—1 Q.

Very near bicurvifera Prout (Ann. Transv. Mus., in the press),
much smaller, costa rather straighter, apex rather less sharp,
termen rather less convex in posterior half, palpus and female
antennal pectinations slightly shorter, markings less reddish
(more olivaceous), postmedian line almost parallel with termen
(in bicurvifera more oblique), hind wing rather shorter, less
unicolorous, face apparently Jess reddish.

* Denks. Akad. Wien, Math.-nat. Kl. Ixxi. 2, Sep. p. 67 (1907). As only
separata in advance seem to have been issued of the paper of Rebel’s containing
this species, and it was not noticed in the ‘Zoological Record, I do not feel to
blame for haying overlooked it in the ‘Genera Insectorum’ and created a
synonym.

MOTHS FROM SOMALILAND. 147

Subfam, ACIDALIIN &.

Genus TRICENTROSCELIS, nov.

Face rounded, markedly prominent, with appressed scales.
Palpus short, shortly rough-scaled. Tongue present. Antenna
in female minutely ciliated. Pectus and femora glabrous. Hind
tibia in female with a single proximal and a pair of terminal
spurs, all of moderate length. Wing-shape and facies of
Acidalia, distal margins smooth. Fore wing with SC* from
cell, anastomosing with SC! and then very strongly with SC**
(i. e. areole double and SC* and SC? arising before and behind the
apex of the distal areole), M’* well separate from R*. Hind wing
with C normal, SC* very shortly stalked or almost connate with
R', M* well separate from R’.

Type of the genus: T'ricentroscelis protrusifrons, sp. 0.

Differs from the Neotropical Scelolophia Hulst (= Calyptocome
Warr. = Crypsitila Warr.), which also has often a 3-spurred
female hind tibia, in the protuberant face and longer cells.

TRICENTROSCELIS PROTRUSIFRONS, sp. n. (PI. II. fig. 21, 9.)

2,21 mm. Face fuscous. Palpus fuscous, not quite reaching
extremity of frons. Vertex and antenna similarly coloured to
wings, but rather paler; antennal ciliation minute. Thorax
concolorous with wings. Abdomen slightly paler, 2nd—4th
segments mostly occupied dorsally by a large fuscous blotch, the
later segments interruptedly marked with fuscous dorsally. Fore
wing with apex and termen somewhat rounded ; reddish brown
(light grey-brown irrorated with rufous and blackish); ante-
median line indicated by rufous and blackish scales, accentuated
by black spots on costa and hind margin and dots on the veins,
arising before one-third costa, bent in cell, becoming oblique
inwards and sinuous ; discal dot small but sharp ; median shade
obsolescent, placed between discal dot and postmedian line,
slightly more distinct as a costal dot, outbent at radials, inbent
at fold; postmedian from costa slightly beyond two-thirds, faint
except at costa and veins, where it is marked by black dots,
angled at SC’, then strongly oblique outwards to R', then more
parallel with termen, but slightly sinuate inwards at R? and
more strongly at fold; terminal dots strong, black; fringe
somewhat dusted with blackish, a slender clear line at base.
Hind wing with termen almost smooth, very slightly sinuous
towards anal angle; concolorous with fore wing, discal dot
slightly larger, median shade much stronger, nearly straight,
proximal to the discal dot, antemedian wanting, the rest as
on fore wing. Under surface slightly paler, the discal and
terminal dots and postmedian line reproduced, though less
sharp; hind wing in addition with median shade indicated
at abdominal margin.

1909: Apr. 6,—1 @ (type).

Superficially similar to some African Acidalia (ewiguaria group)
or Ptychopoda (sinwilinea Prout, ete.)

Os

148 PROF. E. B. POULTON ON

TRAMINDA RUFISTRIGATA (Hmpsn.).

Ephyra rufistrigata Hmpsn. Proc. Zool. Soc. Lond. 1896,
Do AGT, jal 2k whee

1908: Sept. 28,—1 9. 1909: Apr. 14,—1 9.

Described from Aden; distributed as far as British East
Africa, As the male hind tibia has all spurs present, the
species must be removed to Tranunda.

TRAMINDA NEPTUNARIA (Guen.).

Timandra neptunaria Guen. Spec. Gén. Lép. x. p. 3, t. 18. fig. 5
(1858).

Timandra viridaria Walk. List Lep. Ins. Brit. Mus. xxii.
p- 800 (1861). ;

Gnamptoloma neptunaria Warr. Nov. Zool. it. p. 95 (1895).

Traminda neptunaria Swinh. Tr. Ent. Soc. Lond. 1904, p. 562.

Mandera.—1909: Apr. 5,—1 ¢; Apr. 27,—1 @.

Hargaisa.—_1908 : Oct.._-1 ¢.

Widely distributed in Africa.

CHLORERYTHRA RUBRIPLAGA Watr.

Chlorerythra rubriplaga Warr. Nov. Zool. 1. p. 91 (1895).

1908 : Sept. 19,—1 9; Oct. 25,—1 9. 1909: Feb. 15,19;
Feb. 22,—1 @; Mar. 14,—1 3; May 10,—1 2. 1910: Jan.,—
I es

The male and two females (22 Feb. and 10 May) belong to the
plain green form with the oblique red line almost entirely
obsolete; the other four females have the line distinct, the
eround-colour showing the three gradations well known in this
group (green, green irrorated with rufous, rufeseent). Doubt-
fully distinct generically from Vraminda. Widely distributed
in Eastern Africa.

ACIDALIA MINOA, sp.n. (PI. I. fig. 20, 2 .)

2,20 mm. Unfortunately (like so many of the species)
without the male, but showing sufficient peculiarities to render
it safe to describe it. Absolutely without markings, very glossy,
otherwise bearing a good deal of superficial resemblance to a worn
female of Minoa murinata Scop., though with narrower wings ;
similarly coloured; the fore wing beneath with a smoky suffusion,
which is also slightly indieated on the upper surface at certain
angles of light. Head and body slightly more ochreous than
wings, the face and palpus sharing this colour, not—as in the
majority of Acidalia—black or fuscous. Abdomen rather robust.
Venation rather variable, SC’ of fore wing arising from just
before apex of areole or well stalked with the other subcostals ;
C of hind wing not rapidly diverging from SC, sometimes anas-
tomosing at slightly more than a point; SC* in two of the
examples extremely shortly stalked with R'. Termen of hind
wing not at all bent in middle.

MOTHS FROM SOMALILAND. 149

1909: Apr. 7,—1 2 (type); Apr. 9,—19: Apr. 23,—19.
I suspect this may prove to belong to the less specialized
section Pylarge (male hind tibia with terminal spurs).

ACIDALIA MINORATA (Bsd.).

Geometra (Idea) minorata Bsd. Nouv. Ann. Mus. Hist. Nat. 11.
p- 263 (1833).

? Acidalia remotata Guen. Spec. Gén. Lép. ix. p. 458 (1858).

Acidalia consentanea Walk. List Lep. Ins. Brit. Mus. xxi.
p- 745 (1861).

? Acidalin actuaria Walk. ibid. p. 752 (1861).

? Acidalia derusata Walk. ibid. xxvi. p. 1604 (1862).

1908: July 17,—1 2; Sept. 11,—2 2; Sept. 13,—1 @; Sept.
25,—1 9; Sept. 30,—1 9; Oct. 11,—192; Oct. 20,—19; Nov.
24-19. 1909: Jan. 12,—29; Oct. 20,—29; Oct. 29,—1 9;
Nov. 16,—1 2 °

All fifteen females referable, so far as present knowledge is
available, to this very common, very widely distributed, and
moderately variable species. On an average, the lines are less
crenulate and more concise than in the most typical forms, and
as there is some slight variation in the breadth of the wings, as
well as in the tone of colour and strength of markings, I am not
prepared to say that there may not be two or three species mixed.
One or two examples, in the sparseness of their irroration, recall
lactaria Walk. (List Lep. Ins. xxii. p. 744), which, however, is
possibly also only an aberrant form of minorata. Distributed
throughout Africa, except, perhaps, the extreme north-west ;
also eastward to Aden and, if actwaria is really the same species,
to India and Ceylon.

ACIDALIA SpOLIATA Walk. (2).

Acidalia spoliata Walk. List Lep. Ins. Brit. Mus. xxii. p. 744
(1861).

1908: Nov. 22,19.

The single example is ef the mnorata group, larger than that
species, somewhat less reddish and more marked than ier-
nataria Walk. (List Lep. Ins. xxii. p. 746), very likely a deeply
coloured form of spoliata Walk. (from §. Africa), or, perhaps, a
form of the widely distributed Oriental species, nescraria Walk.
(List Lep. Ins. xxii. p. 750).

ACIDALIA HORIOCHR@A, Sp. n.

2, 20-23 mm. Face blackish. Palpus white beneath, dark-
mixed above. Vertex white. Collar pale ochreous. Thorax,
abdomen, and legs concolorous with wings, fore femur and fore
tibia infuseated above. Fore wing of medium breadth, with apex
moderately pointed ; dirty white, irrorated (variably in strength
in the different individuals) with brown-grey ; lines moderately

150 PROF, E. B, POULTON ON

strong, though resolving themselves under the lens into con-
densed irroration ; ; antemedian slender, generally indistinct ante-
riorly, oblique outwards from one-third costa, strongly recurved
in cell, oblique inwards to about one-fourth hind margin,
faintly sinuous; discal dot small but sharp; median line
thicker than the others, well beyond discal dot, very strongly
oblique outwards from costa, very sharply bent subcostally,
thence very slightly obliquely inwards and sinuous, the sinu-
osities being, as usual, at the folds, but never very deep, some-
times scarcely appreciable; postmedian fine, midway between
median and termen or slightly nearer to the former, parallel
with termen except at costa, where it makes a bend, though
less sharply than the median, sinuosities slight or very slight ;
subterminal indicated by absence of irroration, accompanied
proximally by a band of stronger irroration, distally by a band
of browner tone, sometimes partly obscured by the grey irro-
vation, but always noticeable, usually clearest anteriorly, not
rarely showing a bright brown, almost ferruginous spot near
apex; a narrow white line separates this band from the terminal
line, which is black, thickest between the veins, slightly inter-
rupted at the veins and extends rownd the apex for some distance
along the costa, as in the swhmutata group; fringe with a line
of stronger irroration beyond the middle, distally hereto less
strongly irrorated than proximally. Hind wing with termen
not bent at R*; first line absent; median just proximal to
discal dot, obsolete anteriorly ; postmedian slightly sinuous, nearly
parallel with termen; distal area nearly as on fore wing, the
brownish band fading out at apex instead of becoming more
conspicuous, the terminal line not extended round apex ; fringe
as ou fore wing. Under surface more weakly marked, that of |
fore wing somewhat suffused basally, that of hind wing whiter ;
first line and sometimes median shade obsolete or nearly so.

1908: Aug. 24,—2 9. 1909: Jan. 16,—1 9; Jan. 19,—1 9;
Feb. 18,—1 Q; Feb. 19,—1 9; Apr. 22,—1 9; May 10, an Q;
May 29,—1 9; Aug. 17,—1 9; Sept. 17,—1 2 (type); Dee. 30,
—19. 1910: Jan. 8—19; Jan. 12,—1¢9.

A very distinct though unostentatious little species, recog-
nizable especially by the characters printed in italics.

ACIDALIA (PYLARGE) NEPHELOPERAS, Sp. n.

3g, 16-23 mm.; 9,19-23 mm. Superficially like Glossotrophia
romanaria Mill., and rufomixtata Rbr., but structurally an Acidalia
of the section Pylarge. Best described by a comparison with
the well-known 4. submutata Tr., with which it entirely agrees
in the markings (though these are, on an average, more sharply
expressed), including the continuation of the black terminal line
round the apex, and the tendency (sometimes very strong) to
blue-grey clouding in the distal area. Much smaller; ground-
colour varying from ochreous whitish to reddish sand-colour
(much as in pulehellata Fab.), antennal ciliation in both sexes
longer, male hind tibia with a pair of spurs, face pale in lower

MOTHS FROM SOMALILAND. 151

half, termen of fore wing slightly more curved, of hind wing
not suberenulate ; terminal line accompanied proximally by a fine
whitish line (as in pelchellata), and on the fore wing expanded
into a large triangular dot anteriorly to SC’, fringe more strongly
dark-dotted than in suwbmutata; under surface glossy, that of
hind wing scarcely paler than that of fore wing, discal dots
rarely quite obsolete, fore wing generally with noticeably dark-
ened border distally.

1908: Feb. 13,—1 9; Sept. 15,—1 ¢; Sept. 22,—1 9; Nov.
13,—12. 1909: Jan. 12,—19; Jan. 20,—19; Feb. 18,—1 6;
Mar. 24,—1 ¢ (type); Apr. 14,—1°. .

There is also a male from Port Sudan (Mrs. H. NV. Waterfield)
in coll. Brit. Mus., and a larger one from Bhuj Kutch (Z#.-
Colonel C. G. Nurse).

Rather variable in colour and in the strength of the markings.
Related to Acidalia (Pylarge) ocellicincta Warr. (Nov. Zool. viil.
p: 9), from British Hast Africa.

ACIDALIA PULCHELLATA Fab.

Phalena pulchellata Fab. Ent. Syst. iii. (2) p. 171 (1794).

Acidalia addictaria Walk. List Lep. Ins. Brit. Mus. xxi.
p- 749 (1861).

Craspedia addictaria Hmpsn. Faun. Ind., Moths, ii. p. 429
(1895).

? Craspedia rufinubes Warr. Nov. Zool. vii. p. 91 (1900).

1908: May 4,—1 g. 1909: Mar. 24-1 2; Apr. 24,—1 9;
May 6,—1 2. 1910: Mar. 18-19.

Mostly of a more ruddy form than the Indian. The British
Museum collection has one quite similar from Aden.

ACIDALIA TIMIA, sp. n. (Pl. I. fig. 19, 2-)

@, 21-26 mm. Face blackish fuscous, very narrowly pale-
edged beneath. Palpus fuscous above, pale beneath. Antenna
(as in nepheloperas) unusually strongly ciliated for a female, the
cilia fully one-half as long as diameter of shaft. Vertex, thorax,
and abdomen concolorous with wings; collar ochreous. Fore
femur darkened on upper side. Fore wing with apex not very
sharp, termen straight anteriorly, gently curved posteriorly, not
extremely oblique; palest fleshy ochreous, rather glossy (similar
to beckeraria Led., but still paler), without dark irroration, but
in places slightly clouded with less whitish fleshy-ochreous ;
antemedian and median lines (or narrow shades) ochreous, very
feeble, sometimes almost entirely obsolete, the former sometimes
marked with darker dots on SC, M, and SM°; discal dot usually
distinct, sometimes elongate, placed on the median shade ; post-
median line fine and faint, but marked with fuscous dots on the
veins (a larger one at costa), shaped about as in beckeraria ;
darker ochreous, fuscous-dotted spots or patches commonly follow
the postmedian between the radials and at posterior margin ;
terminal line ochreous, very feeble, especially posteriorly ; fringe

152 PROF. E. B. POULTON ON

concolorous, in strongly marked specimens with dark ochreous
or even fuscous-mixed dots. Hind wing with termen smooth;
. concolorous with fore wing, discal dot and postmedian row of
dots present, the latter followed by a not very strong band of
ochreous shading. Under surface glossy, slightly less pale
ochreous (especially the fore wing), without markings.

1908: Mar. 24,—19; Aug. 15,—19; Sept. 26—19. 1909:
Jan. 14,—1 9; Jan. 20,—1 9; Apr. 16,—1 ¢; Apr. 20,—1 9;
May 8,—1 2; Oct. 7,—1 9 (type). 1910: Jan. 8—12.

Apart from the colour, and the presence of dark cloudings
distally to the postmedian line, this neat little species differs from
beckeraria in having the termen of the fore wing, on an average,
less oblique.

ACIDALIA PYRRHOCHRA, sp.n. (PI. II. fig. 18, 2.)

Q, 23-25 mm. Structure of the preceding. Face blackish
fuscous (in all three examples badly abraded below). Shape and
essential markings of timia, of which it might possibly be an
extraordinarily different colour-form, unconnected with the name-
type by any transitions. Strongly rufous ochreous, as in fulvi-
color Hmpsn. (Nat. Hist. Socotra, p. 331), or the most rufous
aberration of nepheloperas Prout, in two of the examples finely
irrorated with blackish, in the other more uuiform ; in place of
the two ochreous patches which characterize timia there 1s a
continuous black-grey clouding proximally to the dentate sub-
terminal line (which is, in consequence, distinctly expressed),
and some slighter shading distally to the same—altogether re-
calling certain forms of marginepunctata Goeze, nepheloperas
Prout, etc., more than timia. Under surface rather paler than
upper, the distal cloudings faintly indicated in greyish.

1908 : Oct. 30,—1 92 (type). 1909: Apr. 9,—1 9; Nov. 16,
OR

The last-mentioned (the example without blackish irroration)
is further aberrant in having the distal cloudings extremely
weak, the discal dot of the fore wing surrounded by an ill-defined
deeper reddish spot, that of the hind wing very minute. Easily
distinguished from the reddest form of nepheloperas by the
absence cf black terminal line and triangular subapical dot, ete.

ACIDALIA LURIDATA (Zell.).

Idea luridata Zell. Isis, 1847, p. 20 (nec Ster.).

Acidalia cenosaria Led. Verh. zool.-bot. Ver. Wien, v. p. 209,
t. 3. fig. 3 (1855) (ab.).

Acidalia luridata Prout, Seitz Macrolep. iv. p. 64, t. 4e (1913).

1908: July 31—19; Aug. 25,—19. 1909: Jan. 12,—19;
Feb. 22,—1 3; May 8,—1 3; Sept. 18.—1 9; Oct. 24-19.

Fairly typical, 7. e. ratber darker than, and not quite so reddish
as, the form cenosaria Led., which is the more general in
S. Europe and Asia Minor. The distribution of the species

MOTHS FROM SOMALILAND. 153

extends from Greece and Northern Egypt to Zerafshan and
N.W. India, and the British Museum has an example from
Yemen, Arabia, but the present specimens extend its known
range in Africa,

ACIDALIA SAGITTILINEA (Watr.).

Craspedia sagittilinea Warr. Nov. Zool. iv. p. 219 (1897).

1909: Feb. 15,—16d; Feb. 16,—1 6; Feb. 17,—1 ¢.

Described from Mombasa, and I have seen a few from other
localities in British Hast Africa. Both these Somaliland ex-
amples are rather less strongly marked (especially beneath),
the median shade faint, placed midway between discal dot and
post-median line.

GLOSSOTROPHIA DISPARATA SOMALIATA, subsp. n.

@, 17-19 ay Name-typical disparata Hmpsn. (Nat. Hist.
Socotra, p. 532, Craspedia) has never been described, its recog-
nition ee depending ona good, though uncoloured figure
(ibid. t. 20. fig. 18), and a note by Rebel (Denks. Akad. Wien.
Math.-nat. KI. Ixxi. 2, Sep. p. 69) to the effect that it belongs to
the confinaria group of Acidalia (i.e. Glossotrophia Prout), and
that the male antenna might almost be called shortly pectinate
(i. e. subdentate with fascicles of cilia). It is the smallest of the
genus; male hind tibia with one spur, hind wing slightly less
regularly rounded than in the typical species (slightly bent at
R’*); sand-colour with dense dark irroration, not “fluted” as in
romanaria Mill., terminal line (except towards apex) broken into
very short, strong dashes, at and round apex fine and less pro-
nounced than in most of the species. ace concolorous. Palpus
dark-mixed on outer side. ‘Tongue moderately long. Subsp.
somaliata (bon. sp.?) is rather larger, fore wing slightly longer,
irroration much hghter, showing a Y Paatille fendeney towards “ihe
“fluting ” ; terminal dashes more slender and less black, apical
line somewhat more pronounced. Palpus with less dark spotting
on outer side.

1908: Sept. 14—192. 1909: Jan. 11,—1 Q (type).
ZYGOPHYXIA TORNISECTA, sp. n.

3,14 mm.; 2, 16-19 mm. Faceand palpus fuscous. Tongue
slender, vather short, Antennal ciliation im male moderately
long. Vertex white. Collar white, with a slight ochreous tinge.
Thorax, abdomen, and legs concolorous with wings. Hind tibia
in both sexes with terminal spurs. Wings less narrow than in
relictata Walk. (the type of the genus); fore wing only slightly,
hind wing decidedly, narrower than in elongaria Rbr., which in
some respects it rather recalls ; hind wing with shallow, rounded
excision from M* to tornus, inner margin consequently shortened.
Dirty white with a tinge of brownish, and with moderately
strong fine brown-grey irroration ; discal dots small, black ; lines
generally not sharply defined (in the male particularly weak),

154 PROF. E. B. POULTON ON

those of the fore wing approximately parallel with the termen,
the slender antemediar n and broad (sometimes strong) postmedian
perhaps slightly more oblique and with a strong proximal bend
at costa, the median touching the distal edge. of the cell-dot,
postmedian slender, proximal subterminal broad, distal subter-
minal very weak; first line of hind wing absent, median shade
rather straight, except the sharp anterior bend; termen with
black dots between the veins; fringe dusted with fuscous, most
thickly opposite the veins, a fine pale line at its base, a broader
one just beyond the middle. Underside similar or more blurred.
1908: Aug. 15.—-192. 1909: Feb. 28,—1¢ (type); Mar. 24,
—12; Apr. 15,—1 2.

PYrYCHOPODA SUBTORRIDA, sp.n. (PI. II. fig. 17, 2.)

2,18 mm. Face and palpus black. Vertex, antenna, thorax,
abdomen, and fore leg concolorous with wings (other legs lost).
Fore wing rather narrow ; light ochreous brown, almost entirely
suffused with vinous, less so basally and costally ; scattered black
irroration; lines black; antemedian from one-fourth costa,
oblique outwards, very acutely angled in cell, then equally oblique
inwards to behind M, thus forming a V-mark, a second, much
shorter angle outwards at fold, but the entire posterior half of
the line iess strong, more dissolved into coarse black dots ; median
line from mid-costa, sinuous in §-shape, the anterior (outward)
curve crossing the deep-black cell-spot ; postmedian from costa
at nearly three-fourths, forming an inward curve or bend at first,
sharply angled outwards at R' (forming a shorter V than the
antemedian), weakly incurved between the radials and boldly
between M’* and SM”, finally oblique inwards at hind margin ;
distal area with blackish cloudings, the most conspicuous being
one at R' (interrupted at the position of the obsolete subter minal
line) and a longitudinally elongate pair on either side of M’;
rio terminal line; fringe very long, its proximal half light
ochreous brown, opposite the veins with amorphous spots formed
of groups of black dots, distal half paler and somewhat greyer,
unmarked. Hind wing rather narrow, with termen almost
smooth, not very strongly convex ; more strongly irrorated than
fore wing, especially at base; first line not developed ; median
thick and somewhat diffuse, proximal to the sharp black cell-
spot; postmedian as on fore wing, but with the angle at R° less
pointed ; distal area with two somewhat sinuous bands of dark
irroration, enclosing a pale subterminal line; fringe as on fore
wing. Under surface considerably paler, costal margin of fore
wing finely dark-dusted, the rest rather smooth and glossy ; both
wings with strong discal spot (larger than above) and a moderately
strong outer line, starting from a slightly enlarged spot at costa
which corresponds to the origin of the postmedian of upper
surface, but oblique outwards, bent about R', thence approxi-
mately parallel with termen, corresponding to the proximal sub-
terminal dark shade of the hind wing above ; fringe unspotted.

or

MOTHS FROM SOMALILAND, 15

1909: Mar. 24,—1 ? (type).

Near torrida Warr. (Noy. Zool. xi. p. 468), termen of hind
wing less protuberant, antemedian line more acutely angled, not
connected with median by dark shading, under surface more
strongly marked.

PrycHopopa nicrosticra Warr. (2).
Ptychopoda nigrosticta Warr. Nov. Zool. iv. +P. 61 (1897).

1908 : Feb. 24,—1 9 (rather worn).

The large discal dots and the markings, so far as they can be
made out, suggest the more mottled forms of this species, but the
former are perhaps somewhat exaggerated, the wings are rather
more coarsely dusted, with stronger dark markings near the
termen, the underside with the cell less infuscated and with the
spot near the tornus apparently wanting. Described from Natal
and occurs in the Transvaal and, perhaps, British East Africa,
but it is by no means certain that these more northerly speci-
mens belong here: they may be slightly narrower winged. Good
specimens, and especially the male, must be awaited.

PTYCHOPODA sp.
1908: Aug. 26—19. Rather rubbed.

Probably new, unless it be a form of subpurpurata Stgr. (from
Syria, etc.). Rather uniform reddish sand-colour, very weakly
marked ; fore wing with traces of a strongly sinuous darker
median line, hind wing with fine, strongly sinuous postmedian
line; both these lines marked with some fuscous dusting.

PrycHOPODA APERTA, sp. n. (PI. II. fig. 16, 3.)

3,17 mm. Face dark fuscous. Palpus short and slender,
dark fuscous. Tongue slight. Antennal ciliation scarcely longer
than diameter of shaft. Head, body, and legs about concolorous
with wings; fore leg fuscous above; abdomen dorsally belted
with fuscous. Fore wing with all the subcostals on a common
stalk, through suppression of distal wall of areole (base of
SC°°); brown with a tinge of reddish and with rather coarse
blackish-fuscous irroration, mainly longitudinal in direction ;
base more strongly irrorated costally; first line ill-defined, bent,
becoming oblique inwards, with some black marking near costa ;
median shade also ill-defined (especially in anterior half), strongly
sinuous, the proximal curve in submedian area being rather deep;
postmedian line well expressed, from a black spot at two-thirds
costa to one at about three-fourths hind margin, forming a slight
outward curve from SC’ to M’ and a stronger submedian inward
curve, marked throughout with small dark dashes on the veins ;
some broad, vague, irregular dark shading between this and
termen ; fringe with large black spots opposite the veins. Hind
wing with termen somewhat sinuate towards tornus; M’ separate
at its origin from R*; concolorous with. fore wing, the median

156 PROF. E. B. FOULTON ON

and postmedian lines strongly expressed; distal area and fringe
as on fore wing. Under surface paler; fore wing very weakly
marked, only the postmedian line showing, and that faintly ;
hind wing with median and postmedian lines, though much
weaker than above; fringes with the dark spots present, but
weaker than above.

1909: Jan. 13,—1¢ (type).

Tn some respects similar to Pé. semilinea Warr. (Nov. Zool. 111.
p. 314), from the Khasia Hills. Remarkable for the subcostal
venation of the fore wing, which is like that of Chrysocraspeda
and almost unique in Ptychopoda; I know of only one species
which shares the peculiarity, namely marginata Swinh. (Tr. Ent.
Soc. Lond. 1894, p. 182), unless chrysocilia Hmpsn. (Ill. Het. vin.
p- 124) is also a Ptychopoda; in any case none of the three has
any connection with Chrysocraspeda, which has strongly pectinate
male antenna, hind tibia with four spurs, cell of hind wing
short, ete.

Subfam. LARENTIINA®.

PSEUDOSTERRHA PHIL#ARIA (Brabant).

Sterrha philearia Brabant, Bull. Soe. Ent. Fr. 1896, p. 384.

Pseudosterrha gaynert N. Rthschd. Nov. Zool. vill. p. 433
(1901); xi. t. 4. fig. 10 (1905).

1908: Sept. 16,—1 9.

Described from Egypt. J suspect, however, it is merely a form
of—or even entirely synonymous with—Ps. pawllula Swinh.

(Proc. Zool. Soe. Lond. 1886, p. 456), from India.

Eupryra (CAMPro@RAMMA) NATALATA (Walk.).

Scotosia natalata Walk. List He Ins. Brit. Mus. xxv. p. 1681
(1862).

Scotosia rubritincta Hmpsn. Bull. Liverp. Mus. ii. p. 38 (1899).

Hargaisa—1908 : Oct. —1<.

Widely distributed in Kast Africa, also Socotra.

Subfam. GEomMETRINE.

HETEROSTEGANE INDULARIA (Guen.).

Stegania indularia Guen. Spec. Gén. Lép. x. p. 46 (1858).

Mandera.—1908: Apr. 27,—1 35; May 2,—1 9; May 28,—1 9;
June 1,—2d5, 492; June 1] Ge Trine 29,2 6; July 8,—
1g; July 17,—1¢; July 24,—16; July 31,—1 64; Sept. 13,—
36,12; Sept.14,—1 dg; Sept. 17,—2d; Sept. 22,4 ¢; Sept. 24,
43,19; Sept. 26,246,199; Oct.1—l1¢. 1909: Mar. 24,—
19; Apr. 8—1¢; May 10,—19. 1910: Mar. 2,—12.

Hargaisa.—1908 : Oct.,—1¢.

The series from Mandera shows the usual range of variation in
eround-colour and intensity of markings, but nearly all are of
small size.

MOTHS FROM SOMALILAND. Way

The male from Hargaisa may perhaps be referred to ab. cali-
data Warr. (Nov. Zool. iv. p- 78), though the ground-colour
remains pale; all three lines are strongly developed ; possibly a
distinct species, as the palpus looks a little stronger and the
antenne (unfortunately damaged) may have less long ciliation.

Described from Abyssinia, but extends through eastern Africa
to the Cape.

ZAMARADA SECUTARIA (Guen.).

Stegania secutaria Guen. Spec. Gen, Lép. x. p. 45 (1858).

Zamarada pulverosa Warr. Nov. Zool. i. p. 158 (1895).

Mandera.—1908: Sept. 23,—1<¢.

Hargaisa.—1908: Oct..—1 6.

Both are small, the Hargaisa example rather worn, but
apparently more weakly dusted than usual.

This species is likewise distributed from Abyssinia to the

Cape.
OSTEODES PROCIDATA Guen., subsp. ERITREENSIS Prout.

Osteodes procidata Guen. Spec. Gén. Lép. x. p. 177 (1858).

Osteodes turbulentata Guen. ibid. (1858).

Aspilates semispurcata Walk. List Lep. Ins. Brit. Mus. xxvi.
p. 1679 (1862).

Gan Libbah.—1908: June 24,—1¢. 1909: Nov. 4,19.

Both with the dark borders strong (subsp. eritreénsis Prout,
Nov. Zool. xxii. p. 348, described from Eritrea).

Distribution as in the two preceding. Warren (Nov. Zool. ix.
p. 529) apparently regarded Guenée’s two forms (from Abyssinia
and the Cape) as separate species, but Guenée’s descriptions and
the material before me suggest that Swinhoe (Tr. Ent. Soc. Lond.
1904, p. 504) is correct in uniting them.

T)-SCALMA SUBCURVARIA (Mab.).

Tephrina subcurvaria Mab, Ann. Soc. Ent. Fr. Ixvi. p. 278
(1897).

Hargaisa.—1908: Oct.,—1¢.

This species, apparently common throughout East Africa from
Abyssinia to Natal, and originally described as from West Africa,
has been mixed in our British collections with observata Walk.
(List Lep. Ins. xxii. p: 963) from the Cape, but I doubt the
correctness of this; the latter, apart from its much darker and
vather more reddish colour, has the median shade of the fore
wing more oblique and the postmedian line rather further from
the termen.

DIscALMA PUERILIS, sp.n. (Pl. II. fig. 15, ¢.)

3,16mm. Face apparently without projecting cone of scales.
Antennal joints scarcely projecting, the cilia scarcely as long as
diameter of shaft. Head, body, and legs concolorous with wings ;
hind tibia dilated, with long hair-pencil, tarsus rather short.

158 PROF. KE. B. POULTON ON

Fore wing with fovea slight, SC'* coincident, touching © at a
point ; dirty white irrorated with brown-grey, leaving clearer a
slight, ill-defined antemedian band (at least towards hind margin),
a fairly broad but exceedingly ill-defined band just distally to the
cell-dot and especially a broad, somewhat sinuous subterminal
line, from costa at less than 1 mm. before apex to tornus, slightly
dark-shaded proximally, especially at costa and hind margin; a
small discal dot near costa at middle; a slightly darkened terminal
line, interrupted at the veins; fringe pale (whitest proximally),
traversed in middle by a fine grey line and cut throughout its
length by brown-grey spots opposite the veins. Hind wing with
termen almost smooth (scarcely waved); similar to fore wing, but
with pale band-like markings scarcely indicated, subterminal line ©
not quite so conspicuous as on fore wing; no discal dot;
indications of a dark spot (end of a line) about middle of inner
margin. Under surface similarly but more faintly marked ;
discal dot of fore wing absent.

1910: Feb. 14,—1 ¢ (type).

A very distinct though modest little species; it may be
pictured by imagining a miniature edition of the North American
“ Seiagraphia” nubiculata Pack., exceedingly washed-out, dark
lines of hind wing absent; structure as in that species, wings
slightly narrower.

DISCALMA CALVIFRONS, sp. n. (PI. II. fig. 14, ¢.)

$,17-20 mm. Face rather protuberant, rough, almost scale-
less, with small horny prominences at upper and again at lower
edge. Palpus rather short, rough-scaled, grey mixed with fuscous,
3rd joint very small and blunt. Antenna with joints projecting,
the ciliation about as long as diameter of shaft, arranged in very
slender even fascicles. Thorax and abdomen concolorous with
wings, the patagia in front more fuscous. Hind tibia with hair-
pencil. Fore wing with fovea; not broad, the termen being
rather straight and moderately strongly oblique ; SC** coincident,
sometimes free, sometimes slightly connected or anastomosing at
a point with one or both of the adjacent veins; white-grey,
irrorated with brownish fuscous; discal dot black, variable in size ;
lines fuscous, irrorated more or less with blackish, sometimes
sharply dark at costa; antemedian from nearly one-fourth costa,
oblique outwards, strongly angled in front of M, then oblique
inwards to M, here sometimes marked with a blackish dot, then
running perpendicularly to hind margin, occasionally with a very
slight curve inwards ; median line very variable in strength and
thickness, usually crossing, sometimes just proximal to, the discal
dot, almost straight except for a small proximal bend at costa;
ostmedian from before three-fourths costa, vertical or (oftener)
slightly oblique outwards, curved or angled at R’, thence about
parallel with termen, marked with blackish vein-dots, sinuate
inwards in-‘submedian area and slightly oblique outwards at hind
margin ; distal area (at least in its proximal half) clouded with

MOTHS FROM SOMALILAND. 159

dark grey, with a distinct, sinuous, whitish, subterminal line,
which is rather thick in places, and especially widens at costa so
as almost to reach apex; proximally to the subterminal an
irregular blackish spot between R® and M', sometimes also
anteriorly to R®; termen with a row of moderately thick blackish
dashes ; fringe weakly dark-chequered opposite the veins. Hind
wing with termen almost perfectly rounded, sometimes with a
slight suggestion of prominence at R* and sinuosity in front of
the same; similar to fore wing, without the antemedian line.
Fore wing beneath somewhat more suffused from base to median
shade ; discal dot weak; the shade between postmedian and sub-
terminal strengthened into a dark band, without the blackish
spot, the subterminal itself obsolete; band between median and
postmedian lines, also the posterior half of distal area (or at least
a patch behind R*) somewhat whiter than above. Hind wing
beneath whitish, with median shade, discal dot and outer band
distinct. Remelle. on an average, rather larger, rather broader-
winged, sometimes more suffused, lines generally weak (except
the Gostal spots), distal dark shade extended to termen, usually
almost obliterating the subterminal line except costally, where
the subapical pale spot persists, at least in part, black spot
proxumally to subterminal line between R* and M? seldom
developed; the hind wing and under surface show corresponding
differences.

1908: Sept. 14,—1 3; Sept. 24,—1 g; Sept. 28,—1 5; Oct. 14,
= OF Octy Nog 5) Oct.) 24-1 3) (type)s 1909); Apr: 10;
7h Or Apr. 14, ie GR Ayoe: poms 2; Apr. 27,—19; May 10,—
19; May 12,—1°.

Variable both individually and (in particular) sexually. More
recalls certain tropical American species (e. g. Macaria nigricomma
Warr. in the case of the male, heliothidata Guen. in that of the
female) than any African species with which I can compare it.
Scarcely a true Discalma.

MACARIA SEMIALBIDA Prout.

Macaria semialbida Prout, Nov. Zool. xxii. p. 351 (1915).

1908: June 21,—1<¢.

Antenna br aan, but a remnant shows that Ths ciliation is
scarcely as long as the diameter of the shaft. Hind tibia strongly

dilated. Tew strong.
Founded on females from British East Africa.

MACARIA OBLIQUILINEATA (Watr.).

Gonodela obliquitineata Warr. Nov. Zool. vi. p. 307 (1899).

Semiothisa obliquilineata Swinh. Tr: Ent. Soc. Lond. 1904,
p. 507.

1908: May 4,29. 1909: Mar. 14,—-1¢; Apr. 11,—1 9;
May 10,—1 9; Nov. 10,—1 9.

Rather variable, on an average slightly smaller than the

160 PROF, E. B. POULTON ON

examples (Abyssinia, White Nile, British Hast Africa) in the
_ British Museum, postmedian line of fore wing rather more curved
at costa. In the fore wing veins SC'™ are coincident, in three
examples free, in three slightly connected with C.

TSPHRINA INCONSPICUA Warr.

Tephrina inconspicua Warr. Nov. Zool. iv. p. 113 (1897).

1908: July 2,—19; Aug. 24,—19.

Rather weakly marked, especially the hind wing, which appears
rather more whitish than in the normal form (Natal to Nyassa-
land); but it is not in quite perfect condition. In the fore wing
the costal end of vein C is obsolete, leaving it to appear that C
and SC are coincident throughout, anastomosing shortly with
SC, A worn female from Arabia (coll. Brit. Mus.) appears to
agree with the Somaliland example, though larger and with
normal venation; thus the species seems to be widely distributed.

TEPHRINA BUTARIA (Swinh.).

Semiothisa butaria Swinh. Tr. Ent. Soc. Lond. 1904, p. 510.

1909: May 8,—1¢.

Known from Central and Hast Africa and as far north as
Abyssinia.

Tepurina nerra (Holland).

Grammodes netta Holland, in Donaldson-Smith, Through
Unknown African Countries, p. 418, fig. 9 (1897).

1908: July 1,—1 4; July 31,—19; Sept. 21,—19; Sept. 26,
—19. 1909: May 21,—1 9; July 8—19; Aug. 17,—16.

As I understand Sir George Hampson has compared Holland’s
type, I accept the determination of the species at the British
Museum, which possesses a single example from Abyssinia. The
fore wing agrees well with Holland’s figure, but the hind wing
has distal borders nearly as in Discalma subcurvaria Mab., or
even more extended to the termen. Male antenna bipectinate.

TEPHRINA CINERASCENS (Btlr.).

Acidalia cinerascens Btlr. Ann. Mag. Nat. Hist. (4) xvi. p. 418
(1875).

Ematurga bilineata Warr. Nov. Zool. 11. p. 129 (1895).

Tephrina cinerascens Swinh. Tr. Ent. Soc. Lond. 1904, p. 511.

1909: May 10,—1@. Rather worn, weakly marked.

Distributed from the Cape to British East Africa, and doubt-
fully distinct from pumicaria Led. (Syria) and fwmosa Hmpsn.
(India). The large, strongly marked Abyssinian form figured
by Guenée (Phal. t. 17. fig. 7) is unknown to me.

TEPHRINA DISPUTARIA (Guen.).

Lubolia disputaria Guen. Spec. Gén. Lép. x. p. 489 (1858).
Tephrina disputaria Hmpsn. Faun. Ind., Moths, 1. p. 209
(1895).

MOTHS FROM SOMALILAND. 16]

Mandera. ec June A 19; June 29,—19; July 5,
19; July 17,— ns ; Aug. 2 3 a Sept. (134 12; Sept. 19,
19; Sept. 26,— 19; oe Hh 12 2,—19. 1909: Jan. 18, 19;
Jan, 19,—-1:9 ; ue 12,—19 ;: Mar. 19,—1 9; Apr. 7,—19?
Apr. 22,—1 9; Aug. 17,—1¢,29; Oct. 5,—19; Nov. 9,—2°?.
1910: Mar. 2,—1 92; Mar. Gia] @. Year ?: Feb. 25,—1 9.

Hargaisa.—1908 : Oct..—4 ¢.

Kixtremely variable, occurring probably throughout the year,
though none was taken in February or December, ‘The great
majority of el females belong to the form sebocellata Warr.
(Novy. Zool. iii. p. 413), described from $. Othman, Arabia, [ follow
Swinhoe in otetienng subocellata to be a form of the variable
disputaria Guen. Two of the males are small, with the distal
part of the median area almost free from dark dusting, the post-
median line more than usually bent. A few females are also
small, but otherwise normal. Two females are smooth-scaled,
the distal area broad, inclining to violet-grey, the black marks on
hind margin of fore wing and those distally to the middle of the
postmedian line very sharply prominent.

Described from Neypt, but widely distributed in North and
Kast Africa, Arabia, India, ete.

TEPHRINA DEERRARIA Walk.

Tephrina deerraria Walk, List Lep. Ins. Brit. Mus, xxiii. p. 962
(1861); Swinh, Tr, Knt. Soc. Lond. 1904, p. 510.

Mandera.—1908 : Oct.15,-1 9. 1909: Apr.7,--1 9; Apr. 10

19°; Apr. 22,—19; May 10,—19; Oct. 7,—16; Oct. 11
1g. 1910: Jan. 8,—1¢.

Buggan. 1908: June 27,—1 3.

In one female the discal spot of the fore wing is reduced, scarcely
forming an ocellus. Generally larger than the preceding, less
brown, postmedian line straighter, otherwise hard to distinguish ;
pevhaps a form of the same. Distributed from the Cape to
Abyssinia,

)

'TEPHRINA PRIONOGYNA, SP. 1. Ply toe lig c
D) | 5 > +

®, 28-33 mm, Antenna strongly serrate, the serrations of
the outer series so long as to form rudimentary pectinations.
Fore wing with SC’ out of GU, free or anastomosing at a point
with S¢ a SC? in one example anastomosing at a point with SO™.
Otherwise extremely like strongly irrorated females of the pre-
ceding * ; discal mark of fore wing reduced to a small weak
dot, median shade usually absent, antemedian line of fore wing
more strongly curved, postmedian curving anteriorly (both vari-
able in distinctness), terminal dots weaker or obsolete, the dark
distal border beneath extended to the termen, or else becoming

* Tn disputaria and deerraria SC! ave coincident, often anastomosing at a point
or slightly connected with SC" Kyven if this difference prove not absolutely
constant, it is at least general.

Proc. Zoou, Soc.—1916, No, XI. 11

162 PROF, E. B. POULTON ON

uniformly lighter there, no tendency towards the central pale
terminal patches which are general in deerraria, termen of hind
wing smooth (in disputaria often slightly more undulate),

1909: Apr. 15,—19; May 9,—19; May 10,—3@ (including
type).

C@NINA TERGIMACULA, sp. n. (PI. II. fig. 12, ¢.)

3 9,24 mm. Face ochreous, lower half with a large fuscous
spot on each side. Palpus with 3rd joint short, ochreous mixed
with fuscous. Male antennal pectinations long and reaching to
near apex—beyond three-fourths (female without head). Vertex,
thorax, and abdomen concolorous with wings, abdomen with a
dark fuscous dorsal blotch anteriorly. Fore wing narrow, costa
and termen almost straight, hind margin somewhat convex, in
female sinuous ; SC? connected by bar with SC*4, R°-M? stalked
(sometimes rather long-stalked) ; pale ochreous whitish, irrorated
and suffused, especially at base and half-way along eosta, with
purple-grey ; a brown or fuscous stripe from beyond middle of
costa, oblique outwards, strongly curved or angled between R*
and R*, oblique inwards (and forming a very slight proximal
curve) to middle of hind margin ; a faint, oblique antemedian line
or shade usually indicated, another midway between postmedian
and termen, bearing in its anterior part two or three darker
wedge-shaped spots, the one before R' the strongest and blackest ;
fringe spotted with fuscous. Hind wing narrow, especially in
temale, tooth at end of SC? in female very aeute and produced,
termen m male weakly, in female more strongly, toothed at R’,
anal angle scarcely produced; concolorous with fore wing or
rather more brightly ‘and shar ply marked ; a broad, dark, olive-
brown, fuscous-mixed, somewhat sinuous Bend from inner margin
near anal angle to costa near apex, finely white-edged distally,
the white broadening anteriorly. Under surface similar, the
markings weaker and more diffuse.

1908: Oct. 3—1 5. 1909: Apr. 7,1 3; Apr. 12,—1 ¢
(type); Sept..—1?.

Genus SESQUIALTERA, nov.

Face slightly prominent, convex, with moderately appressed
scales. Palpus moderate, rather stout, rough-scaled, terminal
joint short. Tongue short. , Antenna in male (?); in female
slightly subserrate, with very minute ciliation. Pectus and
femora moderately hairy. Hind tibia with terminal spurs only.
Fore wing long and narrow (greatest breadth—mid-costa to
tornus—three-eighths of length), costa nearly straight, apex rather
sharp, termen, except close to apex, extremely oblique, slightly
curved, very faintly sinuous, rather longer than hind margin ;
tornus very weak ; cell short, less so at its extremities, DC” being
oblique inwards and DC? extremely oblique outwards ; SC'™ long-
stalked, SC? sometimes anastomosing with C, SC* connected by a

MOTHS FROM SOMALILAND. 163

bar with SC** about opposite the origin of SC*, SC*> long-
stalked from just before end of cell, SC** separating near apex ;
radials normal ; M! long-stalked with R*, M? arising rather near
end of cell. Hind wing only about halt the length of fore wing,
costal margin strongly concave, termen with long projecting
teeth at SC? and R’, otherwise sinuous rather than dentate,
tornus moderate ; cell about one-half, DC incurved ; C approxi-
mated (but not appressed) to SC to near end of cell, then
moderately diverging ; SC’ from close to end of cell; R?® absent ;
R*—M" long-stalked, M? from very near end of cell.

Type of the genus: Sesquialtera ridicula, sp. n.

SESQUIALTERA RIDICULA, Sp. ne (PI. II. fig. 11, 9.)

2, 32-36 mm. Head, body, and fore wing fuscous with darker
irroration. Fore wing extremely weakly marked, a darker cell-
mark and a postmedian line indicated, the latter remote from
termen, strongly excurved at radials; sometimes also a vague,
diffuse, oblique, somewhat curved antemedian line or shade in-
dicated. Hind wing variable, in the type with the basal area
blackish fuscous, the rest more concolorous with fore wing, but
traversed by ill-defined ferruginous-brown bands, in other ex-
amples more uniform, the base being less blackened, the ferru-
ginous scales largely suppressed ; a blackish discal dot, crossed or
closely preceded by the more or less sinuous antemedian (median)
line and followed by a distinct postmedian (darker brown or
blackish) sinuate inwards between radials and in submedian area ;
a brown or black terminal line. Under surface similar to upper,
in the type less variegated in hind wing.

1909: May 12,—1 9 (type).

Also in Hope Department, Oxford, from British East Africa,
5 May, 1913, c. 1°S., 35° K., 5000-6000 feet, forest with open
glades (H. B. Popplewell), 19. Also from Mt. Kenya in Paris
Museum.

PACHYPALPIA SUBALBATA Warr. ;

Pachypalpia subalbata Warr. Noy. Zool. vii. p. 98 (1900).

1908: Oct. 25,—1°.

Described from British East Africa. Known also from German
East Africa. ;

ASCOTIS SELENARIA (Schiff.).

Phalena Geometra selenaria [Schiff.] Schmett. Wien, p. 101
(1775).

Ascotis selenaria Hbn. Verz. Bek. Schmett. p. 313 (1826).

Trigonomelea semifusea Warr. Nov. Zool. xi. p. 475 (1904).

1909: May 10,—1 ¢.

Extraordinarily widely distributed in S. and E, Europe, Asia,
and Africa,

Isles

164 PROF: E. B. POULTON ON

Fam. SATURNID &.

EprrHora ATBARINUS Butl.
1909: Oct. 14,—1¢6,29.

LupiA HANSALI Feld.

1908: Oct.29,—1 ©. 1909: Sept.—l 9. Year?: Apr. 27,
—l ¢ (BM).

Fam. ARBELID&.
SELAGENA EUSTRIGATA, sp.n. (PI. IT. fig. 27, 3.)

3. Head and thorax white tinged with rufous, the dorsum of
thorax with long spatulate ¢hocolate-brown scales ; antennze with
the branches rutous; pectus and legs white, the latter with brown
mixed ; abdomen white, the 2nd segment with dorsal tuft of long
spatulate rufous and chocolate-brown scales, the anal tuft tinged
with rufous and with spatulate black-brown scales mixed. Fore
wing creamy white, with fine dark brown pencillings with white
bars on them defined by black and leaving the veins white below
base of costa, from medial part of cell to inner margin, in end of
cell, between veins 5 and 2 to near termen, and forming a wedge-
shaped subterminal patch between veins 8 and 6; a white patch
with some rufous before and beyond it beyond the discocellulars,
and a white spot with some rufous before and beyond it below
vein 2 near its origin. Hind wing silvery, white. Underside
white ; fore wing with the cell and area beyond it to near termen
suffused with brown with some white strize on it; hind wing
with some red-brown striz on medial part of costa.

1908: Sept. 13,—1 ¢ (type); Sept. 14,—1 ¢ (B.M.);
Sept. 18,—1 3s Sept. 20,—1 dg; Sept. 26,—1 5; Oct. 6,—1 g.
1909: Sept..—2 g¢. Hap. 20-24 millim.

SELAGENA ATRIDISCATA Hmpsn.
1909: Oct. 5,—1 ©.
METrARBELA DIODONTA, sp.n. (PI. I. fig. 28, 3.)

3. Head and thorax dark reddish brown mixed with some
grey-white, the frons whiter; palpi black-brown ; pectus and
legs red-brown mixed with some whitish ; abdomen white tinged
with brown and with chocolate-brown dorsal streaks at base and
extremity, the anal tuft tipped with chocolate-brown. Fore wing
white tinged with brown ; a series of black-brown points below
the costa ; a black-brown subbasal point on median nervure with
a slight dentate dark brown line from beyond it toimner margin ;
a very irregular patch of brown suffusion defined by black-brown
on medial part of inner margin, indented above and below by
white spots towards its extremity ; a down-curved black-brown
streak from middle of cell to lower angle with a point beyond it
in the cell and an irregular oblique bilobate mark defined by
black-brown in upper extremity of cell; an oblique strongly.

MOTHS FROM SOMALILAND. 165

dentate dark line from below apex to inner margin at the medial
patch with more prominent blackish teeth between veins 7 and 5
and some dark suffusion beyond it ; a terminal series of geminate
black points. Hind wing silvery white witha very faint brownish
tinge and faint brownish terminal line. Underside white with a
faint brownish tinge and series of small brown spots on costa of
fore wing.

2. Abdomen more suffused with brown; fore wing more
suffused with brown, the streak in lower end of cell absent
and the oblique dentate postmedial line very indistinct; hind
wing strongly tinged with brown.

1908: Sept. 27,—1 3; Sept. 28,13 Sept. 291g; Oct. 1,
—-1 $6; Oct. 4,—1 Q (type); Oct. 11,—1 ¢ (type); Oct. 14,—
1 g¢ (B.M.); Oct. 20,—1 & ; Oct. 23,—1 6 ; Oct. 24,—1 ¢.
1909: Sept..—1 ¢. Hep, 22-26 millim.

METARBELA PERSTRIATA, Sp. n. (PI. IT. fig. 29, 9.)

2. Head, thorax, and abdomen grey-white suffused with
reddish brown. Fore wing creamy white, thickly irrorated with
dark reddish brown and with numerous rather reticulate lines,
formed by dark reddish-brown strive and with obscure dark
brown spots at middle and end of cel], Hind wing and underside
whitish suffused with brown.

1908 : Sept. 27,1 @ (type). Hap. 22 millim.
ARBELODES RUFULA Hmpsn,
1909: Mar. 14,—1 3; Apr. 8—1 ¢:; May 10,—1 g,

Fam. Cossirp#,

AZYGOPHLEPS IncLUSA WI1k.,

1909: May 13,—1°9.

Duomitus Mrsosticta, sp. n. (Pl. I. fig. 30, 3.)

¢. Head, thorax, and abdomen white mixed with dark brown,
the metathorax almost entirely black-brown; palpi and tarsi
black-brown. Fore wing white irrorated with reddish brown,
the medial inner area with a shade formed by thicker irroration,
extending before middle to above vein 1; a discoidal patch formed
by similar irrovation conjoined beyond lower angle of cell to a
similar shade on terminal area, the cell, submedian interspace
except on terminal area, and an oblique postmedial shade trom
costa whiter ; elliptical black-brown medial spots above and below
vein 1; cilia chequered dark brown and white at tips. Hind
wing white, the cilia chequered with dark brown to vein 2.
Underside of fore wing with the inner area white with a brown
spot below middle of cell; hind wing with the costal area irrorated
with brown.

1908: Apr. 29,—1 3; May 4,—1 ¢ (B.M.); Sept. 12,—1 o.
1909: Oct. 7,—1 ¢ (type). Hap. 28-38 millim.

166 PROF. E. B. POULTON ON

Duomirus sTENrPTERA, Sp.n. (Pl. II. fig. 31, 3.)

3. Head and thorax white mixed with red-brown and some
black, the metathorax almost entirely black-brown ; antenn
dark brown; tarsi black ringed with white ; abdomen whitish
suffused and irrorated with red-brown. Fore wing white with
sparse strong black strie, the inner area tinged with rufous to
beyond middle; a wedge-shaped black-brown patch on costal
area from base to near middle; a rather triangular black spot
below end of cell with its upper extremity somewhat produced ;
a black-brown patch at end of cell extending to the costa ; some
small black spots on costa towards apex; a subterminal series of
small more or less distinct clavate dark marks in the interspaces ;
cilia chequered reddish brown and white. Hind wing white, the
cilia with some brown scales mixed. Underside white; fore
wing with small black spots on costa and blackish spot below
end of cell, the terminal area striated with brown; hind wing
with the costal area finely striated with red-brown.

1908: Apr. 27,1 3 (type). 1909: May,—l 3d. Hap. 28-
38 millim.

Dvomirus sIMILLIMA, sp.n. (PI. II. fig. 32, ¢.)

3. Head and thorax dark red-brown mixed with some whitish
and black, the metathorax entirely black-brown; antenne dark
brown; tarsi black ringed with white; abdomen dark red-brown
mixed with whitish at sides towards base. Fore wing grey-white
suffused with red-brown and sparsely striated with strong black
striz, slighter on basal area, the medial area from costa to vein 1,
the postmedial area from costa to vein 2, and the interspaces of
terminal area whiter ; a rather triangular black spot below the
cell; subterminal sevies of slight clavate dark marks in the inter-
spaces ; cilia chequered dark brown and greyish. Hind wing
whitish suffused with brown, the cilia obscurely chequered with
dark brown. Underside of fore wing suffused with brown, the
terminal area striated with brown, series of small black spots on
costa and below terminal part of cell; hind wing whitish, the
costal area suffused with brown, the costa towards apex and
termen with some brown strie.

1908: Sept. 20,—1 ¢ (type); Sept. 23,--1 g. Hap. 24-

28 millim.

Fam. LASIOCAMPIDS&

‘TRICHIURA OBSOLETA Klug.

1908: Oct. 25,--1 5; Oct. 29,1 g. 1909: Mar. 14,—2 ¢;
Apr. 8,--2 6; Apr. 11,--1 9 ; Apr. 15,1 9 ; Oct. 14,--1 ¢,
1 @; Dee. 30, 1 ¢. -1910: Jan. 2,—1 g; Mar. 10,—1 9;
Mar. 13,—1 c.

ANADIASA SIMPLEX Pag.
1909: Apr. 8,—1 o.

MOTHS FROM SOMALILAND. | 167

CHILENA DONALDsoNtI Holl.

1908: Oct. 13,--1 g$. 1909: Feb. 20,—1 ¢; Mar. 26,--1 d;
Mar. 29,—1 g¢ ; Apr. 1,—1 6; Apr. 5,--1 od ; Apr. 6,--1 9 ;
Apr. 10,—4.d6; Apr. 14,—1 3; Apr. 20,—1¢. 1910: Mar. 20,
=I Cie

Fam. Limacopip4&.

CG@NOBASIS CHLORONOTON, sp. n. (PI. II. fig. 35, ¢.)

3. Head and thorax emerald-green ; antennz fulvous yellow ;
palpi fulvous yellow, brownish at sides ; tibiz on inner side and
the tarsi fulvous yellow, the tarsi with brown points on outer
side ; abdomen pale orange-yellow, the sides and ventral surface
whitish at base. Fore wing emerald-green, the costal edge orange-
yellow. Hind wing white tinged with emerald-green, especially
towards termen; the cilia emerald-green. Underside of fore
wing with the costal half fulvous brown, the inner half greenish
white, the termen and cilia green; hind wing pale orange-yellow,
the terminal area tinged with green ; the cilia green.

1908: Oct. 12—1 3. 1909: Apr. 4,—1 ¢ (type); Apr. 8,
—2 ¢ (1 in B.M.); Apr. 9,—1 ¢6; Apr. 101 5; Apr. 20,
—l1¢; Apr. 23—1 3; Apr. 24,—1 ¢ (B.M.); Sept..—l ¢.
Exp. 18-20 millim.

Ca@NOBASIS FULViCORPUS Hmpsn.

1908: Sept. 26,—1 ¢; Oct. 24,—1 d. 1909: Apr. 4,—1 ¢;
Apr. 5,—1 6; Apr. 7,—3 6; Apr. 8,—2 g,1 9.

Genus FEATHERIA, nov.
Type, /. obvia.

Proboscis absent ; palpi obliquely upturned, short, not reaching
to middle of frons, which is smooth ; antenne of male bipectinate
with moderate branches to apex, of female with short branches ;
metathorax with spreading crest ; tibize with the spurs moderate,
the hind tibie with the medial spurs present; abdomen with
rough hair at base of dorsum. Fore wing with the apex rounded,
the termen evenly curved ; veins 2, 3 shortly stalked, 5 from near
angle; 6 from well below upper angle; 7 from just below angle ;
8, 9 stalked; 10,11 from cell. Hind wing with veins 3 and 5
from near angle of cell; 6, 7 stalked; 8 from middle of cell.

FEATHERIA oByIA, sp. n. (Pl. II. fig. 33, 3.)

g 9. Head and thorax grey-white mixed with reddish brown ;
tarsi ringed with white; abdomen grey-white tinged with red-
brown. Fore wing grey-white tinged in parts with reddish brown
and irrorated with dark brown ; a dark brown point at lower angle
of cell, with an oblique brown line from it to inner margin
slightly defined on outer side by whitish followed by a reddish-
brown shade; an oblique dark-brown fascia from apex meeting
the shade beyond the medial line, slightly ineurved below vein 7,

168 PROF. E. B. POULTON ON

where there is a more or less elongate black spot beyond it,
the oblique fascia followed by a whitish shade arising below
apex ; a terminal series of black-brown striz. Hind wing white
suffused with reddish brown ; a fine darker brown terminal line
and fine white line at base of cilia. Underside white tinged with
reddish brown, the costal areas irrorated with brown.

Mandera.—1908: July 16,—1 g ; Aug. 16,—1 ¢; Sept. 26,
—2 g, 19; Sept. 27,—1 2 (type). 1909: Mar, 24,1 ¢g
(type); Apr. 9,—1 ¢ ; Sept. 6,—1°.

Gan Libbah.—1908: June 26,—1 2. Hap. S 22, 2 28

millim.

PARYPHANTA FIMBRIATA Karsch (ARCUILINEA B.-B.).
1909: Mar. 26,—1¢ ; Apr. 10,—1 d.
ScCOTINOCHROA MINOR, sp. n. (PI. II. fig. 34, 3.)

3. Head, thorax, and abdomen bright chestnut mixed with
fulvous yellow and some dark brown; antennz fulvous. Fore
wing bright chestnut irrorated with dark brown and rough
silvery scales; an obscure dark mark below origin of vein 2;
the postmedial area ochreous whitish with a very ill-defined band
of dark and silvery scales, rather maculate to lower angle of cell,
then excurved ; a curved maculate subterminal band of dark and
silvery scales from below costa to vein 2; a dark brown patch at
apex. Hind wing yellow tinged with rufous, the cilia deeper
rufous at tips. Underside yellow, the fore wing suffused with
rufous, the hind wing tinged with rufous.

1909: Apr. 20,—1 ¢ (type). Hap. 18 millim.

GAVARA LEUCOMERA, sp.n. (PI. II. fig. 36, 9.)

2. Head, thorax, and abdomen white, faintly tinged with
rufous; fore tibiz and the tarsi pale brown ringed with white.
Fore wing white tinged with rufous except on terminal area,
which is slightly irrorated with rufous; a rather oblique rufous
antemedial shade from below costa to inner margin; a black
point at lower angle of cell; an indistinct waved rufous line
from lower angle of cell to inner margin; an indistinct sinuous
rufous line from middle of costa to submedian fold above tornus,
then incurved to inner margin; a distinct diffused rufous line
from costa beyond middle to termen at submedian fold, excurved
at middle; a terminal series of rufous strie. Hind wing white
tinged with rufous; a fine rufous terminal line; cilia white.
Underside white, the fore wing suffused with rufous, the hind
wing tinged with rufous.

1909: Apr. 8,—1 9@ (type). Hap. 16 millim,

Fam. THYRIDID&.

RHODONEURA HAMATIPEX, sp.n. (PI. II. fig. 37, 3.)

3S. Head, thorax, and abdomen ochreous suffused with rufous.
Fore wing ochreous tinged with rufous and thickly reticulated

MOTHS FROM SOMALILAND, 169

with rufous strie, browner at costa; a slight antemedial line
forking towards costa and forming a slight fork towards inner
margin; a narrow rather oblique postmedial band formed by two
lines filled in with rufous except towards costa, the imner line
curved inwards to costa and somewhat angled inwards at lower
angle of cell and vein 1, the outer line excurved below costa, the
band somewhat constricted at submedian fold, a reticulate band
formed of double strie beyond it from vein 7 to tornus; an
oblique double line filled in with rufous across apical area from
costa to termen at vein 4. Hind wing ochreous tinged with
rufous and striated with rufous lines; a slightly curved ante-
medial line, dark point just above lower angle of cell, darker
slightly sinuous medial line, two or three faint postmedial lines,
and a more prominent subterminal line oblique to discal fold,
then sinuous to tornus. Underside of fore wing with a streak
formed by black spots and opalescent silvery scales below middle
of cell and a short streak formed by black dashes beyond upper
angle, the medial part of postmedial band, the subterminal band
towards tornus, and the oblique band except at costa prominently
filled in with rufous.

1909: Apr. 11,—1 ¢ ; Oct. 19,—1 ¢ ; Oct. 30,—1 ¢ (type).
Exp. 22 millim. Closely allied to R. squamigera Pag.

Fam. PYRALID®.
Subfam. CRAMBIN.

ANCYLOLOMIA PECTINIFERA Hmpsn.

1909: Mar. 10,—1 9; Mar. 14,—1 2.

SURATTHA SCITULELLUS WI1k.

1908: Sept. 18—1 2; Sept. 20.—1l 92; Sept. 21,—1 9 ;
Sept. 22,—I 9; Sept. 24-2 9; Sept. 26-—1 9. 1909:
Mar. 13,—1 9; Mar. 14,—1 2 (B.M.); Mar. 19-—19; Mar. 22,
—l1 9.

SURATTHA INVECTELLUS WIk.

1908: Aug. 15,—1d; Sept. 14,—1 6d; Sept. 20,—1 ¢; Sept. 24,
—1 ¢; Sept. 26—1 9. 1909: Feb. 15,—1 9; Feb. 16,—2 9
Feb, 23,—1 ¢ ; Mar. 5,—1 ¢ ; Mar. 11,—1 2 (B.M.); Mar. 13
—3 6,492; Mar. 19,—1 5,12 (¢d B.M.); Oct. 14,—1 9.

?
?

Subfam. SCH@NOBIAN.

Genus CALAMOSCHG@NA, nov.

Type, C. ascriptalis.

Proboscis absent; palpi upturned, in male hardly reaching to
middle of frons, in female to vertex of head; maxillary palpi
minute; frons smooth, rounded ; antennz in both sexes laminate
and almost simple ; hind tibize with the outer medial spur minute.
Fore wing with vein 3 from before angle of cell; 4, 5 shortly

170 PROF. E, B, POULTON ON

stalked ; 6, 7, 8, 9 stalked; 10,11 from cell. Hind wing with
veins 3 and 5 from angle of cell, 4 absent; 8 anastomosing
with 7,

CALAMOSCHGNA ASCRIPTALIS, sp. n. (PI. II. fig. 38, 3.)

¢. Head and thorax pale ochreous ; pectus, legs, and abdomen
ochreous white. Fore wing uniform pale ochreous. Hind wing
glossy white, the cilia tinged with ochreous at base. Underside
white ; fore wing with the costal half tinged with ochreous.

2. Head and thorax slightly tinged with rufous: abdomen

more ochreous; hind wing with the termen and cilia at base
tinged with ochreous.

1908: Oct. 6—1 3S (type). 1909: Oct. 12,—1 92 (type).
Exp. 3 20, 2 28 millim.

Subfam. Puycirin.».

STAUDINGERIA SUB-OBLITELLA Rag.
1909: Feb. 15,—1 9.

EUZOPHERA VILLORA Feld. (stRAMANTELLA Rag.).
1909: June 10,—1 9°.

NEPHOPTERYX METAMELANA Hmpsn.
1908: Sept. 16,—1 2; Sept. 18-1 6.

NEPHOPTERYX ? EMUSSATATELLA Rag.
1908: Sept. 14,—1 9°.

NEPHOPTERYX EUGRAPHELLA Rag.
1909: Oct. 11,--1 @.

NEPHOPTERYX SERRATELUA Rag.

1908: Feb. 24,1 9; Sept. 12,—1 9; Sept. 19,—1 9 (B.M.);
Sept. 26—1 9 (B.M.); Oct. 12,—1 9; Oct. 20,—1 9: Oct. 25,
—1 S$ (B.M.); Nov. 18,—1 9. 1909: Jan. 9,—1 ¢@ (BME):
Mar. 13,—1 9; Apr. 16,—1 9(B.M.); Dec. 31,—1 9.

Subfam. ErrpascHian a.

MACALLA PURPUREOPICTA, sp. n. (PI. IL. fig. 39, 9.)

2. Head and thorax pale grey; pectus, legs, and abdomen
cupreous rufous, the last with some blackish at base of dorsum.
Fore wing with large tufts of raised hair-like scales below the
cell before middle and in middle and end of cell; pale greyish
and white and with some dark irroration beyond the cell,
the area below the cell and vein 3 purplish rufous from before
middle to tornus; a blackish antemedial line from cell to inner
margin, slightly angled outwards at submedian fold, the tufts of
scales in the cell grey-brown; blackish streaks on middle of

MOTHS FROM SOMALILAND. 7k

vein 1 and basal half of veins 2 and 3, and a slight streak beyond
upper angle of cell; an oblique black bar from origin of vein 7 to
vein 5 near termen ; a purplish-rufous patch on terminal part of
costa with oblique purplish-rufous bar from it at vein 7 to vein 5
just before termen ; a terminal series of dark strize except towards
tornus ; cilia white tinged with rufous and with a pale brownish
line near tips. Hind wing semihyaline white, the apical area
suffused with red-brown to vein 4; a diffused purplish-red streak
on terminal part of vein 2; a terminal series of red strie, darker
towards apex ; cilia white, tinged with fiery red at base. Under-
side of both wings white, the costal and apical areas red.
1909: Apr. 9,—1 @ (type). Hap. 24 millim.

Subfam. ENDOTRICHIN#.

ENDOTRICHA CONSOBRINALIS Zell.
Hargaisa.—1908 : Oct.,—2 ¢.

Subfam. PyRALIN®.

AGLOSSA INCULTALIS Zell.
1909: Feb. 17,—1 ¢.

AGLOSSA OMMATALIS Hmpsn.
1909: Mar. 8,—1 9.
AGLOSSA BASALIS W1Ik.

1908: Sept. 17,—1 9 ; Sept. 24,—-1 9; Sept. 26,—1 9 ;
Sept. 30,—1 9. 1909: Feb. 11,—1 9.

TEGULIFERA ZONALIS Warren.
1908: Nov. 3,—1 @.

'TEGULIFERA NIGRICINCTALIS Hmpsn.

1908: Sept. 18,—1 ¢ ; Oct. 1,—1 2; Oct. 13,—1 9. 1909:
Apr. 14,—1 9; Apr. 22,—1 ¢.

TYNDIS PROTEANALIS Hmipsn.

1908: Aug. 24,—19; Aug. 27,—2 g; Sept. 3,—1 6d; Sept. 12,
—1 9; Sept. 15,—1 ¢; Sept. 16,—1 ¢; Sept. 18—1 45,1 9;
Sept. 19,—1 ¢ ; Sept. 21,—1 6.1 9; Sept. 22,1 9. 1909:
Mar. 9,—1 9; Mar. 10,—1 9; Mar. 11,—1¢; Mar. 12,—1¢;
Mar. 15,—3 6,19; Mar.18,—2 2; Mar. 19,—3 9; Mar. 20,
—l1 9; Mar. 21,—1 6,19; Mar. 22,4 9; Mar. 24,—1 9;
Mar. 26,—2 2; Mar. 30,--1 g; Apr. 15,—1 2; Oct. 4,—1 9.
Year ?: Sept. 25,—1 ¢.

ZitHA SUBCUPRALIS Zell.

1908: Aug. 24,—1 g. 1909: Feb. 23,—1 6; Feb. 28,—1d;
Mar. 4,—1 6; Mar. 9,—1 ¢.

172 PROF, E, B. POULTON ON

Bosrra yARIANS Butl.

1908: Sept. 13-1 9 ; Sept. 16,—3 9°; Sept. 19,—1 9.
1909: Mar. 26,1 9; Apr. 8,—2 9 ; Apr. 10,—2 9 ; Apr. 14;
—l 2; Apr. 20,—1 ¢@.

Bostra TENEBRALIS Hmpsn.
1908 : Sept. 14,—1 3; Sept. 15,1 g; Sept. 19,—1 ¢.
Bos?ra PYROCHROALIS, sp.n. (PI. IT. fig. 48, 9 >)

Q. Head, tegule, and abdomen whitish tinged with red-brown ;
thorax fiery red. Fore wing fiery red slightly irvorated with
whitish, the costal edge with some dark scales towards base and
alternating whitish and dark brown points on medial area ; ante-
medial line white, slightly excurved below costa, then inwardly
oblique ; postmedial line white, slightly excurved to vein 4, then
slightly meurved ; cilia purple-brown at base, the tips white with
some red at apex. Hind wing white tinged with red-brown; the
cilia purple-brown at base, white at tips.

1909: Mar. 24,—1 9 (type). xp. 16 millim.
DaTTINIA PERSTRIGATA, sp.n. (PI. IT. fig. 40, 3.)

Antenne of male bipectinate, with long branches to near apex.

¢. Head and thorax creamy white more or less tinged with
brown ; antenne with the branches brown; palpi irrorated with
blackish ; abdomen creamy white with dorsal fulvovs-yellow
bands except at base and extremity, the anal tuft with pale
blood-red subdorsal streaks. Fore wing with diffused blackish
streaks below end of cell, above and below submedian fold and
vein 1 to beyond middle, and on each side of veins 5 to 2; a small
black spot in lower angle of cell and slight point in upper angle;
the streaks partly interrupted by traces of a subterminal white
line with blackish points before it on veins 7, 6; the costal half
of wing sometimes irrorated with blackish ; a terminal series of
blackish points ; cilia chequered with pale blood-red. Hind wing
pure white and somewhat semihyaline.

Ab. 1. Fore wing with the streaks on each side of vein 1 and
veins 5 to 2 beyond the cell with pale blood-red mixed, veins 7, 6
with pale blood-red streaks except on terminal area.

2. Thorax strongly tinged with pale blood-red, the fore and
mid tibiz and tarsi suffused with blood-red, abdomen at sides and
anal tuft blood-red ; fore wing with the costa and cilia blood-red,
diffused blood-red fascize above and below vein 1, the streaks on
veins 7, 6 and on each side of veins 5 to 2 blood-red ; an indistinct
obliquely curved waved subterminal blood-red line between veins
7and1; hind wing suffused with brown, the veins towards termen
and cilia suffused with blood-red ; underside suffused with brown,
the costal areas, veins towards termen, and cilia of both wings
blood-red.

1908: July 1,—1 ¢; July 8,1 ¢g; July 19,—1 g; July 24,
Sm 3 uca 2 fo) OAs 2, (BE Me) eA S lee

MOTHS FROM SOMALILAND. eS

Sept. 21,—1 g; Sept. 23,—1 3; Sept. 24,—1 g; Sept. 26,—1 ¢
(type); Sept. 27,—1 ¢; Sept. 29,—1 ¢. 1909: Jan. 18,2 g,
19 (type); Mar. 14,—1 5,29; Apr. 1—1 ¢; Apr. 14,1 6;
May 12,—1 3; May 21,—2:°d; Sept. 21,—1 o; Oct. 22,2 ¢;
Nov. 25,—1 2. 1910: Mar. 6,—1 ¢; Mar. 9,—1 ¢; Mar. 10,
—l1 $6; Mar. 12,—1 ¢ (B.M.). Hap. § 36-40, 2 42 millim.

DarrintaA oRNATA Druce.

1908: Feb. 24—1 ¢; Sept. 24-1 9; Sept. 291 9;
Oct. 3,--1 g¢. 1909: Mar. 14,1 9; Mar. 18,—1-9; Mar. 19,
—l1 2; Mar. 26,—1 9; Apr. 16,—1 9; Oct. 8—1 9; Oct. 11,
ae i On

DATTINIA PERATALIS, sp. n.~ (Pl. II. fig. 41, 3.)

Antenne of male bipectinate, with long branches to two-thirds
length.

3. Head, thorax, and abdomen ochreous tinged with rufous,
the thorax deeper rufous; antennze with the branches brown ;
frons and palpi deep rufous; legs red-brown, the tibiz and tarsi
ringed with whitish. Fore wing with the basal area rufous, the
rest of wing silvery white with a creamy tinge suffused in parts
with rufous; antemedial line creamy white defined on outer side
by rufous and with some black irroration before it, slightly
waved ; the medial area mostly suffused with rufous, with a
creamy-white patch in and beyond the cell extending to costa;
an oblique black-brown discoidal. bar and slight yellowish spot
below end of cell; postmedial line creamy white defined on inner
side by rufous, excurved to vein 3, then incurved, a patch
of blackish scales beyond it at middle; a silvery whitish apical
patch defined by rather diffused black scales. Hind wing
“ochreous white suffused with rufous; cilia with a white line
at base followed by a rufous line. Underside whitish, the
fore wing and costal area of hind wing suffused and irrorated
with red-brown.

1909: Mar. 14,—1 S$ (type). Hap. 14 millim.

DaTTINIA COSTINOTALIS, sp.n. (PI. II. fig. 42, 3.)

Antenne of male ciliated.

3d. Head, thorax, and abdomen grey mixed with reddish
brown and fuscous, the vertex of head whitish; pectus whitish ;
tarsi brown ringed with white; abdomen blackish brown
ventrally except towards base. Fore wing pale brownish grey
slightly irrorated with blackish ; the costal edge black towards
base; subbasal line black defined on outer side by whitish,
angled outwards below the cell and ending at vein 1; ante-
medial line black, oblique to below the cell, then incurved,
a quadrate patch of blackish suffusion beyond it from costa
to median nervure; a slight dark mark at lower angle of cell;
postmedial line blackish, indistinct except towards costa, ex-
curved to vein 4, then oblique and sinuous, an oblique black bar

174 PROF. E. B. POULTON ON

beyond it from costa; a terminal series of slight brown spots ;
cilia brownish white with two fine brown lines through them.
Hind wing semihyaline white; a fine brown terminal line and
slight line through the cilia.

1908: July 31,—1 ¢ (type). Hp. 18 millim.

CLEDEOBIA RADIALIS Hmpsn.

1908: July 17,—1 9; July 24,—1 3. 1909: Mar. 26,—1 9;
Apr. 8,—1 ¢; Dec. 10,—1 9.

Subfam. PYRAvsTiIn&.

ZINCKENIA FASCIALIS Cram.

1908: Sept. 30,—1 9. 1909: Jan. 11,—19; Jan. 13-19;
May 5,—1 o.

SYLEPTA SABINUSALIS WIlk.
1908: Feb. 24,—1 2. 1909: Oct. 7,—1 2; Oct. 14,1 92.

GLYPHODES INDICA Saund.
1908: Oct. 27,—1 ¢. 1909: May 14,—1 ¢.

AGATHODES MUSIVALIS Guen.
1909: May 10,—1 9.

CROCIDOLOMIA BINOTALIS Zell.
Berbera.—1908: Mar. 4,—1 ¢.

HELLULA UNDALIS F.
1909: Jan. 19,—1 9; Mar. 14,—2 9.

SAMEODES OCELLATA, sp. n. (PI. II. fig. 44, 9.)

Fore wing with scale-tooth on inner margin before middle.

@. Head white, the frons with rufous spot, the antenne and
palpi fulvous red ; thorax fulvous red ; pectus and legs white, the
latter tinged with red-brown ; abdomen white dorsally suffused
with rufous. Fore wing fulvous red; a large rounded white
patch with pale red centre from upper angle of cell to inner
margin, its edges slightly waved and a similar but smaller patch
beyond the cell connected with the costa and extending to vein 4.
Hind wing pale rufous. Underside whitish suffused with rufous.

1908: May 28,—19; Sept. 19,—1 9 (type); Sept. 24,19.
Exp. 16 millim.

LEUCINODES ORBONALIS Guen.
1908: Oct. 29,—1 @.

NoMOPHILA NOCTUELLA Schiff.
1908: Nov. 13,—1 ¢.

MOTHS FROM SOMALILAND. 17/5)

PACHYZANCLA PHAOPTERALIS Guen.
1908: Sept. 20,—1 9.

PACHYZANCLA BASALIS W1k.
1908: Feb. 24,1 9. 1909: Apr. 10,—1 9.

PACHYZANCLA BIPUNCTALIS F,
1908: Sept. 30,—2 9; Nov. 24,—1 9.

PHLYCTHZNODES NUDALIS Hiibn.
1908 : Sept. 26,—1 2; Sept. 29,—1 9.

ANTIGASTRA CATALAUNALIS Dup.
1908: Sept. 18—1 @.

NoorDA BLITEALIS WIk.
1909: May 10,— 2 9.

MEcYNA GILVATA F.

Mandera.—1908: Sept. 3,—1 ¢ : Sept. 11,—1 ¢; Nov. 13,
—2 9; Nov. 14,—1 9; Nov. 18,—1 ¢. 1909: Jan. 14,—1 9;
May 9,—1 93; Oct. 19,—1 ¢; Oct. 22,—2 $; Nov. 10,—1 9.

- Gan Libbah.—1908: June 25,—1 ¢.

PIONEA MELANOSTICTALIS, sp.n. (PI. II. fig. 46, ¢.)

’ 6 Q. Head and thorax grey tinged with brown, the vertex of
head white; palpi red-brown, white at base; pectus and legs mostly
white, the fore tibie and tarsi brown ringed with white; abdomen
grey-brown with white segmental rings, the ventral surface white.
Fore wing whitish tinged and irrorated with brown, the costal
area browner; small antemedial black spots on subcostal and
median nervures, vein 1, and above inner margin ; a black point
in the cell towards extremity and discoidal bar; postmedial line
black, dentate to vein 4, then with oblique bar to vein 2, then
retracted to below end of cell and excurved at submedian fold
and slightly above inner margin; a curved series of blackish
points just before termen and a terminal series. Hind wing
whitish suffused with red-brown ; traces of a curved brown post-
medial line; a terminal series of slight brown points; cilia white
with a faint brown line near base.

1908: Sept. 23,—1 9 ; Sept. 27,—1 9,1 ¢ (type); Oct. 11,
—1 9 (B.M.); Nov. 24,-1 9. Hzxp. 16 millim.

PIONEA RUBRITINCTALIS, sp. n. (Pl. II. fig. 45, 2.)

Q. Head and thorax ochreous yellow tinged with rufous; frons
with white lines at sides ; palpi white in front at base; pectus
and legs white; abdomen reddish ochreous, the ventral surface
white. Fore wing ochreous yellow tinged with rufous; traces of
an oblique rather diffused rufous antemedial line ; a more distinct

176 PROF. E. B. POULTON ON

obliquely curved diffused rufous postmedial shade; cilia white at
tips. Hind wing pale reddish ochreous with traces of a rather
diffused curved rufous postmedial line.

1908: Sept. 24,—1 9. 1909: May 10,—1 9 (type). Hap.
18 millim.

PyRAUSTA INCOLORALIS Guen.
1909: May 3,—1 @.

PyYRAUSTA STHENIALIS, sp. n. (Pl. II. fig. 47, 3.)

Mid tibie of male dilated with a fold containing a tuft of long
hair, the hind tibiz with the outer medial spur minute ; abdomen
very long with the anal tuft long.

Head, thorax, and abdomen pure white, the shoulders with grey
stripes ; frons and palpi towards tips tinged with grey. Fore wing
semihyaline white; the costal area suffused with grey; oblique
slightly curved grey postmedial and subterminal lines. Hind
wing semihyaline white with faint curved greyish postmedial and
subterminal lines.

~1908: May 4,—-3 d, 62; May 6,—19; May 28,—16¢;
Sept. 29,—2 9 (1 in B.M.); Oct. 15,—1 93; Oct. 20,—1 ¢ ;
Nov. 13,—2 6 (1 in B.M.); Nov. 18,—1 ¢ (type). 1909:
Apr. 16,—1 ¢ (B.M.). Hap. 22-24 millim.

PYRAUSTA CONISTROTALIS, sp. n. (PI. II. fig. 48, 2.)

9. Head, thorax, and abdomen pale reddish brown tinged with
grey, the vertex of head whitish; palpi rufous, white at base ;
pectus, legs, and ventral surface of abdomen white, the fore legs
brown in front. Fore wing whitish tinged with reddish brown
and thickly irrorated with dark brown, the costal area rather
browner ; antemedial line indistinct, dark, oblique towards costa,
angled outwards at median nervure and vein | and incurved below
the cell and above inner margin ; a minute dark spot in the cell
towards extremity and curved discoidal striga; postmedial line
dark, waved, excurved from below costa to vein 3, then retracted
to lower angle of cell and erect to inner margin ; a terminal series
of small dark spots; cilia with a dark line near base. Hind wing
whitish suffused with brown especially on terminal area; an
indistinct brown postmedial line, excurved from below costa
to vein 2, where it is slightly angled inwards; cilia white with a
dark line near base.

¢. Browner; fore wing with a faint purplish gloss.

1908: Oct. 23,—1 2 (type).

Also in B.M. from Br. E. Africa, N. Kavirondo, Maramas
Distr., lala (Weave), 1 $. Hap. 22 millim.

SCELIODES LAISALIS Wlk.
1908: Sept. 26.—1 9; Nov. 13,--1 9. 1909: Jan.17,—1 9;
Feb. 25,1 9; Oct. 17,—1 ©.

MOTHS FROM SOMALILAND. 177

CoRNIFRONS ALBIDISCALIS, sp. n. (Pl. II. fig. 49, 3d.)

Antenne of male bipectinate with moderate branches to near
apex ; frontal prominence pointed at extremity, its lower edge
produced to a point before extremity.

Head and thorax red-brown mixed with some white ; antennee
ringed brown and white, the branches blackish in male; abdomen
pale red-brown. Fore wing pale red-brown irrorated with darker
brown especially on the veins ; an oblique whitish shade from base
of costa; a narrow white antemedial band defined by dark scales
and with irregularly waved edges ; a small rather elongate white
spot defined by dark scales in middle of cell and a white discoidal
bar also defined by dark scales, its lower extremity somewhat
curved inwards; a narrow white postmedial band defined by
dark scales and with minutely waved edges, angled inwards at
veins 6, 3, 2 and outwards at submedian fold, excurved at middle
and incurved to inner margin; a slight dark terminal line; cilia
white at base followed by a dark line. Hind wing whitish
suffused with brown especially in female; a slight brown spot
at lower angle of cell and indistinct rather diffused curved
subterminal line; cilia white with a brown line near base.

1908: May 4,—1 92 ; Sept. 28,—1 ¢ (type); Oct. 18,—1 ¢;
Nov. 13,—1 9; Nov. 14,—1d¢. 1909: Apr. 5,—1 9; Apr. 7,—
13,29 (1 in B.M.); Apr. 10,—1 g. Hap. 20 millim.

TEGOSTOMA COMPARALIS Hubn.

1908: June 1,—1 9; Sept. 25,—1 9; Oct. 31,1 2. 1909:
May 9,—1 35; May 10,—4 9°.

TEGOSTOMA SUBDITALIS Zell.
1909: May 8,—1 ¢.

TEGOSTOMA BIPARTALIS Hmpsn.
. 5
1908: Aug. 15,—1 9.

NOCTUELIA GLOBULIFERALIS, sp. n. (PI. II. fig. 50, 3.)

3. Head and thorax white mixed with rufous; palpi red-
brown ; fore tibiz with brown bands near extremities ; abdomen
white with slight rufous dorsal bands and streaks on anal tuft.
Fore wing white suffused with rufous; an indistinct rufous sub-
basal line; antemedial line red-brown, excurved above inner
margin, a round white spot defined by red-brown on its outer
side in and below the cell; rounded white spots defined by red-
brown in and below end of cell ; a postmedial white patch defined
by red-brown except above below the costa, intersected by a red-
brown streak on vein 7 and its outer edge indented by a wedge-
shaped red-brown mark on vein 6, a white patch beyond it at
apex and oblique elliptical white spot defined by red-brown below
it ; a dark brown terminal line; cilia white at base with a brown
line near base and brownish tips. Hind wing white ; an elliptical

Proc. Zoou. Soc.—1916, No. XII. 12

178 PROF. E. B. POULTON ON

yellowish discoidal spot defined by brown and with brown line
from it to above inner margin; a brown postmedial line from
costa to vein 5 and elliptical white spot defined by brown between
vein 5 and submedian fold; a dark brown terminal line; cilia
with series of brown striz near base and brownish tips.

1908: Oct. 29,—1 ¢ (type). Hap. 18 millim.

TINEINA.
By Jno. Harriey Durrant, F.E.S.

OLETHREUTIDS.

Kucosma Hb.
EUCOSMA SOMALICA, sp. n.

Antenne fuscous. Palpi whitish ochreous, more or less mixed
with fuscous on the outer side of the median joint. Head
and thorax whitish ochreous, slightly tinged with pink; tegule
brownish fuscous, mixed with reddish. ore wings elongate,
slightly dilated posteriorly, male without costal fold, apex obtuse,
termen nearly straight, slightly oblique ; whitish ochreous, striate
with fuscous and pinkish, with fuscous markings outlined by
shining pearly scales; the markings, which appear more or less
irrorate with whitish owing to some of the scales being tipped
with white, consist of a basal patch, obtusely angled on the cell,
and an irregular central fascia becoming attenuate, or even
obsolete, toward the tornus, this fascia is outwardly connected
with a subapical quadrate spot; the costa is strigulate with
fuscous, there is also a fuscous apical spot and an interrupted
terminal line; cilia whitish ochreous, with a pinkish gloss, tra-
versed by two pale fuscous shade-lines. Hap. al. ¢ 18-272 mm.
Hind wings with 3-4 stalked; fuscous; cilia whitish ochreous,
traversed by two greyish fuscous shade-lines. Abdomen fuscous
with paler transverse lines. Legs whitish ochreous ; tarsi spotted
with blackish.

Type 3 (7248); 2 (7249), Drnt. Det.

1908: Sept. 19,—1¢ ; Sept. 20,—1 2 (type); Oct. 24,—1 ¢
{type). 1909: Jan. 20.—1 9. 1910: Mar. 6,—1 9.

The female is a little darker than the male, the pearly scaling
tending to become leaden, especially toward the tornus.

TINEIDS&.

Nomi, gen. n.
(vopipos, n, ov = conventional.)
Type: Vonima prophanes Drnt.

Antenne 2, with projecting scales on each joint, giving a
serrate appearance, and bipectinate 3, each pectination ciliate ;
basal jomt without pecten. Labial palpi porrect, clothed beneath

MOTHS FROM SOMALILAND. 179

and at end; terminal joint short, concealed. Mazillary palpi and
haustellum obsolete. Head rough-haired. Thorax smooth—
perhaps slightly tufted posteriorly. ore wings elongate, rather
narrow, apex round-pointed, termen rounded, surface with tufts
of raised scales: newration 12 veins, all separate; 7 to apex, 3-4
basally approximate; 1 furcate at base. Hind wings 1, elongate-
ovate, with small transparent space below cubitus near base ;
cilia 3: neuration 8 veins, all separate; 4-7 nearly parallel.
Abdomen vather slender. Legs: posterior tibize long-haired
above.

NOMIMA PROPHANES, Sp. 0.

Antenne fuscous. Palpi yellowish ochreous. Head and
thorax dark brownish fuscous ; face yellowish ochreous. ore
wings cream-ochreous, with a dartle brownish fuscous basal patch,
and with a rather broad patch of the same colow commencing
before the tornus and extending around the termen to the apex ;
the whole wing is ornamented with glistening spots of raised
scales arranged in transverse lines—these raised spots have some
admixture of bluish leaden-metallic, especially on the dark
patches, and on the ochreous part of the wing transverse lines
of pale greyish scaling occur between them ; Cilia shining, dark
brownish fuscous with a purplish gloss, cream-ochreous along
their base and above the apex ; underside suffused with fuscous,
except on a yellowish ochreous apical patch. Hap, al, 21-25 mm.
Hind wings shining, pale grey with brassy sheen, more or less
suffused with dark fuscous above and beneath in some specimens ;
cilia pale yellowish ochreous. Abdomen yellowish ochreous, dusted
with fuscous. Legs yellowish ochreous, tarsi tinged with fuscous.

Type 3 (7253), Drnt. Det.

1908: Sept. 30,—1¢. 1909: Apr. 10,—1 ¢ (type); Apr. 15,
—l1¢.

ACHTHINA, gen. n.

(axBewvds, 4, 6v = irksome.)
Type: Achthina ctenodes Drnt.

Antenne @ bipectinate 2; basal joint without pecten.
Labial palpi vather short, upeurved ; terminal joint very short.
Maaillary palpi and haustellum obsolete. Head rough. VThoras
smooth. ore wings with costa straight, apex round-pointed,
termen and tornus evenly rounded: neuration 12 veins; 7-8
stalked, 8-9 stalked enclosing apex, 10 out of stalk of 7-9: 4-5
closely approximate, connate or short-stalked ; 3 from angle, 2 at
least twice as far from 3 as 3 is from 4; 1 basally fureate. Jind
wings 1, rather short and broad, apex and tornus bluntly rounded :
neur ation 8 veins; 3-5 approximate, 2 remote from 3, 5 bent
over and closely approximate to 4, or 4—5 stalked ; 6-7 stalked or
separate; media to below 6. Abdomen: female moderate, ovi-
positor exserted. Legs: hind tibiz long-haired above.

The male is at present unknowi, and there is some variation in
12%

180 PROF. E. B. POULTON ON

the neuration, but the pectinate antenne and exserted ovipositor
of the female distinguish this genus from its allies.

ACHTHINA CTENODES, Sp. n.

Antenne and palpi cinereous. Head and thorax cinereous,
with some admixture of fuscous. More wings cinereous, with
some admixture of whitish, clearly and distinctly, but irregularly
strigulate with blackish, to the number of about 12 or 13 complete
lines, some reduplicate in part and with paler intermediate lines ;
cilia cinereous with a fuscous line near their base, and another
toward their tips. Hap.al.20mm. Hind wings fuscous; cilia with
a pale line near their base. Abdomen fuscous. Legs cinereous.

Type 2 (7256), Drnt. Det.

1908 : Sept. 18,—1 2; Sept. 27,1 © (type).

Meuasina Bdv.

MELASINA PSEPHOTA, Sp. n.

Antenne whitish ochreous, spotted with fuscous ; male pecti-
nate 5. Palpi moderate, densely scaled; whitish ochreous mixed
with dark fuscous. Head whitish ochreous., Thoraa whitish
mixed with dark fuscous; with a dark fuscous tuft posteriorly.
Fore wings elongate, costa somewhat arched, apex obtuse, termen
obliquely rounded, with 12 veins, all separate; chalky white,
strigulate and shaded with fuscous, and with the transverse mark-
ings more or less continuously edged with blackish ; a fuscous
basal patch, slightly angulate outward on the fold, is edged with
dark fuscous except on the costa and dorsum, there is however a
dark costal spot before the end of the patch with some trace of
dark spots crossing the wing; at one-third from the base a more
or less irregular, outwardly oblique, fuscous fascia crosses the wing,
generally widening out from the costa and narrowing below the
fold, with outward extension above the fold in the direction of a
dark fuscous spot at the end of the cell, occurring on an oblique
fuscous fascia extending, more or less conspicuously, from costa to
tornus—in some specimens this fascia is connected with a fuscous
costal patch preceding the apex; cilia chalky white, with two
fuscous parting lines more or less interrupted by four or five
whitish bars. Hap. al. ¢ 21-24mm., 2? 34mm. Hind wings
pale fuscous; cilia whitish, with a fuscous line near their base.
Abdomen fuscous; female with long exserted ovipositor. Legs
pale fuscous; anterior and median tarsi barred with dark fuscous.

Type 3 (7260); @ (7261), Drnt. Det.

1909: Jan. 19,—16d; Feb. 16,—1 6d; Feb. 17,—1 4; Feb. 19,
—16; Feb. 20,—1 5; Feb. 21,—1d; Feb. 22,14; Feb. 27,—
13; Mar. 9,—1 3; Mar. 10,—1 ¢; Mar. 13,—2d; Mar. 14,—1 9
(type); Mar. 20,—1 g; Mar. 26,—1 ¢ (type).

Closely allied to recondita Drnt., but the hind wings are
distinctly broader and the termen is more erect above vein 3,
where there is a slight angle, not noticeable in recondita which
has the wings narrower and more pointed.

MOTHS FROM SOMALILAND. 181

MELASINA RECONDITA, sp. n.

Antenne pale fuscous; male pectinate 5. Palpt moderate,
densely sealed ; pale fuscous. Head pale fuscous. Thorax whitish
cimereous mixed with dark fuscous; with a dark fuscous tuft
posteriorly. ore wings elongate, rather narrow, termen oblique ;
whitish cinereous, strigulate with blackish, and with pale fuscous
markings more or less continuously edged with blackish; a basal
patch is indicated by fuscous suffusion ; on the costa at one-third
commences an irregular pale fuscous fascia, contracted (sometimes
interrupted) on the cell, thence widening, but becoming narrow
from the fold to the dorsum; this fascia is connected to a
pyriform costal patch of the same colour by a larger pyriform
patch on the dise with some extension toward the tornus—these
markings usually reach to the costa before the apex, but are some-
times disconnected ; a rather conspicuous discal spot, irregular in
outline, occurs at the end of the cell on the fuscous patch, below
a small, oblong, dark-margined fuscous patch; cilia whitish cine-
reous, with seven or eight broad fuscous bars beyond a narrow
fuscous dividing line. Hap. al. 6 22-27 mm.; 2 33-39 mm.
Hind wings fuseous; cilia whitish, with a fuscous line along their
base. Abdomen fuscous; female with long exserted ovipositor.
Legs cinereous ; tarsi barred with fuscous.

Type 3 (7275); 2 (7276), Dint. Det.

1909: Mar. 9,—1¢; Mar. 10,—2 9 (including type); Mar. 11,
—1d; Mar. 12,—2 $; Mar. 13,—5 g; Mar. 15,—1 6; Mar. 17,
—I¢; Mar. 18—1¢; Mar. 19,—1 3, 189 (type); Mar. 21,—
1g; Mar. 26,—292. 1910: Mar. 16,—1<¢.

EXPLANATION OF THE PLATES.

Puavte I.
Fig. | Fig.
1. Estigmene griseata ...........6.4. Q. | 26. Ozarba semitorrida .............. 8.
2. Secusio somaliensis ............... 2. | 27. By CLOUIIESODRE cosadacssosnses Son SE
3. Chloridea albivenata sens adh Pb = WOPOPMIRE cax0ass2s0ea050006 Q.
4. Thalatha melanostrota............ 6. | 29. ah REMUSATCOM- cnet Go
5, Matopo heterochroa ............... é. | 30. y EXOUUACEM.... 60.4 .0004s 2. 3.
6. Acroriesis ignifusa .............. %. | 81. 4, mesozonata ............... 3+
7. Odontoretha featheri ............ 6. | 32. a endoplaga .......--...+..--- be
8. Athetis discopuncta ............... 2. | 33. Hulocastra argyrostrota ......... 3.
9. 4, ectomelena ............... 6. | 34, Aulotarache plumbeogrisea...... 2
10. Ethiopica ignecolora Shae. 30. Constantiodes pyralina ......... é.
11. 5 pheocausta ............ g 36. Hoplotarache ectorrida ......... 3.
12. Pachycoa olivacea.................. Q. | 37. 5 ceruleopicta...... 6.
13. Rabila albiviridis ............... dé. | 38. Tarache mesoleuca Rear aU Ons
14. Acrapex albicostata es SOs 39. 20 MIOGONG ...crveeeeeeerere Qa
15. Huterpiodes pictimargo ......... 6. | 40. Hutelia grisescens..... ............ 3.
16. % croceisticta ......... 6. | 41. Acanthonyx seriopuncta ......... é.
17. Paratwerta nang ........0...0c00s 6. | 42. Cerocala albimacula............... é.
18. Hnispa flavipars .................. 8. | 43. Auchenisa cerwrodes.............. 3.
19. Hublemma eremochroa............ 6, | 44. Authadistis camptogramma ... &.
20. 53 ochricosta ... @, | 45, Catephia pyramidalis ......0.... b+
21. 5 arenostrotda............ 3 AG. Eh pericyina ataxic Que Ole
22. Toana nigrilineata abner tous, Aide 0 PoOviochrod............--- G-
23. Chionoxanthia leucophea ...... 3d. | 48. - mesonephelé .........40. SO»
24. Gdicodia strigipennis ............ 8. | 49. % eurymMelas ...... 6.0.6.0. 3.
25. 5 melanographa ......... 2. | 50. Lyncestis diascota One

182 ON MOTHS FROM SOMALILAND.

Puate II

Fig. Fig.
1. Asplenia rubrescens ......... +... 6- | 26. Victoria sematopeyas ............
2. Tephrias trigonosema ..........-. Q. | 27. Selagenk eustrigata ...............
3. Plecoptera polymorpha ......... g. | 28. Metarbela diodonta ...............
4. Magulaba grisea eB SBR OCT (oho aller o a FOCERSUIPOGIEG oan on0 200006
5. Naarda nigripalpis ............... 6. | 30. Duomitus mesosticta...............
6. Rhynchina endoleuca ek Mine | 31. rr SCENIPCEN A... 6... ce eee eee
Ue 3 perangulata ......... O | 32. = SimMIUMMING ...... 2020000
8 35 albiscripta............ 6. | 33. Featheria obvid .........cccce cece
9. Aclonophlebia inconspicua ...... 6. | 34. Scotinochroa minor .............
10. Serancia discomma ............-+- © 35. Coenobasis chloronoton ............
11. Sesquialtera ridicula ee!) 36. Gavara leucomera .....0 ec. eee ee
12. Cenina tergimacula ............... 6. | 37. Rhodoneura hamatiper .........
13. Tephrina prionogyna Mag) 38. Calamoschena ascriptalis ......
14. Discalma calvifrons.............. 8 39. Macalla purpureopicta .........
15. 0 FOUIGTFOUIScoa0c%ne~ 000300004 6. 40. Dattinia perstrigata 4.
16. Ptychopoda aperta .............. 6. | 41. % PER CLQULS) meer eree trees

17. subtorrida ......... Oe Az 3 costinotalis 1.00.5...
18. Acidalia pyrrhochra .. Q. | 438. Bostra pyrochroalis ...........0...
19. an CUMAG vocce ves cscencesese 9 | 44. Samneodés ocellata ... 660... 2...1.:0-
20. D MANOD vc. serceenseeees.. G. | 45. Pionea rubritinctalis ............
21. Tricentroscelis protrusifrons ... 2 46. » melanostictalis............
eS
2

22. Hucrostes astigmatica ............ 47. Pyrausta sthenialis ...............

23. Hierochthonia featheri ......... . ABs. 35 conistrotalis............
24. Neromia manderensis ............ 9. 49. Cornifrons albidiscalis .........
25. Prasinocyma perpulverata ...... SG. 50. Noctuelia globuliferalis .........

ON THE INTESTINAL TRACT OF MAMMALS. 183:

6. Further Observations on the Intestinal Tract of Mammals.
By P. CHatmers Mircuett, M.A., D.Se., LL.D.,
F.R.S., Seeretary to the Society.

(Received January 31, 1916: Read February 22, 1916.)

(Text-figures 1-30.)

INDEX.

ANATOMY AND MorpPHoLoey : Page
Gut-patterns of Mammals ..................c:0ccceeeeee. 183
Primitive Mammalian gut ...............cceceeceneneeeeees 185
Gut-patterns of Monotremata...................:cc:00000. 189

NIETSDVNEIN, ceccopcosace sancvonascceccaces | LUGO)
identatay sateres scscan eon ee nc LOO
JEISPIRXCONGIGA, cosoooucacaccse bub coo ore canon AO
Proboserd cays peste sees sacra nasce 210
Cetacea: Riis m8 eee cetpe tere
ANE OGY IAM, csedcoc0 ononodene sedeoveca sea.) tLe}
JEDBEISOCEXOINTIE, sasconpsbaneccosnenceceocss 1
Hod ential Weeeiaeie iss. OS eet S223
JENEROMNTORD, Lsgddaccocoscdotenasenaoenvsae 8
(CHRO AUABA, boecdatconadeosedaees bovedboos | EN)
Carnivore er pe eee oe
Prosi sy eae eo"
SEND Tuize ye RMR re alle rai = Seen, a tater Aa ed t/AG212 10)
GenerallComelusions)) cena. oeeeces cece cee ences eae
Systematic Imferences ...............concssssreeecenceecesses 24D
Mist ofiReterencest passes eke ee eee OU)

In this communication I describe the gut-patterns of certain
mammals that I have been able to examine since the publication
of a larger memoir on the Intestinal Tract of Mammals (Mitchell,
1905), and I discuss further the significance of the facts with which
Tamdealing. J adhere to the purpose stated in the introduction
to my memoir, to “limit my observations to a definite set of
facts, hoping that the examination of a continuous series by one
observer, from one point of view, would yield more information
than might be derived from a wider range of work over a smaller
range of animals.” My object was to approach a conception of
the primitive pattern of the mammalian gut, to show how the
complex patterns in the different groups were related to the
primitive pattern, and to discuss how far such relations throw
light on the systematic affinities of the groups.

In certain cases, most common in the lower types of mammals,
there is no difficulty in observing the pattern. When the gut is
severed near the stomach and at the distal end of the rectum,
there remains only to cut the dorsal mesentery from the rectum
to the stomach and to sever the portal vein and mesenteric
arteries ; the whole structure of intestinal tract, mesentery, and
blood-vessels may then be pinned-out on the dissecting-board
and the pattern observed without further trouble. Text-fig. 27

184 DR. P, CHALMERS MITCHELL ON THE

is a reproduction of a photograph kindly taken for me by my
colleague, Mr. D. Seth-Smith, and shows the intestinal tract of
the Elephant-Seal prepared in this way. It will be seen at
once how closely it corresponds with the simplified diagrammatic
drawings of dissections which are the material of the other
text-figures in my former memoir and in this communication.

In most cases, however, and especially where the alimentary
canal is relatively long and thin-walled, or where different
portions differ notably in calibre, regions of the tract belonging
to one morphological part are held in close adherence to regions
belonging to another morphological part. Some of these adhesions
are individual: such are more common in old animals and in
animals loaded with fat or plainly diseased. Others are permanent
structures, invariably present in the members of the species in
which they occur—as, for example, the connections between
the colon and the duodenum which have been named the
eavo-duodenal and the colico-duodenal ligaments, or the attach-
ment of the omentum to the colon. Sometimes, moreover,
blood-vessels belonging to one region of the gut may traverse
the adhering folds of mesentery and supply morphologically
remote regions of the gut.

In extreme cases the secondary adhesions may be stronger
than the primitive mesentery, and large portions of the latter
may have disappeared. Sometimes, therefore, the pattern can be
displayed only after tedious dissection and the cutting of many
structures not easy to distinguish from the primitive mesentery ;
but when the process has been accomplished, the pattern of gut
and primitive mesentery is revealed,

The mode in which the intestinal tract and its mesentery
are folded in the body-cavity, and the secondary adhesions,
pathological or permanent, that are formed, are of great surgical
importance; and many anatomists, for the most part cited in
my former memoir (Mitchell, 1905), have paid attention to them.
Their bias towards secondary phenomena, with consequent over-
looking of the relations of the gut-patterns that I have tried to
work out, has made it impossible to derive a coherent picture of
the morphology of the mammalian gut from their work.

The literature of surgery gives us a clear idea as to how
secondary connections may be established when living membranes
are in juxtaposition, and it is a fair supposition that such
“accidental” structures may have become permanent features
of the anatomy where they were useful. The intestinal tract is
a muscular tube, constantly undergoing strong peristaltic waves
of contraction. Its contents, sometimes liquid, sometimes
strongly charged with gases, sometimes with solid hard lumps,
are seldom quiescent, but partly from the mere action of gravity,
and partly because of peristalsis, subject the wall and the
delicate suspensory apparatus of mesentery with the con-
tained blood-vessels and nerves to sudden and varying strains.
These strains are of relatively little importance when the gut

INTESTINAL TRACT OF MAMMALS. 185

is short, thick-walled, and of even calibre, as, for instance, in
the case of many carnivorous animals. When the gut is rela-
tively long, and when the thickness of its walls and its calibre
vary much in different regions, as is frequently the case in omni-
vorous and herbivorous creatures, the danger from mechanical
strain is greater. The habit of life of the creature also must be
taken into consideration. Animals of placid gait, and aquatic
animals living in a medium of nearly the specific gravity of their
own bodies, subject the contents of their abdominal cavity to the
least possible disturbance. Animals that run and leap, and
especially climbing animals—as the latter are constantly shifting
from a horizontal to an erect posture,—subject the contents
of their abdominal cavity toa maximum strain. As adhesions
may take place between portions of the gut that, although they
belong to different regions, are in close contact, it is plain that we
may expect to find them varying in correlation with the nature
of the food, the structure of the gut, and the habits of the animal.
We see readily how they may have arisen in many groups in-
dependently, and that they thus afford no definite indication
of affinity. Dr. Beddard, in a communication to this Society
(Beddard, 1908, p. 561), has brought together a valuable set of
observations, old and new, on such secondary features of the
‘gut, and would appear to agree with me that they cannot, as he
phrases it, ‘“‘yield accurate classificatory results,’ as he is able to
arrange them in a series of ascending stages, and to show that
these stages, or some of them, occur independently in different
groups.
The Primitive Mammalian Gut.

In text-fig. 1 1 have drawn the primitive type to which
the varied patterns displayed by the gut of mammals (when
the secondary connections have been severed) can be reduced.
The left-hand diagram (A) shows the pattern as it may be seen
in a very young mammalian embryo; the right-hand figure (B)
shows it as it appears in some of the simpler adult ‘animals,
The whole gut from the stomach (S.) to the distal end of
the rectum (R.) is suspended from the dorsal wall by a con-
- tinuous mesentery (Mes.) containing the blood-vessels. It
consists of three definite regions. The proximal region, from
the point marked 1 to the point marked 2, is the duodenal
region; in birds this is usually characterised by the outgrowth
of a long, narrow, single loop, but in mammals more frequently
appears as a bunch of short loops not clearly marked off from the
beginning of the next region. The second region, from the point
marked 2 to the cecum (C.), I have termed Meckel’s tract;
it corresponds, according to the position of the cecum, with
the whole or the proximal portion of the pendant loop of
human embryology, and its apex is fixed in the embryo by the
-umbilieal cord (text-fig. 1 A, M.). As a very rare abnormality
in mammals, a diverticulum, known as Meckel’s diverticulum,

186 DR. P. CHALMERS MITCHELL ON THE

the equivalent of the yolk-sac diverticulum which persists
throughout life in a very large number of the families of
birds, is to be found at the point marked in the embryo by
the attachment of the umbilical cord to the apex of Meckel’s
tract. Meckel’s tract forms the small intestine, and however
the gut may be lengthened it remains a nearly closed loop, the
point (3) where the post-cecal gut passes into the hind-gut.
remaining extremely close to the point (2) where the duodenal
region passes into Meckel’s tract. It is also of some importance
to notice that the lengthening of Meckel’s tract to form the coils
of the small intestine takes place chiefly on the proximal limb of
the primitive tract; in the vast majority of mammals, however
long the small intestine may be, the distal limb of Meckel’s tract.
remains as an almost straight tube running up until it nearly
meets the distal extremity of the duodenal loop. Meckel’s tract
in mammals differs notably from the similar region in birds. In
birds the tract tends to be drawn out into definite minor loops,
the disposition of which forms characteristic patterns in different
groups, and the distal region of the tract, immediately proximal
to the cea, tends to form a specialised loop, folded over, and
secondarily attached to the duodenal loop. The third region
of the gut is what I term the large intestine or hind-gut; it
stretches from the cecum to the anus, and occupies a greater
portion of the antero-posterior axis of the body than the
duodenal region and Meckel’s tract together.

The ceecum of mammals occupies nearly the same morphological
position as the cea of birds. In birds like the Ostrich, and in
mammals like the Kangaroo or Hlephant, where the hind-gut is
relatively long, but little differentiated, the ceca throughout life
occupy almost exactly the position indicated in text-fig. 1 B.
In birds where the rectum is very short, and especially when
the distal portion of Meckel’s tract is prolonged into a loop,
and in mammals such as Carnivores in which the hind-gut is
very short and straight, the ceeca appear to lie more close to.
the rectum. In mammals in which the hind-gut is highly
differentiated, the ceca occur on the straight portion of the
recurrent limb of the pendant loop at a varying distance from
the point marked 3 in text-fig. 1. Thus, when the cecum is
situated distally, the recurrent limb of the pendant loop gives
rise to the distal portion of Meckel’s tract. When, as is more
usual in mammals, the cecum is attached proximad of the distal
end of the pendant loop, the recurrent limb of the latter gives
rise partly to the distal and usually straight portion of Meckel’s
tract, and partly to the proximal portion of the hind-gut. In
birds the ceca are almost invariably paired, but as an individual
abnormality a single cecum has been recorded in several cases
(Plotus, Faleoniformes, Columbee), and in Herons and Baleniceps
a single cecum is the normal condition. In mammals a single
cecum is the normal condition; but there are many anatomical
facts most easily explained as vestiges of a paired condition

INTESTINAL TRACT OF MAMMALS. ; 187

(Mitchell, 1905), and the paired condition is normal in some
Kdentates, Hyracoidea, and Manatus.

The hind-gut of mammals differs notably from that of birds,
inasmuch as it tends to be drawn out into specialised loops
forming patterns characteristic of different groups. A rather
different nomenclature has been applied to these minor loops of
the hind-gut by different anatomists ; I propose in this communi-
cation, following, I believe, the more exact of my predecessors, to
designate these loops by their morphological position. The highest
point of the recurrent limb of the pendant loop, where it bends
round to pass into the primitive straight hind-gut, represents
what in many mammals forms the transverse colon; a loop of

Text-figure 1.

Diagram of the primitive Mammalian Gut-pattern.

A. Ina young embryo. B. Ina simple adult.

S. Cut junction with the stomach. RR. Cut distal extremity of the rectum.
Mes. Dorsal mesentery. M. Attachment of umbilical cord, position of
Meckel’s diverticulum. C. Caecum. 1-2. Duodenal region. 2-3. Meckel’s
tract. 3-4. Hind-gut, 7. e., large intestine and rectum.

the hind-gut to the right, or proximad, of this is an ansa coli
dextra ; a loop to the left, or distad, of this forms an ansa coli
sinistra. The angle between the ascending colon and transverse
colon in human anatomy, on this nomenclature, might be called
a vestigial ansa coli dextra ; the corresponding angle, where the
transverse colon passes Into the descending colon, would be a
vestigial ansa colt sinistra. A loop of the recurrent limb of the
pendant loop, proximad of these and close to the cecum, may be
called a postcecal loop or paraczecal loop.

It will be seen that my conception of the primitive mammalian
gut differs in two respects from that presented by Dr. Beddard
(Beddard, 1908, p.591). First and most important, I regard the

188 DR. P. CHALMERS MITCHELL ON THE

primitive gut as presenting three definite morphological regions :
a proximal and short duodenal region; the pendant loop, a
nearly closed loop, the outgrowth of a very small part of the
original straight gut, and divided by the insertion of the
umbilical cord into proximal and recurrent, or distal, limbs; and
third, the hind-gut, corresponding with a much longer portion
of the original straight gut. Next, it possesses a cecum, or
possibly a pair of ceca, homologous with the paired ceca of
birds. Unless we accept such a constitution of the primitive
or ancestral mammalian gut, we are driven to the much more
difficult view that these very definite subdivisions or parts have
arisen independently in many different groups of mammals.
I infer, therefore, that where a mammalian gut-pattern presents
less specialisation than what, I have described as primitive, the
condition has come about by secondary reduction.

In comparing the more differentiated gut-patterns with the
primitive pattern, I attach little importance to the secondary
connections between proximal and distal regions; and in this
Dr. Beddard appears to agree with me for the most part. The
ease with which the more important of these can be established,
and the apparent independent establishment of them in different
groups, arise from the morphological fact that, as the pendant
loop is nearly closed, the colic region and the attachment of the
cecum are brought very close to the duodenal region.

With regard to the subsidiary loops that may be formed in
different portions of the gut, in mammals particularly in the
hind-gut, I attach more importance to their morphological
positions, and less to whether or no they form what Dr. Beddard
ealls “ fixed” loops. Apparently that author employs two
separate criteria in applying the designation. The proximal
and distal limbs of his “ fixed” loops are held together by
a very narrow expanse of mesentery; this, however, is a
question of degree, and narrow loops are linked by many
gradations with what cannot be described as specialised loops
at all. N ext, “fixed” loops are sometimes bound down by
extrinsic ligaments or secondary attachments ; such are obvious
adaptations, and appear to come into existence independently in
different groups.

Nor do I attach much importance to the presence or absence
of a spiral disposition of loops or regions of the gut. Spirals are
common growth-forms, and however striking they may appear,
there is little reason to suppose that the resemblances they
produce are other than convergent. They are far from constant,
even in individual life. The intestines of the tadpole, which are
long in proportion to the size of the creature, are coiled in a tight
spiral ; the spiral has disappeared in the adult frog, in which the
intestines are shorter in proportion to the whole length. I have
found the intestines of young marsupials coiled in spirals, and
comparison of my own observations with those of others leads me
to believe that the chief subsidiary loop of the hind-gut in Lemurs

INTESTINAL TRACT OF MAMMALS. 189

is disposed sometimes irregularly,sometimes in a spiral. The most
conspicuous spiral arrangement in the mammalian gut, however,
the colic spiral of Ruminants, appears to be constant.

DESCRIPTIONS ARRANGED SYSTEMATICALLY.

Sub-Class MONOTREMATA.

Order MonorreMAtA.

I have already figured the gut-patterns of Ornithorhynchus
and Hehidna (Mitchell, 1905, figs. 1,2), but my material was then
only rather badly preserved spirit examples. By the kindness
of Dr. Colin Mackenzie, who has brought from Australia a
magnificent set of well-preserved examples of Monotremes and
Marsupials, I have now seen several much better examples of
Ornithorhynchus and EKchidna. The gut-patterns of these animals
are rather more alike one another and the general mammalian
type than I was formerly able to make out.

Text-figure 2.

Intestinal tract of Ornithorhynchus anatinus.

S. Cut junction with stomach. R. Distal extremity of rectum at cloaca.
C. Cecum. C.L. Colic loop (ansa coli dextra).

The duodenum in each case is a well-marked loop, and is
attached by a cavo-duodenal ligament to the hind-gut at the
curved portion of the hind-gut where the recurrent limb of the
pendant loop bends round to join the rectal portion. Meckel’s
tract is suspended round the circumference of an expanse of

190 DR. P. CHALMERS MITCHELL ON THE

mesentery which is rather more elongated in Ornithorhynchus
than in Hchidna. The middle mesenteric vein curves through
the mesentery, following Meckel’s tract and giving off numerous
branches to the rather regular minor loops of the tract. The
cecum, which appears to be functionless, is placed very close to
the apex of the pendant loop; so that nearly the whole of the
recurrent limb of the pendant loop is hind-gut. This is rather
an unusual arrangement, but is present in the Sloths among
Edentates and in the Mystacoceti amongst Cetaceans. Before
the recurrent limb reaches the dorsal line it is thrown into a
small bunch of minor loops forming an asi coli dextra, less
numerous, however, than I figured for Hcehidna in my earlier
memoir, and placed much nearer to the dorsal middle line.
Then follows a point at which the hind-gut reaches the duodenal
region, to which it is attached by a secondary ligament.

Text-figure 3.

Intestinal tract of Hchidna hystrix.

Lettering as in text-fig. 2.

The rectal portion of the hind-gut is larger in calibre and is
thrown into very shallow minor loops.

Sub-Class MARSUPIALIA.
Order MARSUPIALIA.
Sub-Order Polyprotodontia.
Family Notoryctide. Votoryctes typhlops (text-fig. 4).

The gut-pattern is extremely simple, showing a divergence
from the primitive condition by degeneration. There is no
distinction between the duodenum and Meckel’s tract, the latter
being thrown into irregular minor loops; there is no cecum,

INTESTINAL TRACT OF MAMMALS, 191

and the delimitation of Meckel’s tract from the hind-gut is not
marked. The mesentery is continuous, and the mesenteric velns
are arranged as simple branches of the main channel.

Text-figure 4.

Intestinal tract of Notoryctes typhlops.

S. Cut proximal end of duodenum. R. Cut distal end of hind-gut.

The mesentery is dotted; the veins are marked in thick black lines.

Family Dasyuride. Thylacinus cynocephalus (text-fig. 5).
Sminthopsis crassicaudata. S. larapinta.

In the Thylacine (text-fig. 5) the pattern does not differ in any
important respect from that of Votoryctes, there being no cecum
and the three regions of the gut not being sharply marked off,
although the grouping of the tributaries of the mesenteric vein
suggests their presence. The calibre of the whole gut is rather
large and approximately the same throughout. The subsidiary
coils of the proximal portion of Meckel’s tract are rather more
numerous than is represented in the figure.

The two species of Sminthopsis showed a pattern almost
identical with that of Motoryctes. Dr. Beddard (P. Z.8. 1908,
p- 561, text-figs. 111 & 113) has described and figured the
intestinal tracts of Antechinomys laniger and Phascogale mac-
donellensis. It is clear that these small Dasyurids display a
gut-pattern in all essential respects identical with that of
Notoryctes. In the example of Phascogale, however, although
apparently full-grown, Meckel’s tract was so simple a loop that

192 DR. P. CHALMERS MITCHELL ON THE

Dr. Beddard compared it with the pendant loop of mammalian
embryology, and was so fortunate as to find a remnant of the
umbilical cord passing to the apex of this loop. I have never
found this structure in any full-grown marsupial, and think that
Dr. Beddard’s example was an individual peculiarity ; but it is
interesting to note that it occurred precisely at the position in
which I always look for it, and its presence confirms the validity
of text-fig. | A as a diagram of the primitive mammalian gut-
pattern.

Text-figure 5.

Intestinal tract of Thylacinus cynocephalus.

Description as in text-fig. 4.

The polyprotodont marsupials display gut-patterns of great
simplicity. In some (Didelphys, Peragale) the condition, is
practically identical with text-fig. 1 B (Mitchell, 1905). In
others, such as those described above, a condition of greater
simplicity has been reached, due to the obliteration of the
distinction between the regions and the loss of the cecum. This
simplicity is to be regarded as secondary, as, otherwise, it would
be necessary to suppose that the distinction into definite regions
and the presence of a definitely placed cecum had been acquired
independently in many different groups.

INTESTINAL TRACT OF MAMMALS. 193

Sub-Order Diprotodontia.

Family Phascolarctide. Phascolomys mitchell (text-figs. 6, 7).
Phascolarctos cinereus (text-fig. 8).

In the Wombat the duodenal region is just distinguishable as
one or two loops proximal to Meckel’s tract. Meckel’s tract_is

Text-figure 6,

Q)
. yy ( ASS

VI-=

Intestinal tract of Phascolomys mitchelli.

C. Cecum. C.L.1. Colic loop (ansa coli dewtra). C.L.2. Colic loop (ansa coli
sinistra). Other references as in text-fig, 4.

very distinct, its proximal portion being broken up into a large
number of regularly disposed minor loops. Distally it joins the
Proc. Zoou. Soc.—1916, No. XIII. 13

194 DR. P. CHALMERS MITCHELL ON THE

expanded proximal portion of the hind-gut, and the cecum lies
on the outer side of the curve at the point of junction. The first
portion of the expanded colon runs up towards the dorsal line,
then follows a large colic loop (C.L. 1), and finally a rectal portion
of smaller calibre, thrown into rather regularly disposed minor
loops (U.L. 2). A secondary connection forms a cecal ligament
attaching the cecum to the small intestine and to the duodenal
region. Another secondary connection forms a strong colico-
duodenal ligament, attaching the colic loop to the duodenal
region. The latter was severed to make it possible to lay out
the gut so as to display its pattern.

The peculiar cecum of the Wombat has been described and
figured by Owen (Owen, 1868, p. 417, fig. 315) and by Flower

Text-figure 7.

Junction of the small intestine, cecum, and hind-gut in Phascolomys mitchelli.

Tle. Cut end of ileum. Col. Cut end of colic loop. V. Vermiform appendage.
C. Cecal pouch at the proximal eud of hind-gut. Part of the wall of the
hind-gut has been removed to show the apertures of the ileum and of the
vermiform appendage on a raised projection. X. Beginning of the solid part
of the appendage.

(Flower, 1872, p. 647), and Flower’s figure has been reproduced
by Oppel (Oppel, 1897, p. 567). Owen’s figure corresponds
exactly with the portion of text-fig. 6 marked C., but it has been
drawn from the other side of the gut (the right side). Flower’s
figure has obviously been drawn from a mounted preparation ; it
also shows the right side, but it has been turned upside down.
It is on a larger scale, and part of the side-wall has been
removed to display the mode of junction of the cecum with the
small intestine and the hind-gut. As Flower’s figure is in a
publication that is not now readily accessible, I reproduce as
text-fig. 7 a drawing from my own dissections. It will be seen

INTESTINAL TRACT OF MAMMALS. 195

that the cecum consists of two portions. There is an upper
elongated papilla (text-fig. 7, V.) attached to the ileum by a
mesentery which is not shown in Owen’s figure. This is the
so-called “‘vermiform appendage” of the Wombat. The free
portion is a solid mass of tissue closely similar to the tissue
composing the human appendix; but the proximal portion,
beginning just at the point (text fig. 7, X.) where the appendage
blends or is embedded in the wall of the gut, is hollow. Owen
regards this as the tip of the large cecum indicated by the
letter C. in text-fig. 7.

In Flower’s figure the wide pouch marked C. in text-fig. 7 is
lettered cecum. The author expresses doubt as to whether or
no the vermiform appendage of Owen is to be regarded as
a remnant of an originally expanded cecum, but does not
refer to the fact that only the proximal portion of the vermiform
appendage is hollow. He describes and figures, however, the
relations of the appendage to the ileum in precisely the form in
which I found them. The apertures of the ileum and of the
colon into the dorsal side of the colon lie close together on
a projection enclosed by a raised lip in such a fashion that it is
impossible to regard the so-called vermiform appendage as a
continuation of the globular proximal end of the colon. The
interpretation that seems to be least doubtful is to regard the
projection marked V. as the true cecum, the greater part of
which has become transformed to a solid vermiforn: appendage.
On this view, the cecal pouch C. is merely one of the ssccu-
lations into which the colic loop is constricted, as Owen pointed
out, by two parallel bands. Owen states that he found another
of these sacculations, close to the terminal one, so well marked
as almost to be regarded as another cecum. In one of the two
examples of the Common Wombat that I dissected, I found
another extremely well-marked sacculation forming a _ cecal
pouch towards the distal extremity of the colic loop. In
dissecting the intestines I came upon it first, and until the
whole pattern was unfolded, and the true cecum in its proper
morphological position displayed, thought that I had found a
Wombat in which the cecum had no vermiform appendage.

The length and complexity of the gut-pattern of the Wombat
is in relation with the rough unnutritious diet of the animal.
The pattern, however, is seen to be a simple elaboration of
the primitive type. Apart from the peculiarity of the cecum,
the most interesting feature is the elaboration of the first
portion of the hind-gut into a colic loop. This loop corresponds
with the similar loop in Phascolarctos (text-fig. 8), and, like it, is
an ansa coli dextra, and differs from the expansion on the hihd-
gut of other large Diprotodonts, e. g. Dendrolagus (text-fig. 9),
which is an ansa colt sinistra.

By the kindness of Dr. Colin Mackenzie, I have been able to
examine the intestinal tract of two well-preserved examples
of the Koala (Phascolarctos cinereus). The duodenal region

13*

196 DR. P. CHALMERS MITCHELL ON THE

is not sharply marked off, although in the diagram (text-fig. 8) °
this want of separation is exaggerated. Meckel’s tract is com-
posed of a number of very closely packed minor loops suspended
at the periphery of an oval expanse of mesentery. Its distal
portion bends sharply up towards the dorsal line, and then
bends downwards as if it had been dragged out of place by
the enormous cecum. The cecum is relatively, and in a full-
sized Koala possibly absolutely, the longest cecum of any
mammal. At its proximal end its cavity is directly. continuous

Text-figure 8:

Intestinal tract of Phascolarctos cinereus.

S. Cut proximal end of duodenum. R. Cut distal end of hind-gut. C.C. Caecum.
C.L.1. Colic loop (ansa coli dextra). C.L.2. Colic loop (ansa coli sinistra).
X.X. Cut ends of cecal blood-vessel.

with that of the hind-gut, and is many times larger than the
cavity of the ileum. It tapers gradually towards its apex. The
ileum opens into the dorsal wall of the cecum, where the latter
is continuous with the hind-gut, by a small round aperture
protected by a raised lip. Dr. Mackenzie called my attention to
a pair of pouches placed symmetrically on the lateral walls of the
gut, just where the cecum joined the hind-gut. These could be
felt before the gut was opened as a pair of thickenings which

INTESTINAL TRACT OF MAMMALS. 197

Dr. Mackenzie had ascertained to consist of lymphoid tissue. On
opening the gut, each pouch was seen to have a wide aperture
towards the hind-gut, the blind apex pointing forwards towards
the apex of the cecum. These pouches suggest strongly the
presence of an original pair of ceca, the apices of which have
fused to form the prodigiously long cecum. The cecum is
supported by a mesentery superficial to the primitive mesentery
and suspending it to the duodenal region. It is represented
as severed in text-fig. 8, and the cut ends of the cecal vein
are marked at X.X.

Immediately distad of the cecum is an enormous colic loop,
very wide in calibre and suspended at the periphery of an
oval expanse of the primitive mesentery, continuous with the
mesentery suspending Meckel’s tract. This portion of the hind-
gut must be taken as an outgrowth of the recurrent limb of the
pendant loop, and is therefore an ansa coli dextra. It is
followed by a stout-walled portion of the gut, rather smaller
in calibre, and curving round from the colic loop to the rectal
portion. It is at this point that the intestinal tract returns to
the dorsal middle line, and a very strong secondary “ligament ”
attaches it to the omentum and to the duodenal region. Distad
of this the calibre of the gut is again reduced, and the rectal
portion is enormously expanded and thrown into a regularly
placed set of minor loops attached to a meso-rectum which is
more semicircular in shape than in the diagram. This expanded
portion of the rectum must be regarded as an ansa coli sinistra.

The gut of the Koala, in relation with the diet of leaves, is very
long and very capacious. It is divided into four regions, nearly
equal in capacity, and each “ bunched up” on an expanse of
mesentery. To display them on a flat diagram they had to be
slightly distorted, as well as unfolded. Comparison of the figures
of the gut-patterns of other marsupials, however, shows that in
the Koala there is only an exaggeration of familiar features,
and the pattern resembles that of the Wombat very closely. It
is interesting to notice that the gut-patterns of the ruminants,
in which also the whole gut has become much enlarged in
correlation with the diet, are strikingly different.

Family Macropodide. Dendrolagus ursinus (text-fig. 9).

The duodenal region passes insensibly into Meckel’s tract,
the latter being thrown into minor folds, which are more closely
set than in the figure. The example that I dissected was very
young; it was born in the Society’s Gardens, but died before it
had left the marsupial pouch of the mother. The coils of
Meckel’s tract were closely packed, and in the undisturbed
condition displayed the double spiral represented in the drawing
(text-fig. 9,1). The unconvoluted distal end of Meckel’s tract
was constricted as it entered the dilated hind-gut between a
normal but rather small cecum (text-fig. 9, C.) and a smaller

198 DR. P. CHALMERS MITCHELL ON THE

exeal pouch of the kind frequent in Macropodide, and probably
the remnant of the other member of an original pair. The distal
portion of the pendant loop then passed up towards the dorsal
middle line without trace of the ansa coli dextra marked C.L, 1
in text-figs. 6 & 8, but the proximal portion of the hind-gut
immediately distad of the pendant loop was thrown into a set of

Text-figure 9.

Intestinal tract of Dendrolagus ursinus.

1. Spiral arrangement of part of the small intestines.
Other lettering as in text-fic. 6.

minor folds, forming together an ansa coli sinistra. This was
supported by the mesorectum and supplied by vessels from the
rectal vein and artery. I found a strong cecal ligament, passing
from the ceecum to the proximal portion of Meckel’s tract, and a
short colico-duodenal ligament from the proximal part of the
colic loop to the duodenal region. These contained no blood-

INTESTINAL TRACT OF MAMMALS, 199

vessels, and were severed before the drawing was made from the
dissection.

In the Diprotodont marsupials the gut-pattern remains in a very
simple condition, but the hind-gut is specially elongated. In all
the examples that I have dissected this elongation affects the
region immediately distad of the pendant loop, and may be in
the form of a few wavy expansions or a more concentrated bunch
of minor loops. These are all supported by a simple expansion
of the mesorectum, and represent gradations from a merely
expanded rectum to what would be regarded as a definite ansa
coli sinistra. They are marked C.L. in the figures of Diprotodonts
given in my former memoir (Mitchell, 1905, figs. 6, 7, & 8) and
C.L.2 in the figures of this communication. Examination of
the Wombat and of the Koala have enabled me to ascertain that
in these animals another “colic loop” is present. This is marked
C.L. 1 in text-figs. 6 & 8, is developed on the distal limb of the’
pendant loop, and represents an ansa coli dextra.

Sub-Class MONODELPHIA.
(EDENTATA.)
Order TUBULIDENTATA.

Family Orycteropodide. Orycteropus capensis (text-fig. 10).

In my former communication (Mitchell, 1905) I had to depend
on a description given by Flower. Since then I have had the
opportunity of dissecting the intestinal tract of an Aard-vark.
The proximal part of the gut is marked off as a duodenal region
from Meckel’s tract. The proximal part of the latter is a tube of
nearly even calibre and of very great length (nearly thirty feet),
thrown into minor loops arranged round an oval expanse of
mesentery and corresponding with the proximal limb and apex
of the pendant loop. The first portion of the recurrent limb is
nearly straight.- The whole tract is drained by the middle
mesenteric vein, which curves round the mesentery, receiving
numerous tributaries from the minor loops.

Meckel’s tract opens into a relatively large cecum, the proximal
portion of which is expanded and globular. On opening the
cecum the ileo-cecal aperture is seen to lie on the summit of a
projecting process surrounded by a circular lip that may contract
so as to occlude the aperture. A prominent ridge or flap in the
wall of the cecum passes from the proximal extremity of
the hind-gut in the direction of the ileo-ceecal aperture, and
suggests a former division of the cecum into two cecal pouches.

The first portion of the hind-gut is much expanded and
slightly sacculated. It corresponds with the distal end of
the distal limb of the pendant loop. The gut, after reaching
the point nearest to the duodenum, bends sharply backwards,
and is then expanded to form first a definite wide loop and
then a set of minor loops, finally ending in a short straight

200 DR. P. CHALMERS MITCHELL ON THE

rectum. The wide loop appears to belong to the part of the
hind-gut distad of the pendant loop, and therefore represents an
ansa coli simstra. As Flower has pointed out (Flower, 1872),
the total length of the hind-gut is only about seven feet. Czcal
and colico-duodenal ligaments are both present, but have been
removed before the diagram was made.

Text-figure 10.

Intestinal tract of Orycteropus capensis.

Lettering as in text-fig. 6.

The gut-pattern of Orycteropus, except for the specialisation of
the hind-gut, has not moved far from the primitive condition, the
two chief changes being the lengthening of Meckel’s tract and of
the hind-gut.

Order PHOLIDOTA.

Family Manide. Manis tricuspis.

I have already figured the intestinal tract of the White-bellied
Pangolin (Mitchell, 1905, fig. 9). I have had the opportunity of
examining another example of this mammal. The pattern was
in all essential respects identical with that of the former example,
but the duodenal loop was not so distinctly marked off from
Meckel’s tract, and the subsidiary coils of the latter were rela-
tively larger and more numerous than in my figure; the tract was
very much longer than the hind-gut. A small colico-duodenal
ligament was present, but when that has been removed, as in the
figure, the primitive mesentery is seen to be complete.

INTESTINAL TRACT OF MAMMALS. 201

Order XENARTHRA.
Family Myrmecophagide. Zamandua tetradactyla.

I have been able to examine another example of the Tamandua
Ant-eater. The duodenal region and Meckel’s tract were almost
exactly as represented in the figure I formerly gave (Mitchell,
1905, fig. 11), except that the minor loops of the tract were
rather more numerous and more thickly set. The distal end of
the tract entered the expanded proximal end of the hind-gut
between a well-marked pair of shallow pouches corresponding
with, but not so elongated as, the pair of ceca in the Armadillo
(text-fig. 11, C.). The hind-gut was relatively rather longer, and
not quite so large in calibre. It displayed a colic loop attached
to the duodenal region by a colico-duodenal ligament, but distad
of the pendant loop and corresponding with an ansa coli sinistra.

Family Dasypodidee. Dasypus villosus (text-fig. 11).

I have been able to examine the alimentary tract in a very
young example, little more than a foetus, of the Hairy Armadillo.
The duodemal region was represented by two proximal loops not
well separated from Meckel’s tract.

Text-figure 11.

Intestinal tract of very young Dasypus villosus.

S. Cut end of the gut next the stomach; R. Id., next the rectum. C. Paired
ceca. C.L. 2. Colic loop (ansa coli sinistra).

Meckel’s tract was very long, and was suspended on an
elongated fold of mesentery. The proximal limb of the loop thus

202 DR. P. CHALMERS MITCHELL ON THE

formed was broken up into a numerous set of closely disposed
minor loops. The distal or recurrent limb was straight for the
greater part of its length, and as it approached the dorsal line,
entered the expanded hind-gut between a pair of ceca relatively
longer than in the adult and disposed on the right and left sides
of the gut. The hind-gut distad of the pendant loop had a
distinct colic loop, attached to the duodenal region by a ligament,
removed before the figure was drawn.

The patterns of the intestinal tracts of the Tubulidentata,
Pholidota, and Xenarthra afford no evidence in favour of the
existence of a super-order ‘‘ Kdentata.” Such resemblances as
they present are best explained as a common inheritance from
the primitive type, and so afford no evidence of affinity. In the
Xenarthra, the most characteristic features are the existence of
paired ceca, which seem to be more conspicuous in the young
than in the adult, and the tendency to a great elongation of
the loop formed by Meckel’s tract and the proximal portion of the
hind-gut, a tendency which is better marked in some of the
examples described in my former memoir, than in the young
Armadillo figured here. The hind-gut varies considerably both
in the different groups and even individually. The distal limb
of the pendant loop always approaches the duodenum closely, and
distad of this the hind-gut may pass nearly straight back to the
rectum, may form a shallow, or a well-marked and complex loop.
In both Marsupials and Edentates, the hind-gut appears to be
still in a variable or almost experimental stage.

Order HyracoipEa. Dendrohyrax dorsalis (text-fig. 12).

The pattern of the intestinal tract of the Hyracoidea is the
most remarkable to be found amongst mammals, and deserves
special attention, because of the difficulty that has been found in
assigning its due place to the Order amongst the mammalian
Orders. I have already described and figured (Mitchell, 1905,
p. 461) the intestinal tract of Hyrax capensis; since then I have
been able to examine an adult example of Dendrohyrax dorsalis
and another very young example of H. capensis, and to compare
my own observations and interpretations with those of Dr. Beddard
(Beddard, 1908 and 1909). The pattern of the tract of the 'Tree-
hyrax (text-fig. 12), when the secondary connections have been
severed and the tract laid out aecording to the method I pursue,
corresponds in all essential respects with that of other Hyra-
coids. As Owen long ago (Owen, 1832) correctly stated, the
whole tract, from the duodenum to the distal extremity of the
rectum, is suspended by the primitive mesentery from the dorsal
wall of the body-cavity. In Dendrohyrax I found interruption
in the proximal part of the mesocolon (extending from the point
marked X in the text-figure towards the recurrent limb of the
pendant loop), a gap that I did not notice in D. capensis. The
duodenal region is a distinct loop, well separated from Meckel’s
tract.

INTESTINAL TRACT OF MAMMALS. 203

The proximal portion of Meckel’s tract, as is usually the case
in mammals, is sub-divided into a number of closely-set irregular
loops. Then follows the large cecal pouch, assumed by most
authors to be the representative of the normal mammalian
cecum, but which I regard as peculiar to Hyraw (text-fig. 12,
A.C.). The gut as it leaves this is much expanded and closely
adherent to the wall of the cecum. It then runs a nearly

Text-figure 12.

Intestinal tract of Dendrohyrax dorsalis.

S. Cut end nearest the stomach. R. Cut end nearest anus. A.C. Accessory or
median cecum. C.1, C.2. Paired ceca. C.3. Fourth cecum. C.L.2. Colic
loop (ansa coli sinistra). XX. Severed ends of rectal vein. The portion of
the recto-colic mesentery edged with a broken line is where the mesentery was
cut; the more proximal portion edged with an unbroken line was free from the

dorsal bedy-wall.

straight course parallel with the long axis of the cecum, to
which it is bound by a fold of mesentery, and bearing on its
morphologically ventral or larger curvature a much smaller
cecal pouch (text-fig. 12, C. 3), noted by Lonsky (Lonsky, 1903)
and confirmed by Beddard, and now by myself as present in
Dendrohyrax, absent in H. capensis. The tract now passes up-
wards towards the dorsal middle line, forming what I take to be

204 DR. P. CHALMERS MITCHELL ON THE

the recurrent limb of mammals generally, and bearing on this a
symmetrically placed pair of conical ceeca, which I take to be the
representatives of the normal mammalian ceca, paired as they
are in some Edentates and in the Manatee. Distally the large
intestine forms first a colic loop, thrown into minor folds, from
its position to be regarded as an ansa coli sinistra, and a rather
long rectal portion. The posterior mesenteric vein, supplying
the distal portion of the hind-gut, has to be severed in order to
lay out the intestinal tract in the fashion of this memoir, and its
cut ends are indicated at XX, in text-fig. 12.

The difficulty in interpreting the gut-pattern of Hyra# comes
about from the presence of the cecal pouches. The small pouch,
marked C. 3 in the diagram, appears to be more due to the con-
traction of the gut immediately proximad of it than to any
special outgrowth of the gut itself, and as it is absent in at least
one species of Hyrax, I regard it as a character without morpho-
logical significance. The very large thin-walled pouch marked
A.C. is present in all the species that have been examined. It
is a large thin-walled sac somewhat puckered by two bands
of muscle which, when it is fully expanded, give it an almost
bi-lobed appearance, somewhat exaggerated in the figure of
Hyrax capensis in vay earlier memoir (Mitchell, 1905, p. 461).
The entrance and the exit of the gut lie close together at the
proximal end.

The entrance of the gut into the accessory cecum is protected
by a raised lip. George (1874, pl. 13. fig. 3), who regarded the
accessory cecum as the true cecum, calls this entrance of the
gut into it the ileo-cecal valve, and figures it as guarded by a
flap so placed as to prevent the passage of the contents of the
fore-gut into the cecum. I found no trace of such a structure,
and I do not understand how, if it were present, it could act.
On the other hand, the arrangement I found, by the contraction
of the lip, would prevent the regurgitation of the contents of
the cecum into the proximal part of the intestinal tract. The
aperture of exit leading to the distal portion of the gut is wider,
and is surrounded by a shallower lip. The portion of the intes-
tine into which it leads is closely adherent to the wall of the
cecum, and the cavity is at first slightly convoluted, forming
what might be described as a separate chamber of the cecum,
but in H. dorsalis this is not so well marked as in the figure
given by George (1874, pl. 13. fig. 4). There is a general
resemblance between this cecum and the normal ceca of those
mammals in which the cecum is capacious and relatively short.
The normal cecum of mammals, however, always appears to
be a forward continuation of the hind-gut, the one cavity being
directly continuous with the other in the simplest fashion, except
in those cases in which it is slightly complicated by vestiges of the
presence of the second cecum of an original pair. This is unlike
the complicated relation of the unpaired cecum of Hyrax to the
gut that leaves it. A comparison has been made between this

INTESTINAL TRACT OF MAMMALS. 205

cecum of Hyrav and the normal cecum of the Rhinoceros. I
find none but the most general and vague resemblance. The
cecum of Hyrav is irregular and varying in shape, supported by

Text-figure 13.

The ceca of Hyrax dorsalis.

I, Anterior or accessory cecum. II. Paired or distal ceca. S. Cut end of intes-
tinal tract towards stomach. H. Cut end of intestinal tract towards anus.

e W. Cut edge of gut where a portion of the wall has been removed to display
the interior. A.C. Accessory or anterior cecum. C.3. Cecal pouch distad of
A.C. C.1, C.2. Paired ceca. En. Entrance, Ex. Exit of gut.

two bands of muscle, which in certain conditions of distension
give it an almost bi-lobed shape, communicates with the leaving
portion of gut in a complicated fashion, and has nearly fluid

2.06 DR. P. CHALMERS MITCHELL ON THE

contents *. The cecum of the Rhinoceros is a short cone taper-
ing to a point and regularly sacculated along three bands of
muscle, communicates with the hind-gut, of which it appears to
be the forward continuation, by a simple wide aperture, and its
normal contents are solid.

The paired ceca are conical outgrowths, placed symmetrically
on the sides of the hind-gut. As shown in the figure (text-
fig. 13, IL.), their cavities are widely continuous with that of the
hind-gut, and what I regard as the distal end of the ileum
enters the hind-gut exactly in the middle line between them.
In D. dorsalis the ileum is lined by longitudinal lappets which
cease abruptly between the ceca, the ning membrane of these
being smooth, and that of the hind-gut studded with filiform
papille. The contents of the caeca consist of fecal matter of the
same consistency and appearance as that in the hind-gut. Some
confusion has crept into the literature regarding the orientation
of the paired ceca. Asin the case of the colic ceea of birds
and of mammals, whether there be a single cecum or a pair,
the ceca are the forward continuations of the hind-gut, and
their apices are directed forwards, parallel with the ileum,
towards the proximal extremity of the whole gut. As, however,
the tract hes folded within the body-cavity, in the undisturbed
condition, the portion of the gut to which the ceca are
attached ascends froin the distal and ventral region of the
body towards the dorsal and anterior middle line, it may be said,
in the phrase of Kaulla (Kaulla, 1830), that the ceeca apice pelvem
spectant. The paired ceca, in fact, he on the recurrent limb of
the pendant loop, the position in which the true ceca of all
mammals le. This morphological position, which in my opinion
is sufficient to identify the paired ceca of the Hyracoidea as the
homologue of the true ceca of mammals, is quite apparent if the
various diagrams I have given in this memoir, and in my earlier
memoir, be compared. But the homology is equally plain from
another consideration. When the abdomen of any mammal is
opened, the cecum, if it exist, is found with its attachment to
the gut towards the right side of the body, more anteriorly or
posteriorly placed according to its place on the recurrent limb of
the original pendant loop. If the cecum be very large, and
especially when it is long and coiled, 1 may extend towards the
left side of the body, reaching well across the middle line. If it
be very small, its position on the right side is obvious. Asa
supposed resemblance between the unpaired cecum of Ayrax
and the normal mammalian cecum of the Rhinoceros has been
alleged against the homology I make, I may refer to the figures
of the undisturbed abdominal viscera of the Rhinoceros given
by Garrod (Garrod, 1873, fig. 5) (Beddard & Treves, 1887,

* From observations on a living Hyrax, which was in my possession for nearly
eighteen months, I infer that the contents of the intestines may pass directly from
the aperture of entrance to the aperture of exit of the accessory czecum, and that the

latter gradually fills with a fluid and is discharged at infrequent intervals (usually
about fortnightly), apart from the normal daily defiecation of solid feces.

INTESTINAL TRACT OF MAMMALS. 207

fig. 2). In these drawings the position of the cecum on the
right side is well shown. Dr. Beddard’s own generalised dia-
grams of the mammalian gut (Beddard, 1908, text-figs. 122 &
123) show the same point. Now, if Dr. Beddard’s own diagram
of the alimentary tract of H. capensis (Beddard, 1908, text-
fig. 115) be examined, it will be seen that he represents (and all
my observations confirm him on this point) the paired ceca in
the undisturbed condition as lying on the right side, in the true
position of the normal mammalian cecum, with which he does
not homologise them, and the unpaired cecum as attached to the
gut nearly in the middle line, much to the left of the paired
cca, and therefore in a position in which the normal mammalian
cecum never lies. As a matter of fact, the accessory cecum
of the Hyracoidea, both in the young and the adult, lies in
a region of the body-cavity always occupied in mammals by the
coils of the small intestine.

Although Dr. Beddard (1908, p. 595) makes the general state-
ment that the series of facts (@ofation of the out in the body-
cavity, mesenterial attachments, formation oF ‘fixed ” loops)
cannot yield any accurate classificatory results, he appears to
rely on precisely such facts in his endeavour to show that the
unpaired cecum of Hyraxz is homolegous with the normal cecum
of mammals, and that the intestinal tract of the Hyracoidea is
to be associated with that of the Perissodactyle Ungulates. As
he himself has shown conclusively, rotation of the gut occurs in
almost every group of mammals, and therefore its presence, or
even the stage to which it has reached, does not assist us in the
attempt to detect relationships. I have already (supra, p. 184)
shown that it is necessary to distinguish carefully (a point that
Dr. Beddard has overlooked) between the secondary connections
and the primitive mesentery, as the former are almost certainly
convergent adaptations. Even assuming, however, that the
ligaments might yield evidence of affinity, those that are present
in the Hyracoidea do not support Dr. Beddard’s argument. <A
strong wide ligament attaches the unpaired ceecum to the portion
of the gut which leaves the cecum. This is more extensive in
HZ, capensis (Beddard, 1908, text-fig. 115, 1) than in D. dorsalis,
in which it extends no further than the additional small cecal
pouch (text-fig. 12, C. 3) present in that species. Dr, Beddard,
in directing attention to this, points out that the single cecum
of mammals, however small, is usually, possibly invariably,
attached to the adjacent wall of the gut by such a ligament. It
happens, however, that the mesentery of the true cecum in other
mammals passes between the true cacum and the ileum, that is
to say, the portion of the gut entering, not leaving the cecum.
I do not know of any exception to this relationship, which is in
correspondence with the appearance that the cecum presents of
-being an anteriorly directed outgrowth of the hind- gut, running
forwards roughly parallel with theileum. This nor sna mesentery,
stretching between the cecum and the ileum, is absent in the

208 DR. P. CHALMERS MITCHELL ON THE

case of the unpaired cecum of Hyraw, yet present in Perissodac-
tyles, as in most other mammals. That there is in Perisso-
dactyles (see infra, p. 222) also an adventitious set of fibres
binding the true cecum to the proximal end of the hind-gut,
affords no indication of affinity. Another secondary ligament
stretches from the duodenal region to the portion of the gut
immediately distad of the paired ceca of Hyrax (Beddard, 1908,
text-fig. 115, ¢.d.). The possibility of this attachment being
formed depends, in my opinion, on the fact that at this point
the recurrent limb of the pendant loop nearly reaches the dorsal
middle line, and therefore approaches the duodenum very closely.
If any importance can be attached to its presence, it clearly
marks the region just distad of the paired ceca as the beginning
of the hind-gut, and corroborates my orientation of the gut. A
third secondary ligament well developed in the Hyracoidea is that
between the omentum and the transverse colon (Beddard, 1908,
text-fig. 115, O.). This also, so far as any significance can be
attached to its presence, identifies this portion of the gut, distad
of the paired ceca, and indicates the homology of these organs
with the normal mammalian cecum. Dr. Beddard himself sees
the weight of this objection to his argument, but endeavours to
get out of the difficulty by discussing the varying disposition of
the corresponding attachments in different Rodents. When one
is trying to prove the affinity of Hyrax with the Rhinoceros on
the ground of the attachment of certain ligaments, the argument
does not appear to be much strengthened by showing that these
attachments are not the same in Dasyprocta as in other Rodents.

So far as I am able to follow it, Dr. Beddard’s third point,
relating to the presence of an ansa paracecalis in Hyrax com-
parable with the ansa paracecalis of Perissodactyles is uncon-
vincing. The portion of gut (Beddard, 1908, text-fig. 113, p.a.)
which he thus designates in Hyrax, just distad of the un-
paired cecum, is plainly extremely different from the huge and
extremely definite colic loop, consisting of a closely applied
proximal and distal limb of very wide calibre, held together by a
very narrow expanse of the primitive mesentery, which forms,
perhaps, the most characteristic feature of the gut-pattern of the
Tapirs, Horses, and Rhinoceros (Mitchell, 1905, figs. 23-25, C.L.,
and text-fig. 20, infra). As it happened, I found no definite
structure comparable with the loop figured by Beddard in
H. capensis or in D. dorsalis. If any comparison with the colic
loop of Perissodactyles were to be made, on the assumption
that the unpaired cecum of Hyrax is identical with the cecum
of Perissodactyles, the analogue would be the whole expanse of
the gut from the unpaired cecum to the point where the recurrent
limb approaches the duodenum.

To sum up. If the accessory cecum were absent, anatomists
would have found no difficulty in identifying the paired ceca of °
Hyracoids with the normal mammalian cecum, a structure
which, although usually unpaired, frequently shows vestiges of a
primitively paired -condition, and less frequently is actually

INTESTINAL TRACT OF MAMMALS. 209

paired. In their structure, morphological position on the gut,
position as seen when the abdominal cavity is opened, and
attachments, they correspond with the normal mammalian
cecum. ‘The accessory cecum of Hyracoidea differs from the
normal mammalian cecum in structure, morphological position on
the gut, position in the undisturbed body-cavity, and attach-
ments. The attempt, based on minute details of structure, to.
identify the unpaired cecum of Hyracoids with the unpaired
cecum of a Perissodactyle such as the Rhinoceros, makes the
presence of paired ceca still more inexplicable. Jadhere, there-
fore, to my identification of the paired ceca of Hyracoidea with
the normal mammalian cecum. Owen (Owen, 1832) definitely
compared the paired ceca of Hyrax with the paired ceca of
Edentates and of birds, and the unpaired cecum with the
“additional single cecum, anterior to these, found only in a few
species (of Birds).” his appears to be the most reasonable
interpretation of the facts. J am unaware of any reason for
refusing to identify the paired ceca of Hdentates (and of the
Manatee) with the normal mammalian structure, and I have
shown good reason for identifying the normal ceca of birds with
the mammalian cecum or ceca. I have shown (Mitchell, 1901)
that what Owen calls the “anterior ceecum, found only ina few
species,” which, of course, is the remnant of the yolk-sac, is of
frequent occurrence in adult birds, that its constant presence is a
character of many groups, and that in certain cases (Mitchell,
1903) it is transformed from a vestigial structure to a well-
marked glandular organ. ‘The corresponding structure in mam-
mals, known as Meckel’s diverticulum, is a rare abnormality,
but has been recorded as occurring in just over 2 per cent. of
human bodies. It is a diverticulum of the small intestines
lying almost exactly in the region where the unpaired cecum of
Hyrax is found. If this identification be correct, the unpaired
cecum, obviously functional in the adult Hyracoids, has acquired
an importance that is unknown in any other group; but this
is a supposition less difficult than the view that the Hyracoids
display a loop of the gut identical with that of Perissodactyles.
generally, a cecum corresponding in minute detail with the
eexcum of the Rhinoceros and paired ceca peculiar to them and
the Edentates.

Examination of the intestinal tract of D. dorsalis, and con-
sideration of the points raised by Dr. Beddard, therefore, confirm
the view I stated formerly (Mitchell, 1905, p. 463). The general
pattern of the intestinal tract of the Hyracoids suggests no
affinity with the patterns exhibited by Rodents and Ungulates.
The simple duodenum, the nearly circular Meckel’s tract, and
the hind-gut* divided into a simple colon and rectum merely

* Tt is, of course, plain that by “hind-gut” I imply the region distad of the
paired czeca, as I reckon the part of the gut between these and the unpaired cecum
as part of the small intestine. When Beddard (1908, p. 583) stated that my de-
scription of the hind-gut was “ incorrect’ he was merely restating his belief that the
unpaired czecum was the true cecum, and that all the gut distad of this, including
what he took to be a paraczecal loop and the paired czeca, was hind-gut.

Proc. Zoou. Soc.—1916, No. XIV. 14

210 DR. P. CHALMERS MITCHELL ON THE

conform with the general mammalian plan. ‘The presence of the
paired ceca, on my view that paired ceca are a primitive mnam-
malian feature, does not help us with the placing of the group.
The most striking resemblances are with the patterns displayed
by the Edentate group Xenarthra and the Manatee among the
Sirenia. But it must be remembered that the common possession

of a primitive simplicity is no guide to affinity.

Order ProposcrpEa. LHlephas maximus (text-fig. 14).

I have been able to examine the intestines of a young Indian
Hlephant, and J find that the pattern, in all essential respects, is
identical with what I have already figured for the African

Text-figure 14.

@

fw Ls
Intestinal tract of Hlephas maximus.

S. Cut end of gut next stomach. R. Cut end of gut next anus. C. Cxcun.
C.L. 2. Colic loop (ansa coli sinistra).

Elephant (Mitchell, 1905, fig. 16). There is a separate duo-
denum; Meckel’s tract is supported on a nearly circular expanse
of mesentery, its proximal portion being thrown into numerous
minor loops, and its distal portion, forming the first part of the
recurrent limb, is inserted to the dorsal edge of a moderately
large conical cecum. The hind-gut is not much shorter than the
fore-gut, is of larger calibre, and thrown into comparatively

INTESTINAL TRACY OF MAMMALS. 211

large minor loops with a very short straight rectum. Although
the primitive mesentery suspending the whole length of the
intestinal tract is continuous, a strong secondary connection
forming a cavo-duodenal ligament attaches the proximal part of
the colon to the duodenal region, and has to be severed before
the gut can be laid out to display its pattern. The inner dorsal
wall of the cecum displayed a median fold, running along the
dorsal wall somewhat in the fashion of the typhlosole of the
earthworm. It is conceivable that this may indicate an original
paired condition. In the case of these very simple patterns, it
is rather easy to see resemblances which may have little signi-
ficance, but it is undoubtedly notable that the pattern of the
Proboscidean gut in no way suggests that of the true Ungulates,
and very strongly recalls that of the Sirenia (Mitchell, 1905,
ines, 15).

Order CEeracka.

Sub-Order Mystacoceti. Balenoptera physalus (text-
fig. 15).

By the kindness of Mr. J. Erik Hamilton, I have had the
opportunity of dissecting a young embryo of the Common
Rorqual (Balenoptera physalus) taken from an adult captured
at Belmullet, Ireland. The duodenal region (text-fig. 15) is not
sharply separated from Meckel’s tract. The latter is of even
calibre, and is thrown into a very large number of short, regu-
larly disposed minor loops suspended at the periphery of a much
elongated oval expanse of mesentery. These loops extend to the
extremity of the tract, and just where the recurrent limb of the
usual pendant loop begins its straight course towards the duo-
denal region, there lies a single small cecum. The hind-gut
consists of the almost straight recurrent limb, a short transverse
colon very close to the duodenum, but so far as I could make out,
suspended at this point only by the primitive mesentery, and of
a rather long nearly straight rectum.

The ceeum is short, but rather wide; its cavity is continuous
with that of the hind-gut, and separated by a simple semi-lunar
flap from the entrance of the ileum.

T have already described and figured the gut-pattern of one of
the Odontoceti (Mitchell, 1905, fig. 17). The Toothed Whales
have no cecum, and the whole length of the gut, from the
stomach to the anus, is suspended on a straight dorso-ventral
mesentery, all of it, except a very short rectum, being thrown
into closely-set minor loops. I ventured on the opinion, however,
that this almost reptilian simplicity was not primitive, and,
judging from the description given by Flower (1872, p. 428), I
suggested that the gut-pattern of Whalebone Whales would
approximate more closely to the common mammalian type. This

is actually the case. The characteristic mammalian pattern
14*

Pale DR. P. CHALMERS MITCHELL ON THE

appears in the Rorqual; the whole proximal limb of the pendant
loop is thrown into very numerous minor folds; the cecum is
placed more proximally on the loop, that is to say, nearer the
tip of the loop than in most mammals; with the elongation of
the mesentery suspending Meckel’s tract, the recurrent limb,
composed in this case almost entirely of hind-gut, is unusually

Text-figure 15.

Intestinal tract of embryonic Balenoptera physalus.

S. Cut end of gut nearest stomach. R. Cut end of gut nearest anus. C. Caecum.

long, and the rectum, although straight, is also long. In the
Toothed Whales, partly in relation to the diet of fish, Meckel’s
tract has become enormously long and its minor loops very
numerous, the caecum has disappeared, and the recurrent limb has
shortened until no trace of it remains. The complexity of the
stomach is so elaborate and so alike in Toothed Whales and

INTESTINAL TRACT OF MAMMALS, alts

Whalebone Whales, that were there no other reason for associating
these creatures, 1t would be impossible to place them far apart, and
it must be inferred that, so far as the gut-patterns afford indica-
tions, the Toothed Whales are more highly modified than the
~Whalebone Whales. If weare to seek for indications of the aftini-
ties of the Cetacea, it must be from the Mystacoceti, and not from
the Odontoceti, that we start. The difficulty is that a very simple
and primitive gut-pattern affords few indications. It is plain
that the Cetacean gut-pattern shows no trace of special resem-
blances with the patterns of the Ungulates or of the Sirenia.
There is some indication of similarity with the gut-patterns
of the aquatic Carnivores (see Mitehell, 1905, fig. 32, and
text-figs. 26 & 27, infra), but the more distal position of the
cecum (7. e., the eveater distance from the apex of the pendant
loop) and ane lengthening of the hind-gut in the Carnivores
present notable difference. Unfortunately, we do not know the
gut-patterns of extinct mammals, but, so far as may be judged
from Carnivores and Insectivores, it seems probable that the
Creodonts had an alimentary tract showing a simple pattern
much like those suggested in text-figs 1 A and 1B of this
memoir. The most notable peculiarity in the Cetacean pattern
is the position of the cecum towards the apex of the pendant
loop, a pecuharity that oecurs also in the Monotremes and some
of the Edentates. The lengthening of the gut and mesentery in
the longitudinal axis of the hone the ereat inerease in the
number of the minor loops on Meckel’s uate and the retention
of the importance of the primitive mesentery aie such adaptive
characters as might be expected in animals that had taken to an
aquatic life. The gut-pattern of the Cetacea, then, is compatible
with the view that Cetacea represent a very primitive stock,
long adapted to aquatic life.

Order ARTIODACTYLA.
Sub-Order Non-Ruminantia.

Family Hippopotamide. Hippopotamus amphibius (text-
HS, ALO)

The duodenum and Meckel’s tract are not sharply marked off
from one another. This part of the gut is extremely long (in
text-fig. 16 it has been somewhat simplified), and is thrown into
numerous minor folds compactly crowded on the periphery of an
oval expanse of mesentery. There is no cecum, but an increase
of calibre towards the apex of the pendant loop seems to mark
the point where, on the recurrent limb of that loop, the fore-gut
passes into the hind-gut. The distal portion of the recurrent
loop is thrown into a set of very large minor loops, attached to
the edge of the mesenterial expanse opposite to that suspending
Meckel’s tract, and therefore representing an ansa coli dextra.
The distal end of this colic loop, or series of minor colic loops,

214. DR. P. CHALMERS MITCHELL ON THE

approaches the duodenal region, where it is attached, by a strong
rather wide ligament, partly to the duodenum and partly to the
omentum. The gut then bends sharply round to form the
straight rectum of moderate length.

Text-figure 16.

Intestinal tract of Hippopotamus amphibius.

S. Cut end of gut nearest stomach. R. Cut end of gut nearest anus.
C.L.1. Colic Loop (ansa coli dextra).

Family Suide. Babirussa babirussa (text-fig. 17).

In the Swine, the duodenal region is better marked off, and
consists either of a single or a double loop. Meckel’s tract Ss
very like that of the Hippopotamus, being of even calibre, very
long, and disposed in closely packed minor loops. From the apex
the recurrent limb of the pendant loop runs dorsally a short
distance and then bends to enter the large cecum, which is a
forward continuation of the cavity of the hind-gut. Distad of
the cecum, the pendant loop, in‘the region occupied by a set of
large folds in the Hippopotamus, is developed into an enormous
double spiral, really composed of a very large single loop, the
proximal limb having a larger calibre than the distal limb. This
spiral, in some of the Swine, e. g., the common pig, and Babirussa
is a conical mass, in shape not unlike the shell of a whelk. In
Phacocherus and in Dicotyles the colic spiral was much flatter,
more like a coiled watch-spring. The hind-gut on leaving the

INTESTINAL TRACY OF MAMMALS. 215.

spiral runs up close to the duodenal region, and then bends over
to forin a relatively long but nearly straight rectum. ‘The spiral
loop, which from its position is an ansa coli dextra, is very capa-
cious and very heavy, and the portion of primitive mesentery
that supports it, and that carries the enormous blood-vessels.
supplying it, is reinforced by a strong band of fibres fastening it
partly to the duodenal region and partly to the omentum. A
similar, but much slighter secondary connection, ties the portion
of the hind-gut most contiguous to the dorsal middle line (distal
extremity of the pendant loop) to the duodenal mesentery.

Text-figure 17.

Intestinal tract ot Babirussa babirussa.

Lettering as in text-figs. 15 and 16.

Sub-orders Traguloidea, Tylopoda, and Pecora.

I have examined the intestinal tracts of several mammals
belonging to these three closely related sub-orders, since I
formerly gave an account of the patterns displayed in the
various families concerned (Mitchell, 1905), but as I have little
of general interest to add, I shall review the group as a whole.

The duodenal region is usually well separated, forming a long
distinct loop in nearly all, but rather less marked in the
Traguloidea and Tylopoda. Meckel’s tract is invariably enor-
mously long, of even calibre, and thrown into a very large number
of minor loops closely set round the periphery of the usual

216 DR. P. CHALMERS MITCHELL ON THE

mesenterial expanse. These minor loops cease towards the apex
of the original pendant loop, the first portion of the recurrent
limb being nearly straight, until it bends over to enter the
cecum. The cecum is always present and is capacious, but not
of great relative length. In Moschus, as an exception. it is very
long and narrow. Its cavity is a forward continuation of the
cavity of the hind-gut. I have already shown (Mitchell, 1905,
p- 518) that the relation of the ileum to the cecum and hind-gut
often presents appearances best explained on the supposition that
the normal cecum is the surviving member of an original pair of
ceca. I figured a mass of lymphoid tissue in the case of Gazella
marica, so situated that it seemed to represent a degenerate
second cecum. In an example of Moschus moschiferus that I
have examined since, the same portion of the gut was occupied
by a distinct cecal pouch, the aperture to which was marked off
by a V-shaped ridge. The iliac aperture lay on a vaised lip
between this and the wide aperture of the true cecum.

Text-figure 18.

Tleo-czecal region in Moschus moschiferus.

€. Cut ceeum. H. Cut hind-gut. S.J. Cut small intestine. W. “ Window” cut
in the wall of the ileo-cwcal region. J.A. Aperture of ileum to czxcum and
hind-gut. C.2. Secondcecum. ©.2 A. Aperture of second cecum to hind-gut
protected by V-shaped ridge.

I can suggest no explanation of this arrangement other than
that the second member of a prunitive pair of ceca is less
vestigial than is usual.

On leaving the cecum the hind-gut has a much smaller calibre
than is usual in mammals, being little wider than the distal end
of the ileum. The length and pecuhar arrangement of the
hind-gut form the most Coleraeterictic feature of the intestinal
pattern of this group of Artiodactyles. Immediately distad of
the cecum, there is usually a rather narrow single loop, which
I called the postcecal loop (Mitchell, 1905, fig. 22, P.C.L.).
Dr. Lonnberg (Lonnberg, 1907, p. 241) objects to this name,
inasmuch as he himself (lnnberg, 1903, p. 7) had termed a
similar loop in various ruminants the a@nsa proximalis. Dr.
Beddard in a later paper (1909, p. 181) calls this loop the ansa
paracecalis. The name is of little importance, but paracecal or
postezcal describes its position better Its presence is variable ;

INTESTINAL 'TRACT OF MAMMALS. PAWL

I did not find it in Traguloidea or Tylopoda, but it is present in
most of the true ruminants. Dr. Beddard describes it as
practically absent in M/adoqua and as spirally twisted in JMoschus.
In a Musk-deer that I examined, it was long, but showed no
trace of a spiral.

Distad of the postczcal loop, the recurrent limb of the pendant
loop is disposed in a spiral coil characteristic of the true
ruminants and equally weil marked in the Tylopoda. It is this
region of the gut that forms a set of large coils in the Hippo-
potamide, and a spiral arranged to form a solid conical mass in
the Suide. In the Traguloidea, as has been already described
by me (Mitchell, 1905) and confirmed by Beddard (Beddard, 1909),
the colic spiral is very small and is not flattened. In the
communication just cited, Dr. Beddard describes a somewhat
similar very small spiral in the minute antelopes of the genus
Madoqua, but in Tylopoda and all the true ruminants, except
Madoqua, of which the alimentary canal has been described, the
spiral is nearly flat and consists of a varying number of turns.
This flat spiral, in the undisturbed condition, is folded against
the mesentery that supports Meckel’s tract in the fashion that
the contiguous pages of « closed book touch one another. The
spiral is much smaller than the expanse of the tract, and, in the
undisturbed condition, it appears to be surrounded by the curved
line formed by the minor loops of the tract. ‘This arrangement,
which is familiar to anatomists, is well represented in some of
the figures given by Dr. Lénnberg and Dr. Beddard (e. g. Lonn-
berg, 1907, fig. 4; Beddard, 1909, text-fig. 14). The spiral coil is
composed of a long narrow outgrowth of the hind-gut, rolled up
from its apex, and the primitive mesentery belonging to the
spiral has coalesced with the mesentery supporting Meckel’s
tract in so complete a, fashion that “ short-cireuiting” blood-
vessels appear to supply these two very different regions of the
intestine indifferently. Moreover, especially where the coil is
large, secondary bands of fibres unite the coil firmly with the
intestinal region against which it is pressed. Unfortunately,
Dr. Lounberg does not appear to have considered these primary
and secondary attachments, and Dr. Beddards figures (e. g.
Beddard, 1909, text-fig. 15) do not distinguish between the five
different sheets of membrane to which the connections between
adjacent portions of the spiral coil may be referred, that is to
say, the double layer of the primitive mesentery of Meckel’s
tract to which the spiral coil is adherent, the double layer of the
primitive mesentery of the coiled loop that forms the spiral, and
the adventitious layer of connective-tissue fibres which assists in
holding the coil in its place. This absence of distinction would
be of no moment if the figures were, like my diagrams, intended
merely to represent the general morphology of the gut-pattern,
but it is another matter when the attempt 1s made to distinguish
between species and species by the characters of the spiral coil.
Dr. Loénnberg, who has made such an attempt, has devised an

218 DR. P. CHALMERS MITCHELL ON THE

ingenious method of figuring the spiral. He selects what he
takes to be the apex of the Toop, and up to this point tints the
entering limb of the intestine black, leaving the limb of exit
from the apex outwards round the spiral in grey (Lonnberg,
1907, fig. 4). Dr. Beddard has adopted Lénnberg’s method and
has carried it further. In a set of diagrams (Beddard, 0).
text-fig. 13) he represents the colic spirals of six animals and
arranges them in two series, each series indicating what he
descrihes as a distinct type of spiral. In each case he has
selected what he takes to be the apex of the loop, and, like
Lonnberg. shades the entering limb black, the limb of exit grey.
In one series, containing Madoqua phillipsi, Cephalophus dorsahs,
and Moschus moschiferus, the entering limb of the intestine is on
the smaller curve of the spiral as it approaches the apex, and if
the apex happens to point towards the end of the long axis of
the spiral, away from the point of entrance, then the enters
limb finishes on the cecal side of the apex. In the second series,
containing Tragulus stanleyanus, Cephalophus maxwelli, and
Antilocapra americana, the entering limb of the intestine Hes on
the larger curve of the spiral as it approaches the apex, and if
the apex happens to lie towards the end of the long axis of the
splval away from the point of entrance, or be imagined to have
grown round to that point, then the entering limb finishes on
the opposite side of the apex from what happens in the first
type. This distinction between the types of spiral is stated by
Dr. Beddard to be so important that the presence of one type in’
one species of Cephalophus, and of the other in another species
of that genus, 1s a generic distinction, confirming certain
undesignated differences in external characters which “ appear
to him to be quite as great as those which distinguish certain
other genera of Antelope.”

It is plain, however, that the reality of the distinction on
which Dr. Beddard relies, depends on the actual point selected
as the apex of the spiral. In text-fig. 19 I have reproduced
the drawings which Dr. Beddard gives as the first examples of
each type (Beddard, 1909, text-fig. 13) 1, 2), with the alteration
that they are reverse as in a mirror, to make easier comparison
with Dr. Lonnberg’s figure (Linnberg, 1907, fig. 4) and my own
diagrams, and with an ‘addition to which I shall refer presently.

Obviously, if Dr. Beddard had continued the black shading
representing the ingoing limb of the intestine from the point
marked X, where he left it in the figure of Jragulus, to the
point I have marked XX in the same figure, the two ‘‘ types ”
of spiral would have been in every way identical. Anyone who
has attempted to follow the closely adpressed limbs of a compli-
eated ruminant spiral on the actual specimen, will appreciate
that the fixing of the actual apex is a difficult judgment and not
a substantial basis for the discrimination of types or the deter-
mination of genera.» The judgment is the more difficult, because,
as I have already explained, any two contiguous portions of the

INTESTINAL TRACT OF MAMMALS. 219

spiral may be united either by their own primitive mesentery,
by the primitive mesentery of Meckel’s tract, against which they
are fixed, or by adventitious fibres. ‘There is, however, a definite
morphological criterion. The primitive mesentery of the loop
which is coiled into a spiral, whether it be retained in whole or
in part, fused with or replaced by the mesentery of Meckel’s
loop or adventitious fibres, must have been attached along the
primitive dorsal line of the gut, that is to say, the side of the
hind-gut opposite to that on which the cecum lies, the side into
which the ileum opens. In text-fig. 19 I have dotted in the
primitive mesentery, and it will be seen at once that in the figure
of MONO geE Dr. Beddard has adjudged the apex correctly COE

Text-figure 19.

is ZA.

Diagrams of Beddard’s types of colic spirals.

1. Madoqua phillipsi. 2. Tragulus stanleyanus.

Modified from Beddard (1909, text-fig. 13, 1, 2). The distal end of the ileum, the
cecum, and the entermg limb of the intestine in black; the outgoing limb IS
unshaded. X. Beddard’s apex, the true apex inl. XX. The true apex in 2.
The dotted surface is the primitive mesentery of the loop.

but that in the figure of Tragulus he has adjudged it incorrectly.
If in that figure the point marked X were the apex, then the
mesentery would be attached to the wrong side of the gut. If,
on the other hand, the mesentery be considered, the point that I
have marked XX is seen to be the true apex, and the blackening
of the ingoing limb should have been continued from X to XX,
so abolishing the distinction between the two types. Precisely
in the same way, in Dr. Beddard’s figures of Cephalophus maxwelli
and Antilocapra americana (Beddard, 1909, text-fig. 13, 2a, 26)
and in Dr. Loénnberg’s figure of the Elk (Lénnberg, 1907, fig. 4),
from which Dr. Beddard’s method was taken, the point that has

220 DR. P. CHALMERS MITCHELL ON 'THE

been selected as the apex would place the mesentery on the
wrong side of the gut. The supposed distinction in type does
not exist.

I do not doubt but that an intensive study of these raminant
coils may lead to very interesting results. It is important to
realize, however, that a naive comparison and description of such
complex structures may be extremely misleading.

Distad of the colic spiral the recurrent limb of the pendant
loop undergoes a further complication before it reaches the
dorsal middle tine. ‘Che outgoing limb of the spiral, still with
its mesentery adherent to the mesentery of Meckel’s tract,
pursues a circular course, following the line of the secondary
coils of Meckel’s tract and lying fon this and the spiral
itself until it reaches the duodenal region, where its suspension
is usually reinforced by a colico- duodenal ligament, and then
bends round to form the rectum, which passes backwards towards
the anus suspended in the usual. fashion by its own primitive
mesentery. The adherence of this special coil to the mesentery
of Meckel’s tract is so close, that I have never been able to
dissect it off with any portion of its own meson and I
suspect that this mesentery has disappeared. In the diagrams
that I have Bivens of ‘Traguloidea, 'Tylopoda, and Pecora (Mitchell,
1905, figs. 19-22), ADs portion of the gut is marked 8.F.,
supra-meckelian fold, and is displayed as dissected off and free
from mesentery. ‘This region appears to be simplest in the
Traguloidea and the Tylopoda, but in an example of the White-
tailed Gnu (Connochetes gnw) | was surprised by finding it reduced
to a single quite narrow loop. In the Giratte it is very com-
plicated, “forming, instead of a wavy line round Meckel’s tract, a
set of irregular loops in the space between the tract and the
spiral coil, rather like a similar series that Dr. Lénnberg has
figured in the case of a foetal Elk (Lénnberg, 1907, tig. 4). In
some of the deer, sheep, and goats that I have examined, the
general course of this loop is a sweeping curve concentric with
the curve of the minor loops of Meckel’s tract, but at the distal
eud, just before bending over to form the rectum, it gives rise
to a quite definite, scraight, and rather narrow loop, stretching
across towards the ‘spiral coil and sometimes even crossing a
portion of the coil.

Lam reluctant to suggest homologies between the minor loops
found on the very peculiar hind- -gut of this group of Artio-
dactyles and the minor loops found in the hind-gut of other
groups, as it seems to be plain that we should bave first to trace
such loops down to their form in the ancestral Artiodactyle,
Rodent, and Primate, and so forth, before instituting any valid
comparison between their appearances in the higher members of
these different groups. In the very general sense, however, that
the distal or dorsal extremity of the recurrent loop corresponds
with the transverse colon, and a specialized outgrowth to the
right of this may be named an ansa dextra, a specialized out-

INTESTINAL TRACT OF MAMMALS. 221

growth to the left an ansa sintstra, then both the spinal loop and
the supra-meckelian fold of Pecora, Tylopoda, and Traguloidea
may be taken to represent ansce dextre.

Farther work, and the consideration of the points raised by
writers who have tollowed me, have not given me any reason to
modify the general summary I gave in 1905 (Mitchell, 1905,
p. 476):—“The Ruminant Artiodactyles display a pattern
peculiar to the group, and characterised by the enormous length,
special modification, and arrangement of the hind-gut. In all,
the hind-gut displays three well-marked regions: a spiral loop
simpler in 7ragulus, in” (most of) ‘the others forming a-closely-
coiled, flat, watch-spring like arrangement, folded over on the
mesentery that supports Meckel’s tract; a supra-meckelian fold
which, in the characteristic and most specialised cases is stretched
round Meckel’s tract just at the line where the minor folds leave
the mesentery, and, which is drained by branches from the vessels
of Meckel’s tract ; anda rectal portion, the degree of convolution
of which varies nearly divectly with the size of the animal.

“The non-ruminant Artiodactyles display a pattern funda-
mentally similar to, but less complicated than, that of ruminant
forms. Meckel’s tract is almost identical in its disposition.
The spiral coil of the hind-gut” (usually, not in the Hippo-
potamus) “is present and. is very large, but its calibre is wider
in proportion to its length, and the coiling is not so flat. There
is no more than a trace of the supra-meckelian fold, so that the
hind-gut, although long, is less differentiated.”

Order PerissopactyLa. (Text-figure 20.)

I have little to add to the account [ have already given
(Mitchell, 1905, p. 476, figs. 25, 24, 25) of the gut-pattern of
the Rhinoceros, Tapirs, and Equide. For convenience, I repeat
as text-fig. 20 the figure I have already given (Mitchell, 1905,
fig. 25) of the gut-pattern of Hquus granti. I have added at
M.the portion of primitive mesentery between the cecum and
the ileum, and at XX have marked the line along which the
cecum 1s tied by adventitious fibres to the colic loop. I have
verified these points on the domestic horse, as no exampie of a
zebra was available. In the three families the pattern is quite
definite and remarkably uniform. The duodenum is a distinct
loop, Meckel’s tract is relatively short and compact, the cecum
is very capacious, but relatively smaller in the Rhinoceros than
in the others, and situated rather high up on the recurrent limb
of the pendant loop. It is greatly exceeded in capacity by an
enormous narrow loop, each limb of which is large in calibre,
formed as an outgrowth of the distal portion of the recurrent limb.
The hind-gut then bends round to form the relatively simple
rectum. The cecum is connected with the distal extremity of
the ileum by a short double fold of mesentery, the usual remnant
of the primitive mesentery which is found in this situation in

2, DR. P. CHALMERS MITCHELL ON THE

most mammals. It extends from the base of the cecum only
along a very small proportion of the length of that organ. In
addition, a layer of stout fibres, quite distinct from true mesentery
in appearance and relations, binds together the two limbs of the

Text-figure 20.

Intestinal tract of Equidee.

Shehtly altered from Trans. Zool. Soc. 1905, fig. 25. S. Cut end of intestine
nearest stomach. R. Cut end of intestine nearest anus. CC. Cwcum.
C.L. 1. Colic loop (ansa coli dextra). M. Remnant of primitive mesentery
between ileum and base of cecum. XX. Line of attachment of the adventi-
tious fibres (removed in the figure) which tie the cecum to the colic loop.

enormous colic loop, and passes over from them to the cecum.
‘This secondary attachment is least strong in the Rhinoceros, but
in the Tapirvide and Equide ties down the greater part of the

INTESLINAL TRACT OF MAMMALS. 223

length of the cecum to the colic loop. There are also strong
cavo-duodenal and colico-duodenal ligaments.

The remarkable similarity of the gut-patterns of the three
families of Perissodactyles contrasts strongly with the fact that
there is no resemblance between the Perissodactyle and the non-
ruminant and ruminant Artiodactyle patterns. ‘The Swine are
omnivorous with a tendency towards vegetable diet ; the Hippo-
potamus and all other Artiodactyles are, like the Perissodactyles,
vegetarian in diet. In all the hind-gut is capacious in relation
to the diet, but the pattern, none the less, follows affinity rather
than function.

Order RoventiA. Dipus egyptius (text-fig. 21).

I have little to add to the account I gave in 1905 (Mitchell,
1905, figs. 26-30) of the intestinal gut-patterns displayed by
Rodents. The gut tends to be relatively long, no doubt in
association with the chiefly vegetarian diet. The duodenal loop
is usually very well marked off from Meckel’s tract, the latter
always being supported on an oval expanse of mesentery, and
varying only to the extent to which it displays minor loops.
The cecum is usually capacious, long, and sacculated. Remnants
of an originally paired condition are frequent. The cecum,
especially when long, tends to be coiled in a spiral, and this
coiling may involve not only the distal portion of Meckel’s tract,
but the proximal portion of the hind-gut.

Even when the gut is relatively short, traces of the spiral
condition are frequent, suggesting that in some Rodents,
especially small omnivorous types, the gut has been shortened
secondarily from the longer condition normal in the group.

The cecum is placed rather high up on the recurrent limb of
the pendant loop. The remaining portion of the latter varies in
a remarkable degree, both in species and in individuals. The
most common condition is the presence of two rather narrow
colic loops, but these may be reduced to a single loop or there
may be three (text-fig. 21, C.L. 1, 2,3). The most proximal
loop (C.L. 1) is the portion that tends to be involved in the
spiral twisting of the cecum, and is what has been termed a
paraceecal or post- cecal loop. The two more distal loops (C.L.
2,3) may be spirally twisted, either separately or together, but
in the more common case they are untwisted. I cannot regard
this occasional spiral arrangement as indicating any homology
between these loops and the spiral of Artiodactyles, or as
suggesting any special aftinity between Rodents and Artio-
dactyles. The colic spiral of the Artiodactyles, especially of the
Ruminants, 1s an extremely definite formation, invariably present
in the adult and appearing at a very early stage in embry onic
life. In Rodents it varies from individual to individual, may
involve one or two loops, and is often inconspicuous or absent in
small or relatively young individuals. The spiral formation that

224 DR. P. CHALMERS MITCHELL ON THE

occurs frequently in Rodents and is extremely rare in other
groups, so that it may be designated as a Rodent peculiarity,
affects the cecum.

The colic loops may be pressed against the mesentery of
Meckel’s tract in such a fashion that their own primitive mesentery
may be partly obliterated and replaced, either by the mesentery
of the tract, or by adventitious fibres. In Dipus (although in
this respect text-fig. 21 is somewhat simplified and diagrammatic)

Text-figure 21.

Intestinal tract of Dipus egyptius.

S. Gut end of the gut nearest to the stomach. R. Cut end of the gut nearest to
the rectum. C. Ceeum. C.L. 1, 2,3. Colic loops. C.L. 1. Paraczcal loop.
C.L. 2 & 3. Anse coli dextre.

and in other forms with a relatively simple gut, it is still possible
to dissect away the colic loops and to unfold them to show their
primitive pattern with a minimum of cutting. In other forms,
especially those in which the loops are long, the adherence
between them and the tract is so elaborate as to recall the
condition in Artiodactyles, and the gut cannot be laid out to
show its primitive pattern without extensive destruction of
mesentery, blood-vessels, and secondary attachments. I am

INTESTINAL TRACT OF MAMMALS. 2.25;

convinced, however, that such vague resemblances between
Rodents and Artiodactyles are convergent.

At its most dorsal extremity the recurrent limb of the pendant
loop sweeps round to be continued backwards as the rectum.
In Dipus and in many other Rodents the rectal portion is
relatively simple. It may be much lengthened, especially in its
proximal portion, and this lengthening may take the form of a
single rather narrow loop, an ansa coli sinistra, as for instance,
in Hystrix, or, aS 18 more common, a much-contorted loop or
number of loops, as in Lagostomys and Dolichotis. I do not
attach much importance to this distinction, as I have found both
forms in different examples, both apparently adult, of Atherura,
and in very young and adult examples of some other species.

Variability appears to be a marked character of the subsidiary
loops of the hind-gut in Rodents. Three writers have given
a good deal of attention to the matter. Tullberg, with whose
work, unfortunately, I was unacquainted when I wrote in 1905,
published a most valuable monograph on the group (Tullberg
1899), in which a long section and many plates are devoted to
descriptions of the gut of a very large number of Rodents.
Tullberg devoted himself chiefly to the gut and its attachments
as seen in the unfolded condition when the abdominal cavity is
opened, but there are few features that cannot be understood
from his careful figures. My work followed in 1905, and later,
Dr. Beddard (Beddard, 1908), following the method of Tullberg,
rather than mine, called attention to a good many differences
that he had noted in examination of some of the species that
Tullberg had described, and added descriptions of the conditions.
he found in other species not described by Tullberg. I have
tabulated the results of the three writers. It would be a waste
of space to give the details; it is enough to say that the colic
loops of Rodents appear to differ individually and at different.
stages of growth, in number, attachments, degree of spiral
coiling, relative length, and distinctness (7. e., definite narrowness,
or width and minor expansions). I hesitate, therefore, to
follow Tullberg, even in his cautious use of these structures in
the classification of Rodents themselves, and I think it an
unwise adventure to pursue the comparison of the individual
loops from Rodents to other groups. With the reservation that
these colic loops are rather inconstant, it is possible to distinguish
them up toa point. Immediately distad of the cecum lies what
Tullberg calls the paracecal loop, corresponding with what I
have termed the postczcal loop. This may be absent, imperfectly
formed, definite, nearly straight, twisted with the cecum, or
showing an independent spiral. Next come the two loops of the
recurrent limb that are most commonly present in Rodents.
These are termed by Tullberg ansce dewtre 1 and 2. Frequently
only one is present, especially in young examples of a few days
old. Dr. Beddard, unfortunately, has confused the matter by
labelling the upper or more distal of these the ansa sinistra

Proc. Zoor. Soc.—1916, No. XV. 15

226 DR. P. CHALMERS MITCHELL ON THE

(Beddard, 1908, text-fig. 116); the term sinistra belongs to a
more distal region of the gut, and Tullberg’s definitions, descrip-
tions, and figures make this point quite clear. These ans
dextre may be definite and narrow, spirally twisted separately or
together. The three loops C.L.1, 2, 3 in the diagram of Dipus
(text-fig. 21) represent a paraceecal loop and two anse dextre.
Distad of the highest point of the recurrent limb, and to the
left of the equivalent of the transverse colon, there may be
another region of expansion. When this subsidiary loop is
simple and narrow, Tullberg recognises it as distinct and calls it
the ansa sinistra ; when it is thrown into irregular minor folds,
he leaves it undesignated. As I have already pointed out, I
have found both conditions of this expansion in different examples
of the same species, and therefore do not attach much importance
to it. But, definite or irregular, if it be named at all, ansa
sinistra 1s the correct name. It is absent in Dipus.

The gut-pattern of Rodents, then, displays usually a separate
duodenum, a well-defined Meckel’s tract, a cecum frequently
spirally twisted, and an elongated hind-gut, variable in the
number and nature of the subsidiary loops which may be
developed.

Order Insecrivorna. J/acroscelides species? (text-fig. 22).
Talpa ewropea (text-fig. 23).

Taking the examples of animals grouped together as Insectivora
that I had been able to examine when I wrote before (Mitchell,
1905) and those that I have seen since, I cannot make up a
series approaching completeness. Putting together my own
observations with what I am able to gather from other writers,
I think that three types of different degrees of simplicity
can be distinguished among the gut-patterns of Insectivora.
In Macroscelides (text-fig. 22) the duodenal region cannot be
recognised as separate from Meckel’s tract. The latter is
thrown into rather simple short loops round the whole of the
descending limb anda small portion of the recurrent limb of
the pendant loop; then follows a long, nearly straight portion
running up towards the dorsal line. The cecum is enormous,
and is ‘attached high up on the recurrent limb of the pendant
loop. The distal portion of the pendant loop is expanded to
form a very large nearly closed colic loop, thrown into a number
of minor loops. In the undisturbed condition this lies folded
against the mesentery of Meckel’s tract, but I found no secondary
connection. The recurrent loop then bends round to form the
straight rectum. The superficial resemblance between this
pattern and that presented by some of the smaller Diprotodont
Marsupials, such as Phalangista vulpina (Mitchell, 1905, fig. 5)
is extremely close. When I had finished the drawing I thought
that it had a familiar look, and on hunting through my notes,
I found that, from the point of view of this memoir, it would

INTESTINAL TRACT OF MAMMALS, 227

have been almost unnecessary to draw a second figure but for
the fact that the colic loop is an ansa coli devtra in Macro-
scelides, and probably is not so in Phalangista.

In Talpa europea (text-fig. 23) the pattern is rather simpler.
The duodenal region is rather more distinct, although in the
diagram this is over-emphasised ; Meckel’s tract consists of a

number of rather long minor. loops occupying the whole of the
proximal limb of the pendant loop. There is no trace of a
cecum, and nothing eise to indicate where the ileum passes into
the hind-gut. The recurrent loop runs straight up towards the
dorsal middle line, and, just before bending round to form the
straight rectum, gives rise to a single very narrow and long
colic loop (an ansa coli devtra) which, in the unfolded condition,
is bent over towards Meckel’s tract and shows a trace of spiral
twisting.

Text-figure 22.

‘Intestinal tract of Wacroscelides species.

S. Cut end of the gut nearest stomach. R. Cut end of gut uearest rectum.
C. Cecum, C.L.1. Colic loop (ansa coli dextra).

In Erinaceus and in Centetes (Mitchell, 1905, fig. 31), the
duodenum and Meckel’s tract are not distinct. The latter is
arranged in very regular minor loops round the periphery of a
nearly circular expanse of mesentery. There is no trace of a
cecum, or of a colic loop, but the recurrent limb runs up towards
the dorsal middle line, and then bends over to form a short
straight rectum.

In the three types the whole gut is suspended on a continuous
mesentery, and the three appear to show stages in the attain-
ment of a secondary simplification, the stages of which are, first,

ay

228 DR. P. CHALMERS MITCHELL ON THE

the loss of the cecum, next the obliteration of distinction
between the fore-gut and the hind-gut, and the disappearance
of the colic loop. I found no adventitious connections in any of
the types, but Tam unable to attach much importance to the
presence or absence of these. Nor can I lay stress on the folding
of the gut on itself ; this is certainly present in Macroscelides
and Talpa; Beddard states that it occurs in Hrinaceus, and
found it present in one example of Centetes, absent in
another.

Text-figure 23.

—

Ly.

Q
S =
Ss Ss

R\

Sie
rae

Intestinal tract of Talpa europea.

Lettering as in text-fig. 22.

The similarity of type between the pattern of Macroscelides
(and probably of Tupaia, according to Flower and Hunter) and
the pattern of Phalangista, recalls Dr. Broom’s association of
these animals with Diprotodont Marsupials and removal of them
from the Insectivora (Broom, 1902, 1915). I must point out,
however, that the pattern shared by the two is a very simple
derivative of the primitive mammalian type, and on this ground
alone, I would not be disposed to argue close afiinity amongst
the animals that display it. Moreover, if, as seems to me most
probable, the colic loop of Phalangista resembles that of Bettongia
and the Kangaroos, and belongs to the region of the hind-gut
distad of the pendant loop, then the resemblance of Macroscelides

INTESTINAL TRACT OF MAMMALS. 229

to Marsupials is not so close as to Monotremes. This is a point
to which I had paid no special attention in 1905. Dr. Broom’s
conclusions are derived from investigation of the organ of
Jacobson, and certainly the conditions that he has found appear
to form a broader basis for systematic conclusions. I am not
quite certain, however, as to whether or no he means to imply
that the “‘ Ceenrhinate”’ type of organ, which he finds to occur
in Talpa, Sorex, Hrinaceus, Gymnura, Centetes, and other
normal Insectivora, as in Chrminona and Ungulata and most
higher mammals, is a derivative of the more aeeserall * Archieo-
saan ” type which he has found in Vupaia, Macroscelides,
Diprotodonts, ete. If he means that the Archzorhinate type is
the primitive type, and therefore to have been present in the
common stock, the fact that it has been retained by certain
forms is no conclusive argument for placing these forms together.
As he finds it to occur in Monotremes, on the one hand, and
in Dasypus, Orycteropus, and Rodents on the other, I suspect
that it is, ike the presence of a primitive gut-pattern, a character
that must be used with caution in classification.

Without carrying further this question of breaking up the
Insectivora, I may sum up by saying that the gut- patterns of
the group start from an extremely simple type, and show
successive stages of secondary simplification.

Order Cutroprera. Lhinopoma microphyllum (text-fig. 24).
Artibeus planirostris (text-fig. 25).

Since I wrote in 1905 I have had the opportunity of examining
the intestinal tract in some other Bats, of which the most
interesting was an example of Rhinopoma mucrophyllum (=
ft. hardawickit). ‘The latter and Jlegaderma spasma were the
two Bats in which Owen found a cecum present (Owen, 1868,
p. 429). In Rhinopoma the duodenal region is well separated
from Meckel’s tract. Meckel’s tract makes up the greater
portion of the gut, and consists of a number of very irregular
minor loops, arranged so that they nearly complete the circum-
ference of a circular expanse of mesentery, suspended by a narrow
stalk to the mesentery of the duodenum in front and to that of
the hind-gut posteriorly. In other words, the whole of the
recurrent limb of the pendant loop is occupied by Meckel’s tract,
and it is only where it bends backwards to form the short and
nearly straight rectal portion that the attachment of the caecum
marks the transition from fore-gut to hind-gut. The cecum is
short and conical. The hind-gut may be Nenanded as without
a colon, but consisting merely of a rectal portion.

In the unexpanded condition, the subsidiary coils of Meckel’s
tract are irregularly folded over the mesentery so that they
make up a large irregular mass visible as soon as the abdominal
cavity is opened. The duodenum is also folded backwards, and
cannot be seen until the mass of the fore-gut has been pushed

230 DR. P. CHALMERS MITCHELL ON THE

aside, whereupon it is visible, stretching backwards in close
association with the rectum, but without secondary attachment
either to that or to the mesentery of Meckel’s tract. The
primitive mesentery is retained along the whole length of the
gut, quite unobscured by secondary attachments.

The position of the cecum beyond the extremity of the
pendant loop, and thus approaching the condition in most birds,
especially the higher types of birds, is curious and very unlike
the common condition in mammals,

Text-figure 24.

Intestinal tract of Rhinopoma microphylliun.

Lettering as in text-fig. 22.

In the other Chiroptera that I have examined there was no
eecum, but the general form of the pattern when unfolded and
the mode of arrangement in the undisturbed body-cavity were
closely similar. The hind-gut was straight and relatively longer,
its proximal end approaching very close to the duodenum, In
most species the minor loops of Meckel’s tract were irregularly
folded and lobed as in #hinopoma, but in Artibeus (text-fig. 25)
they were relatively long (longer in proportion than in the
diagram), and very straight. In the folded condition, the long
straight loops, closely packed together and bent over from the
edge of the mesenterial expanse to which they were attached,
suggested a spiral conformation at first sight.

In an example of Pteropus mediuns that I have recently

INTESTINAL TRACT OF MAMMALS, DSi

examined, the duodenum consisted of two short loops marked off
by their larger calile. The proximal limb of the pendant loop
and the beginning of the recurrent loop were thrown into long
and very irregular minor loops, distad of which the recurrent
limb had a straight course up to the middle dorsal line, where
it bent round sharply to pass into the straight rectum.

In 1905 I had not seen an example of a Bat witha cecum, and
was content merely to point out the general similarity between
the simple gut of Bats and of such Insectivores as Centetes, with
the caution, however, that in neither case could it be asserted
safely that the simplicity was primitive. Iam now able to add

Text-figure 25,

Intestinal tract of Artibeus planirostris.

S. Cut end of intestine nearest stomach. R. Cut end of intestine nearest stomach.

to the comparison. In Insectivores and Chiroptera the gut is
relatively short, disposed on a continuous primitive mesentery,
and in the extremer types shows little differentiation. In the
Insectivores, however, the simplicity has been reached from a
condition in which the eczcum was developed a considerable
distance from the distal end of the pendant loop, and the upper
portion of the recurrent limb possessed a large colic loop. In
the Chiroptera, the whole of the pendant loop gives rise to
Meckel’s tract, the cecum being placed distad of the passage of
the pendant loop into the straight rectum. So far, therefore, as
evidence is afforded by the gut-pattern, there is no reason to
associate Chiroptera with Insectivores,

232 DR. P. CHALMERS MITCHELL ON THE

Order CARNIVORA.
Sub-Order Pinnipedia. Odobenus rosmarus (text-
fig. 26). Macrorhinus leoninus (text-fig. 27).
The pattern of the intestinal tract of the Seals and their
immediate allies is distinguished by simplicity, great length of
Meckel’s tract, reduction of the cecum, relative shortness of the

Text-figure 26.

QS,
Nes
SSS)

Intestinal tract of Odobenus rosmarus.

I. The tract as a whole. S. Cut end of gut nearest stomach. R. Cut end of
gut nearest anus. C. Ciecum.

If. Enlarged view of junction of ileum and hind-gut with cxcum, — Ile. Cut end
of ileum. Col. Cut end of colon. C. Cxecum. Part of the side-wall has been
removed to show the protrusion of the ileum into the cxco-colic cavity.

bo
“15

INTESTINAL TRACT OF MAMMALS,

hind-gut, and simple suspension from a continuous mesentery.
B : : ;

In the Walrus (text-fig. 26) there is no clear separation of the

duodenal region from Meckel’s tract. The latter has an almost

Text-fgure 27.

Intestinal tract of Macrorhinus leoninus.

From a photograph by Mr. D. Seth-Smith. A pocket-knife has been inserted into
the cavity of the duodenum where it was separated from the stomach ; the cut
distal end of the rectum has been bent up towards the duodenum to bring it
into focus.

even calibre throughout its length, and is very long (its length,
although relatively less than that of the Elephant- Seal, has been
vather under-estimated in the diagram). It is thrown into a

234 Dk. P. CHALMERS MITCHELL ON THE

large series of irregular minor loops, nearly completing the
periphery of an oval expanse of mesentery. The recurrent
limb of the pendant loop also enters into the formation of
Meckel’s tract, and the short, rounded cecum lies just where the
pendant loop bends round ab its highest point to pass into the
hind-gut.

The distal end of the ileum projects through into the cavity of
the hind-gut at an angle, the projection being much greater on
the cecal side. The cavity of the cecum is, as is usual, a forward
continuation of the cavity of the hind-gut. There is no trans-
verse colon, the gut, at the highest point of the distal end
of the recurrent loop, bending round sharply to the rectum.
There is, in fact, practically no true colon, but the rectum is
considerably longer than the course that it has to traverse, and
is thrown into a number of minor loops.

I have recently had the opportunity of examining the intestines
of a young Elephant-Seal (Macrorhinus leoninus). By the kind-
ness of my colleague, Mr. D. Seth-Smith, I am able to reproduce
as text-fig. 27 a photograph of the intestinal tract removed from
the body and laid out to display its pattern. For this purpose,
owing to the simplicity of the gut in this group and the absence
of secondar ‘y connections, it was necessary only to sever the
primitive mesentery that stretches from the duodenum to the
rectum.

As in the Walrus, the duodenal region is not sharply marked
off from Meckel’s tract. The latter is of even calibre, and is
thrown into extremely numerous minor folds arranged so as
almost completely to surround an oval expanse of mesentery.
Its length is enormous. The Elephant-Seal that I examined
measured six feet nine inches from the tip of the snout to the
tip of the tail: the small intestine measured with the tape, when
the mesentery had been detached, one hundred and seventy-five
feet six inches ; the hind-gut was only two feet four inches long.
When it has nearly reached the level of the duodenum, the
distal limb, without any change of calibre, bends sharply round,
and after a course of nearly a foot, suddenly changes in calibre.
At this point there is nothing that can be called a cecum, and it
is doubtful whether the change from fore-gut to hind-gut can be
placedaccurately. If it is where the gut expands. then the position
is quite abnormal amongst mammals: if, as in the Walrus, it is
at the extremity of the recurrent limb of the pendant loop, then
the sudden change of calibre in the course of the hind-gut, is also
unusual, The hind-gut, after widening, bears an enlarged simple
loop, which in the photograph is represented in an “unnatural
position ; the distal end of the rectum was bent forwards to bring
it into the picture. It appears to belong to the region distad of
the pendant loop, and is an ansa coli sinistr a.

The gut patterns of the Walrus and of the Elephant-Seal do not
differ notably from those of the Sea-Lion and of the true Seals
that I have already described (Mitchell, 1905, p. 493, fig. 32).

INTESTINAL TRACT OF MAMMALS. 235,

The great increase in size of Meckel’s tract, the position of the
cecum, if present, as is usually the case, close to the distal
extremity of the recurrent limb of the pendant loop, and the
presence of a distinct expansion of the hind-gut, although that
is relatively short, are the most salient features. The pattern 1s
quite different from that of the Manatee. It resembles the
pattern of the Cetacean gut only in the extreme length of the
small intestine; it differs notably in the position of the cecum
and in the hind-gut. As I shall show presently, it resembles
closely the pattern of terrestrial Carnivora, the difference being
chiefly the lengthening of Meckel’s tract, which is best explained
as an adaptation to diet,

Sub-Order Fissipedia. Proteles cristatus (text-fig. 28).

In 1905 I deseribed and figured the patterns of the gut of a
number of Fissipede Carnivora (Mitchell, 1905, p. 495, figs. 33

to 38) and have very little to add. I have had the opportunity,

Text-figure 28.

Intestinal tract of Proteles cristatus.

§. Cut end of gut neareststomach. R. Cut endof gut nearest rectum. C. Caecum.

however, of examining an example of the Aard Wolf (Proteles
cristatus), an animal that is now seldom seen in menageries. So
great is the uniformity of pattern amongst the terrestrial carni-
vores that Proteles may serve asan example of all. The duodenal

236 DR. P. CHALMERS MITCHELL ON THE

region is rather distinct from Meckel’s tract, The latter forms
the longest part of the gut, but is relatively shorter than in most
types of mammals. It is thrown into rather simp‘e minor loops
which reach to the extremity of the proximal limb of the pendant
loop, but in the recurrent limbare replaced by an almost straight
portion running up towards the duodenum. The cecum is
situated high up on the recurrent limb, and in Proteles is short,
although in the example I examined it was not so globular as
was described by Flower (Flower, 1869), and had a slight spiral
twist not noted by that author. The axis of the twist was con-
nected with the distal extremity of the ileum by a very small
fold of mesentery. The cecum is frequently absent in Carnivores,
but when present, and especially when it is relatively not very
small, frequently displays a spiral twisting.

Distad of the cecum the hind-gut bends round sharply,
increasing in calibre and displaying a rather considerable
expansion before it passes into the short straight rectum. The
hind- gut is relatively short in all the Carnivores, and a notable
feature is the reduction or absence of the transverse colon, the
recurrent limb of the pendant loop bending round to pass back-
wards either with a very sharp turn, or at most a narrow sweep.
From re-examination of all my original laboratory notes and
drawings, I am disposed to think that at least in the great
majority of Carnivores the expansion of the hind-gut, when
present, belongs to the portion of the gut distad of the pendant
loop, and is therefore an ansa coli sinistra not homologous with
colic loops developed on the recurrent limb of the pendant
loop.

In the Bears, however, as I have already shown (Mitchell,
1905, fig. 34), there is a definite colic loop present, and as this is
an expansion of the recurrent limb of the pendant loop, it must
be regarded as an ansa coli devtra. In the new-born cub of a
Brown Bear, this loop was more definiteand more elaborate than
in the example of Ursus malayanus that I formerly figured.
The Otter shows a somewhat similar condition. There was no
trace of any secondary connection linking the colic region to the
duodenum.

The primitive mesentery is retained in a nearly complete
condition in most Carnivores, and in association with the
relative shortness of the gut in the terrestrial forms, secondary
attachments between proximal and distal portions of the gut
appear to vary even individually, and never attain the physio-
logical importance that they may be presumed to have in many
other groups. As I have already stated, I do not believe them
to have systematic importance.

It is clear that the Pinnipedes and Fissipedes exhibit gut-
patterns that are fundamentally similar, although the resem-
blance is decreased by the adaptive lengthening of the gut in
the aquatic forms.

INTESTINAL TRACT OF MAMMALS. Baye

“Order Prosimim. Chiromys madagascariensis (text-fig. 29).
Lemur species ? (text-fig. 30).

I have had the opportunity of examining an example of the
Aye-Aye, and find that the pattern of the intestinal tract of that
animal resembles extremely closely the patterns exhibited by the
genus Lemur, much more closely than the patterns of other
Lemurs. In this matter [ completely confirm the conclusion of
Dr. Beddard (Beddard, 1908, Chiromys). A comparison of the
diagram of Chiromys (text-fig. 29) with that of a species of the
genus Lemur* (text-fig. 30) makes this resemblance plain. In

Text-figure 29.

Intestinal tract of Chiromys madagascariensis.

S. Cut end of gut nearest stomach. R. Cut end of gut nearest rectum. C. Cxcum.
C.L.1., C.L.2. Colic loops (anseé coli dextra et sinistra).

both the duodenal region is represented’ by two minor loops,
which in the Lemur were marked by a much greater calibre.
Meckel’s tract was relatively rather short, and thrown into:
simple minor loops occupying the proximal portion of the
original pendant loop, but not reaching far beyond it. The
fore-gut enters at a right angle to the cavity of the very
large cecum which is in wide continuity with that of the
dilated proximal portion of the hind-gut. In Lemur the

* The Lemur was an example of what has been called in the Society’s Gardens.
LI. brunneus, the Black-headed Lemur, but the nomenclature of the Lemurs. :
requires revision.

238 DR. P. CHALMERS MITCHELL ON THE

extreme distal end of the hind-gut was much expanded and
was followed by a narrower portion strongly sacculated. I do
not attach much importance to this; I have found the corres-
ponding region sacculated, apparently as an individual peculiarity,
in several éxamiples of Lemurs and Monkeys, and Dr. Beddard
(Beddard, 1908, p. 576) has recorded a similar but more extensive
sadculation in the case of a Baboon. he distal portion of the
recurrent limb of the pendant loop gives rise to a long loop
(C.L.1 in text-fig. 29, Cl.1 in text-fig- 30) of which the proximal
and distal limbs are ‘held together by a very narrow expanse of
mesentery. This loop, lying just to the right of the representa-
tive of the transverse colon, corresponds with what is termed
the ansa coli devtra. Its presence asa defined narrow loop is
most marked in Chiromys and the genus Lemur. Dr. Beddard
states that it is absent in J/icrocebus (Beddard, 1908, p. 579),
and although I cannot agree with that author in making, in fact
or in theory, so sharp a distinction between loops that are
wide and loops that are narrow, I do agree that the asa coli
dextra is absent in Chiroc ellen: Dr. Beddard also attaches
importance to the fact that the ans coli deatra is straight in
Chiromys and Lemur, and spivally twisted in Galago, Loris,
NVycticebus, Indris, and probably Perodicticus. As I have stated
already in this communication, | cannot follow Dr. Beddard in
attaching much importance to the presence of a spiral mode of
packing any portion of the intestinal tract, unless this common
erowth-form attain a precise complexity. Nor can I agree that
the spiral of Lemurs can be taken as the rolling up of a defined
narrow loop. On reference to the original laboratory sketches
from which the diagrams of the patterns of Galago and Pero-
dicticus (Mitchell, 1905, figs. 41, 42) were made, and from
further observations made since, iL find that the loop i in question
may be very wide and irrecular, presenting, when dissected out,
2a number of minor loops, as in at least one example of Galago,
or two or more Joops as in Perodicticus. These, however, are
folded against each other and against Meckel’s tract, and rudely
twisted up. I agree, however, “that it is possible to contrast
Chiromys and Lemur with other genera possessing a well-marked
ansa coli dextra, by saying that in the former the loop in
question is characteristically narrow and straight, and in the
latter that it tends to be spirally twisted. 1 should add to
this, that in the latter it also tends to be wider and more
irregular, and that the spiral twisting varies considerably in its
extent.

Immediately distad of the ansa coli dewtra, whether that be
straight or twisted, the recurrent limb of the pendant loop
reaches its highest point, and then, in the region corresponding
with the transverse colon, sweeps backwards to form the rectum.
The proximal ponuign of this gives rise, both in Chiromys and in
Lemur (text-figs. 29, C.L.2; 30, Cl. 2), to a well-marked loop
neither so long! nor so narrow relatively as the ansa coli dextra,

INTESTINAL TRACT OF MAMMALS. 239

but forming an ansa coli sinistra. Dr. Beddard does not refer to
this in the case of Chiromys, although it is indicated in his
figure (Beddard, 1908, p. 150), but this omission is no doubt due
to the fact that he does not recognise a loop as a distinct entity
unless it has attained a certain degree of definiteness, and
especially when it is “ fixed” by some ligament other than its
primitive mesentery. I apprehend that the expansion of the region
of the hind-gut, just distad of the summit of the pendant loop in

Text-figure 30.

Intestinal tract of Lemur ? species.

Lettering as in text-fig. 29.

my figure of Chirogaleus (Mitchell, 1905, fig. 40), vepresents in a
still less defined condition the ansa coli sinistra, and is not
identical with the ansa coli dextra marked C.L. in figure 39 of
the same comniunication. Burmeister’s figure of the intestinal
tract of Varsius (Burmeister, 1846) is not easy to interpret, as
the gut has heen freed from the mesentery, but it seems probable
that there is no trace of an ansa coli dextra or sinistra but a
vather wide sweep representing a transverse colon. A figure
given by Klaatsch (Klaatsch, 1892, pl. xxiii. fig, 8) confirms this
interpretation. The rectal portion of the hind-gut in Prosimiz
runs a straight course to the anus, distad of the ansa coli sinistra,
if that be present.

As Dr. Beddard has described, there ave several secondary
connections or ligaments in the intestinal tract of Prosimiz.

24() DR. P. CHALMERS MITCHELL ON THE

These appear to me to vary considerably from individual to
individual, but the most notable of them are an attachment of
the omentum to the part where the distal limb of the pendant
loop bends round to pass into the rectal portion of the hind-gut,
and various attachments between the duodenum and the colon.
In 1905 I summed up the description of the gut-pattern in
Prosimize as follows :—‘t The duodenum is seldom well distinct:
from Meckel’s tract; the latter varies in length, probably in
relation to diet. The cecum is always present, and is usually
very capacious. The hind-gut (except in Z'arsiws, where it is
extremely reduced and still shows signs of former differentiation)
is relatively extremely long, sometimes being as long as, or
longer than, the fore-gut. It is, moreover, of greater calibre.
It shows a well-marked colic region which may be a long narrow
loop, or a complex set of folds, and a distinct rectum.” I now
add to this a few points. The duodenum is frequently marked
off by its greater calibre. The hind-gut is much reduced in very
small Lemurs such as Chirogalews and Mierocebus ; in others it
shows a definite ansa coli dextra developed from the distal
portion of the pendant loop, usually long, narrow, and straight
in Chiromys and Lemur, wider and more irregular and tending
to be spirally twisted in at least most other genera. An ansa
coli sinistra, shorter and usually wider than the ansa devtra, is
frequently present on the proximal part of the rectum.

Order SIMI&.

I have no new observations to report, although I have
examined a number of other Apes and Monkeys. For con-
venience I may quote my former summary (Mitchell, 1905,
p. 515):—“ The duodenum and Meckel’s tract together form a
series of loops which differ from group to group in their relative
complexity, arranged round about three-quarters of the circular
outgrowth of mesentery. The cecum is always present and
appears to have been originally capacious and of nearly equal
calibre throughout its length; but it is in process of shortening
throughout the group, being, as a rule, shorter in the Old World
Monkeys than in the New World Monkeys and Anthropoid
Apes (if in the latter case the vermiform appendix be reckoned
with the cecum). The state of the case may be put in another
way. The originally long, capacious cecum of the Simie is
retained by the greater number of the Platyrrhine Apes ; in the
Catarrhine Apes, except the Hylobatide and Anthropomorphe,
it tends to become shorter without the formation of a vermitorm
appendix. In the two groups last named, its proximal portion
has remained capacious, but the greater part of its original
length has been transformed without shortening into the thick-
walled vermiform appendix.”

IT should add to this that the presence of a rather well-
pronounced transverse coion is the normal condition in the group,

INTESTINAL TRACT OF MAMMALS, QA1

and that the rectal portion is usually rather longer than the
length that it has to traverse and is thus thrown into occasional
minor folds.. The colon presents no definite expansions that can
be compared with the ansa coli dewtra et sinistra, but the width
of the transverse region suggests derivation from a condition in
which both these loops were present. It is not difficult to see
in the pattern of the Simiz a condition that might have been
reached by reduction from the Prosimian pattern.

. General Conclusions.

In my earlier memoir (Mitchell, 1905) I dealt at some length
with the inferences that seemed to follow from my observations,
and | propose now to deal only with matters that call for addition
or modification. It may be useful to say, in the first place, that
the figures I gave formerly and those in this communication are,
in the strictest sense, diagrams. That is to say, they are inter-
pretations, not exact reproductions of the precise details of the
individual specimens. So far as I know, they give a fair
presentment of the significant features of the different patterns.
They form, I hope, a good basis for intensive study of the
details. It must be remembered, however, that I have attempted
to represent the primitive continuous mesentery of the gut, and
that, in actual fact, especially in the more elaborate types of
intestinal tract, portions of this mesentery have disappeared.
Notwithstanding the work of Klaatsch (18%2) and others, much
intensive study of individual types is still required to trace the
precise portions that have been lost or retained. Moreover, I am
certain that detailed study of the blood-vessels, after careful
injection of fresh material, would yield useful results. It is
well known that the mesenterial arteries and veins vary con-
siderably in man, and doubtless this also is the case in other
mammals. None the less, the general arrangement of the blood-
vessels appears to me, on such slight study as J have been able to
give, to follow the main morphological features of the gut-pattern,
and in a number of cases where one region of the gut is difficult
to distinguish from another, as, for Singita ae. in the Bears, where
there is no cecum to mark the boundary between the ileum and
the ansa coli dextra, the arrangement of the vessels in two
groups clearly delimits the regions. I hope that in my diagrams
the main features of the grouping of the blood-vessels are given,
but very much more work than | was able to give is required.

The Cecum.—Further work has confirmed me in the opinion
that the cecum of Mammals is one member of a primitive pair,
homologous with the paired ceca of Birds. I have already
sufficiently stated the facts that lead to this conclusion (Mitchell,
1905, p. 515), but I may refer to a curious side-light on the
subject. Baleeniceps is one of the few birds in which ‘the normal

Proc. Zoou. Soc.—1916, No. X VI. 16

249, DR. P. CHALMERS MITCHELL ON THE

pair is represented by a single cecum, and in this case it happens
that the surviving cecum is thin-walled and relatively capacious.
If the figure of the cecum of that bird (Mitchell, 1913, text-
fig. 123) be compared with the normal unpaired cecum of
Mammals, it will be seen that the resemblance is very close.

Position of the Cecwm.—The most common position for the
ceca in Birds is distad of the pendant loop on the straight
portion of the hind-gut close to the cloaca. This position I
associate with the progressive shortening of the hind-gut, which
is a striking feature in avian anatomy as we turn from less
specialized to more specialized. types. In the lower types, in
which the rectum is. relatively longer, the ceca are placed more
proximally on the hind-gut. In the Ostrich (Mitchell, 1896,
fig. 4), for instance, where the fore-gut and hind-gut are more
nearly equal in length, the ceca occupy a position almost identical
with that of the paired ceeca in the Manatee or the single cecum
of the Elephant. In no case, however, are they proximad of the -
distal end of the pendant loop. Among Mammals the most
frequent position is about the middle of the recurrent limb of
the pendant loop. It is a striking coincidence, however, that in
the only Bat with a cecum that I have seen, the position is so
close to the distal extremity of the pendant loop, that it may be
described as occupying an avian position. In Varsius (according
to Klaatsch, 1892, pl. xxi. fig. 8) the czecum is not at the distal
end of the pendant loop. In the Carnivores, among which, as
among Birds, there is a progressive degeneration of the hind-
gut, the cecum, although on the recurrent limb of the pendant
To op, 1s very close to its distal extremity. In the Cetacea the
other extreme is present; the cecum lies almost at the pr oximal
end of the recurrent limb of the pendant loop. The various
positions of the cecum in Birds and in Mammals nearly overlap,
but the most frequent position in the one case is distad of the
pendant loop, in the other somewhere on the pendant loop, a
state of affairs congruous with the idea that the various conditions
have come about by divergent modification from a common
type.

Form and Function of the Cecwm.—I1 Bau nothing to add to
my former discussion (Mitchell, 1905, p. 522). Only in a most
general sense can there be said to be a correlation between diet
and the presence, length, and capacity of the cecum. ‘There are
many exceptions to any general statement, and it seems as if
ancestral history were at least as potent a factor as actual diet.

Secondary Relations between Proximal and Distal Portions of
the Intestinal Tract.—Vwo different kinds of connection may
exist between proximal and distal regions of the intestinal tract.
The connection to which 1 have paid most attention, and

INTESTINAL TRACT OF MAMMALS. 243

which I gave a long account (Mitchell, 1905, p. 524) was that in
which blood- vessels belonging to one region of the gut supply
another region with splined a may he. in contact, although
morphologically remote. In Birds the folding of the gut prings
the distal portion of Meckel’s tract im alos contact with the
duodenum, and it frequently comes about that branches of
the duodenal blood-vessels may form the main supply of the
portion of Meckel’s tract just proximal to the ceca, and may
have to be severed before the whole gut can be unfolded. In
Mammals the connection, when it exists, links the colic region to
the anterior part of the gut. I wish to modify the table ‘Teg gave
only by omitting Ornithorhynchus; from examination of aune her
example, I am far from certain as to the existence of a true
“ short-circuiting ” blood-vessel, and the point could be settled
only by examination of fresh injected material. The cases, then,
in which this peculiar condition of the blood-vessels certainly
exists are the Traguloidea, Tylopoda, Pecora, Rodents, Lemurs,
and Simiz. If one considers it, it is a curious circumstance that
in the development of man a branch of the superior mesenteric
artery should leave its normal course and thrust itself out
to reach the transverse colon. Instead of explaining this as
an instance of some marvellous coordinating vitalistic power, I
prefer to think that it is a legacy from the past, and that the
ancestors of the Simize had a more complex colon with loops
pressed against the mesentery of Meckel’s tract, as occurs in
some of the Lemurs. Jn this connection it is interesting to note
that Klaatsch found a Lemur-like stage of the colon in the
embryo of Hapale (Klaatsch, 1892, p. 671, fig. 12, cited by
Beddard, 1908, p. 598).

There are also connections of a more mechanical kind between
different portions of the gut. These are the various ‘‘ligaments”
and attachments to which I have frequently referred in this
communication. They were not included in the table in my
paper of 1905. Notwithstanding the elaborate work of Klaatsch
(Klaatsch, 1892), and Dr. Beddard’s later discussion (chiefly
Beddard, 1908, p. 568 eé¢ seqwitw), I cannot form a. clear con-
ception of the distribution of these structures among Mammals,
and I have not myself made a connected investigation of them.

Loops of the Hind-gut.—\ have already drawn a contrast
between the gut-patterns of Birds and Mammals, depending on
the broad fact that, even when allowance has been made for the
homoplastic modifications associated with diet (Mitchell, 1905,
p- 526), in Birds Meckel’s tract and in Mammals the hind-gut
tend to display specialized subsidiary loops of systematic im~-
portance. In Birds, however, the loops of Meckel’s tract have
reached a high degree of stability, so that they vary little within
well-defined systematic groups, whereas in Mammals the loops of
the hind-gut vary much more within narrow systematic limits,

as if they. were in much closer relation with habit or diet. The
1§*

244 DR. P. CHALMERS MITCHELL ON THE

facts do not seem to justify too close an identification between
the specialized loops in one mammal and another. I propose,
however, to give a tentative summary of the conditions.

In Monotremes there is an ansa dextra near the distal end of
the pendant loop, and the rectum is straight.

In Marsupials the rectum is relatively long and may be thrown
into minor loops. In the Polyprotodonts there are no other
expansions of the hind-gut. In the Diprotodonts the usual
condition is the presence of a complex ansa sinisira, and there
may be in addition, as in the Phascolarctide, an equally complex
ansa dextra.

In the Edentata the rectum is always relatively long; in the
Pholidota there is no further expansion. In the Tubulidentata
and Xenarthra there is also an ansa sinistra.

In the Hyracoidea, Sirenia, and Proboscidea the rectum is
relatively long, especially in the region just distad of the pendant
loop, and therefore forming an ansa sinistra.

In the Cetacea the rectum is straight, and there is no ansa.

Among the Ungulata vera, the rectum is always longer than
the distance between the distal end of the pendant loop and the
anus, and in the majority of the sub-groups the lengthening
is most marked proximally, although, perhaps, not enough
specialized to be regarded as corresponding with an ansa
simstra. An ansa paracecalis or postcecalis is present, just
distad of the cecum; in most of the Pecora, absent in the others,
but its presence, in addition to the well-known colic spiral, makes
it impossible to identify the latter with the paracecal loop.
The recurrent limb of the pendant loop always forms at least
one large ansa dextra; this is complex in the Hippopotamus,
and forms a spiral in the Swine, Traguloidea, Tylopoda, and
Pecora, and a very long narrow loop in the Perissodactyla.
In the Traguloidea, Tylopoda, and Pecora there is a second
more distally placed ansa dextra, folded closely against Mecket’s
tract between the colic spiral and the minor loops of the tract.

In the Rodentia there is almost invariably a paracecal loop
often spirally twisted, with the cecum or independently of it,
always at least one and frequently two anse dextre, which may
be straight, or spirally twisted, together or independently, and
pressed against Meckel’s tract. An ansa sinistra is frequently
present, either as a definite narrow loop, or as a complex loop,
and the latter condition grades off into a wavy condition of the
rectum, which in all Rodents is longer than the distance it has
to traverse.

In the Insectivora the rectum is short and straight, but a
definite ansa dextra is usually developed.

In the Chiroptera the whole hind-gut forms a short straight
rectum, and there are no anse.

In the Carnivora the rectum, although relatively short, is
usually longer than the distance it has to traverse (between the
distal end of the pendant loop and the anus), and very often

INTESTINAL TRACT OF MAMMALS. 245

presents minor loops at its proximal end, which may be grouped
so as to form an ansa sinistra.

In the Bears there is, in addition, a definite ansa dextra.

In the Prosimiz the rectum is always longer than the distance
that it has to traverse, and at its proximal end, close to the distal
extremity of the pendant loop, there is fr equently a special
expansion, forming an ansa sinistra. An ansa dexctra, usually
large in size, sometimes narrow, sometimes complex and ee
doubled, sometimes straight and sometimes spirally twisted, 1
present in all except a few very small forms.

In the Simiz the rectum is always longer than the distance it
has to traverse. The proximal region of the hind-gut, composed
of the pendant loop distad of the cecum, the colic apex and the
proximal portion of the primitively straight rectum are gradually
approaching the human condition of nearly straight ascending,
transverse, and descending colons, the appearances suggesting
that this condition has been reached through a more prosimian
stage in which there were definite anse coli deatre and sinistre.

Systematic Inferences.—In this section I propose to deal only
with the facts to which I have myself paid attention. Un-
fortunately Iam unable to follow, from Dr. Beddard’s descriptions
and figures, exactly what he means by the “stages of evolution of
the intestinal part of the alimentary tract,” and so cannot
attempt to correlate them with my own results. Stage I.
(Beddard, 1908, p. 591, text-fig. 120 A) represents a condition that
is at least more primitive than in any known mammal. Two
figures are labelled Stage II. (doc. cit., text-figs. 120 B and 121), and
differ in that the second fgue shows rotation of the gut; butin
each figure the so-called ‘‘cavo-duodenal ligament” is “Geer and
labelled, although in the text its existence is stated to be due to
the rotation. In the later figures large portions of the gut are
represented as without any mesentery, and much of the mesen-
tery that is represented shows relations which I am unable to_
follow. Zoologists who wish to follow what is known as to the
mode in which the rotation of the gut affects the primitive
mesentery will find admirable descriptions and figures in the
ordinary text-books (as, for instance, Professor D.J. Cunningham’s
‘Text-book of Anatomy,’ 1902, pp. 1056, 1057, figs. 711, 712).
Klaatsch (1892) is still the best authority on the secondary
ligaments and attachments; but I cannot always follow him ir
the discrimination between portions of the primitive mesentery
and secondary attachments, and suspect that much further
investigation is required.

I am inclined to think, however, that rotation is due largely to
simple mechanical causes, and that it is therefore an event that
may have occurred repeatedly and independently, the resem-
blances caused by it being due not to inheritance from one
ancestor in which rotation had occurred, but to a similar effect
producing similar results on similar material. As Meckel’s tract

946 DR. P. CHALMERS MITCHELL ON THE

lengthens, its closely bunched set of minor loops, developed
chiefly on the proximal limb of the pendant loop, must push
their way towards the middle line dorsal to the distal locp,
which in primitive mammals hangs more freely down in the
gut. .
Taking only the characters presented by the gut-patterns as a
basis, it appears that the most primitive or generalized type had
a duodenum not well separated from Meckel’s tract, Meckel’s
tract, consisting of minor loops developed along the proximal
limb of the pendant loop, up to about the apex where the yolk-
sac was attached, a moderately straight recurrent limb bearing
towards the middle of its length a functional cecum (or more
probably a pair of functional ceca), a sharply bent colic flexure
close to the duodenum, where the pendant loop passed into the
rectal portion of the hind-gut; that rectal portion considerably
longer than the length that it had to traverse, and thrown into
specially long minor loops at its proximal extremity. Apart
from rotation, the gut was suspended on a continuous primitive
mesentery, and the blood-vessels supplied the regions of the gut
to which they belonged. Changes from this primitive condition
occur in two directions: the pattern may be secondarily re-
duced and become even more simple, or it may become more
elaborate.

When characters are used for the purpose of classification, it
may be convenient, in the absence of other information, to place
creatures in the same group because they have retained ancestral
conditions, but if the classification is intended to state the degree
of affinity, then it must be remembered that there is no a priori
reason to suppose that amongst the descendants of a common
ancestor, the groups that have retained an ancestral character
are more closely related than the groups that have lost it. On
the other hand, the common possession of a well-marked elabora-
tion of the primitive type appears to present some ground for
unplying affinity.

As in my summary of 1905, I associate the Marsupialia,
Xenarthra, and Tubulidentata as displaying, on the whole, the
most ancestral type of gut-pattern, with the proviso that this
association does not imply close aftinity but merely the retention
of a common inheritance. I think it safer to exclude the Galeo-
pithecidee from this association, as my information with regard
to that Order is second-hand. I note with regard to the Mar-
supials, however, that they contain two departures from the
ancestral type. In some of the small Polyprotodonts the gut-
pattern is extremely reduced, with complete loss of ceca and
obliteration of clear distinction between the different regions. In
other Marsupials, such as the Phascolarctidee, the hind-gut has
attained an elaboration recalling that of higher types. In the gut,
as In many other parts of thei structure, the Marsupials appear
to forecast, on a lower level, and in a more fluctuating condition,
elaborations that become definite and ‘ fixed” in higher types.

INTESTINAL TRACT OF MAMMALS, 247

As it is difficult to suppose that the different types of organs in
higher Mammals have arisen separately from the corresponding
types in Marsupials, we are faced with the poset any that
organisms may have inherited the possibility of isplaying
definite variations that have not actually appeared in their
ancestral history, a possibility which, as Arthur Willey has
shown (Willey, 1911), has not yet been sufficiently considered in
systematic zoology.

I have also to note that the characters of the gut-patterns
afford no ground for grouping together the so-called Edentates.

It is interesting to note that amongst other primitive characters
this first group of mammals contain abundant relics of what L
take to be the primitive paired condition of the ceca,

The Monotremes have not moved far from the primitive type,
but in a definite direction. The duodenum is distinct; the
cecum is single but degenerate, and is placed very close to the
apex of the pendant loop, the distal limb of which displays a
compound ansa coli dextr a, and the rectal portion is relatively
short and straight.

The Pholidota, although not far removed from the primitive
type, have lost the cecum, and have a longitudinally striated
hind-gut which is unique.

The Hyracoidea, Sirenia, and Proboscidea have not moved far
from the common type, and J realize that their association may
depend very largely on their retention of primitive characters.
In all, the primitive mesentery is very complete and continuous,
and although the hind-gut is long, there are no specialized loops.
The unpaired cecum of Hyracoidea, if [ am correct in refusing
to identify it with the unpaired cecum of other mammals, is a
peculiarity unique among mammals; but apart from that, and
taking the paired ceca as the true representatives of the normal
structure, there is a very close resemblance between the pattern
of Hyracoidea and that of the Manatee. The chief difference is
the further increase of length of the hind-gut, distad of the
pendant loop, in the Manatee. The Dugong is said to havea
gut-pattern of the .same type as that of the Manatee, except
that the cecum is single, and sucha pattern leads directly to
that of the Elephants, in which the cecum is unpaired and the
hind-gut thrown into long irregular minor loops, so that it
appears to be almost as long as the fore-gut. There is no
trace of any of the peculiarities of pattern found amongst the
true Ungulata.

In considering the Cetacea, the first feature of importance 1s
that the Odontocete pattern is most easily explained as a
derivative by reduction of a pattern such as is found in the
Mystacoceti. The Mystacocete pattern, although peculiar, is
not very far removed from the primitive type, but the great
elongation of Meckel’s tract with the ceecum at its apex, the long,
nearly straight, recurrent limb of the pendant loop, and the long
but nearly straight rectum, make up a divergence from the

248 DR. P. CHALMERS MITCHELL ON THE

primitive type quite unlike the divergence found in any other
group, and support the supposition that the Cetacea diverged
from other Eutherians at a very remote period.

I am not now prepared to associate the Perissodactyla and the
Artiodactyla on the evidence afforded by gut-patterns. Both
groups may well have arisen independently from the common
stock. In all the Artiodactyles there has been a great lengthening
of the portion of the hind-gut formed from the distal limb of the
pendant loop. In Hippopotamus this lengthening is irregular ;
in the Suide it forms a definite spiral. In the Traguloidea,
Tylopoda, and Pecora this spiral reappears but gradually be-
comes more specialized and more intimately associated with the
mesentery and blood-vessels of Meckel’s tract. There may be
another expansion of the same region proximad of the spiral,
forming an ansa paracecalis, and another distad of it forming a
second ansa coli dextra.

In the Perissodactyla the whole of the recurrent limb of the
pendant loop distad of the large cecum gives rise to an enormous,
narrow, ansa coli dextra, an arrangement quite different from
that in any other group, It is certainly remarkable to find that
the herbivorous Perissodactyles have developed a type of gut-
pattern extremely like that of the herbivorous Artiodactyles,
unless we are prepared to think that adaptation plays only a
secondary part in the matter.

Among the Rodents we might expect to find convergent
resemblances with Artiodactyles, but these are quite superficial.
The mode in which the hind-gut is lengthened, the spiral twisting
of the cecum, its relation to the paracecal loop, the frequent
doubling of the ansa coli dewtra, and the frequent appearance
of an ansa coli sinistra compose a general picture quite different
from that of the Artiodactyla or Perissodactyla. So far as gut-
pattern is concerned, the Rodents may well have arisen as a
distinct outgrowth of the primitive stock.

The gut-patterns of Insectivora are consistent with the con-
ception that there has been a secondary reduction or simplification
within the group from such a modification of the primitive
Mammalian type as is seen in Macroscelides, The pattern of
Macroscelides might belong to any simple Marsupial or Mono-
treme-like creature; it differs from the Marsupial patterns
most closely resembling it, by the presence of an ansa coli
dextra instead of an ansa coli sinistra, and from the Monotreme
pattern in having the cecum some distance from the apex of
the pendant loop instead of very close to it,

The examination of one of the Chiroptera in which the cecum
is present has enabled me to distinguish between the very simple
patterns of Insectivora and of Chiroptera. In Chiroptera the
whole of the pendant loop becomes Meckel’s tract, and the hind-
gut is reduced to an extremely short and straight rectum, No
doubt the bird-like shortening of the hind-gut is a secondary

INTESTINAL TRACT OF MAMMALS. 249

divergence from the primitive type, but this would not affect the
position of the cecum, and quite certainly the gut-pattern offers
no argument for any close association between Chiroptera and
Insectivora.

The gut-pattern of Carnivores, notwithstanding the enormous
lengthening of the fore-gué in the Fissipedia, has moved little
from the primitive type, consisting of a fore-gut developed from
the greater part of the pendant loop, and a hind-gut, which,
although relatively short, is always longer than the length that
it has to traverse and not infrequently presents an expansion
that may be termed an ansa coli sinistra. The Bears are the
only exception to the coherence of the picture; they have no
cecum, but the anatomical relations seem to show that their
cecum was originally placed much more near the proximal end
of the recurrent limb of the pendant loop, and that the distal
portion of that loop has given rise to an ansa coli dextra absent
in the other groups. The true explanation may be that the
apparent simplicity of the pattern in other Carnivores has come
about by reduction,

The Prosimian pattern is not far removed from the primitive
type, but tends to the development of at least one minor expansion
of the recurrent limb of the pendant loop, an ansa coli dextra,
which may be straight or spirally coiled, and there may also be
an ansa colt sinistra.

The Simian pattern is best understood as derived from the
Prosimian pattern by reduction.

If the gut-patterns were our sole source of information as
to the inter-relationships of existing mammals, I do not think
that we could get much further than is set forth in the annexed
table, in which little stress must be laid on the vertical
arrangement :—

— Pholidota.

Monotremata.
| f Hyracoidea,
| J Sivenia.
Proboscidea.
Mystacoceti—Odontocet.
Marsupialia. Hippopotamidee. :
Sapnagas hin. { ae ie " } Traguloidea.
Tubulidentata.

{ Tylopoda.

Pecora.
Perissodactyla.

Prosimiz—Simie.

Rodentia.

Insectivora.

| Chiroptera.

__ Carnivora.

250 DR, P. CHALMERS MULCHELL ON THE

List of References.

Brpparp, F. E.—On the Anatomy of Antechinomys and some
other Marsupials, with special reference to the Intestinal
Tract and Mesenteries of these and other Mammals.
PEAT selsvsyproole

Zd.—Some Notes on the Anatomy of Chiremys madagascariensis,
with references to other Lemurs. P. Z. 8. 1908, p. 694.

fd.— Contributions to the Anatomy of certain Uneulater in-
cluding Zapirus, Hyrax, and Antilocapra. P. Z. 8. 1909,
p. 160.

Bepparb, F. E., & Treves, F—On the Anatomy of the Sondaic
Rhinoceros. Trans. Z. 8. 1887, vol. xii. p. 183.

Broom, R.—On the Organ of Jacobson in the Elephant-Shrew.
P. Z.8. 1902, vol. i. p. 22¢

f¢d.—On the Organ of Taeobeon and its Relations in the
Insectivora. P. ZS. 1915, p. 1573 et é. ce. p. 347.

Burersrer, H.—Beitrage zur Kenntniss der Gattung Zarsius.

: Berlin, 1846.

Frower, W. H.-—On the Anatomy of Proteles cristatus. P. ZS.
1869, p. 474.

/d.—Lectures on the Comparative Anatomy of the Organs of
Digestion in the Mammalia. Medical Times & Gazette,
1872. ,

Garrov, A. H.—'The Visceral Anatomy of the Sumatran Rhino-
ceros, § P. ZS!) 1873) pr 92.

GeorcE, M.—Monographie Anatomique des Mammiféres du
genre Daman. Ann. Sei. Nat. 1874, ser. 6, Tome I.

Kauuia, H.—Monographia Hyracis. Tiibingen, 1830.

Kuaatscu, H.—Zur Morphologie der Mesenterialbildingen am
Darmkanal der Wirbelthiere. Morph. Jahrb. 1892,
pp. 385 et 609.

LonnBerG, Dr. E.—On the Soft Anatomy of the Musk-Ox
(Ovibos moschatus). P. Z. 8. 1900, p. 142. —

Jd.—Material for the Study of Ruminants. Nov. Act. Reg. Soc.
Sei. Upsala, Ser. 3, Vol. xx. 1903

JId.—Some Comparative Notes on the Anatomy of the Elk (dlces
alces). Zool, Stud. Tullberg, Uppsala, 1907 peek

Lonsky, F.— Beitriige zur Anatomie und Entwic eta geseeschichte
des Darmrohres und des Urogenitalsystemes von Hyrax.
Jen. Zeitsch, 1903, vol. xxxvii. p. 579.

MircHett, P. CuHaAtmers.——On the Intestinal Tract of Birds.
BEA Sels9ey pulse:

Jd.—On the Intestinal Tract of Birds. Trans. Linn. Soe. Lond.
(Zool.) 1901, vol. viii. p. 173.

Zd.—On the Occasional Transformation of Meckel’s Diverticulum
in Birds intoa Gland. P. Z. 8. 1903, p. 352.

/d—On the Intestinal Tract of -Manmals. Trans. Z. 8. 1908
vol. xvii. p. 437

INTESTINAL TRACT OF MAMMALS, 251

MircuEi, P. CoaAtmMErs.—Observations on the Anatomy of the
Shoe-bill (Baleniceps rex) and Allied Birds. P. Z. 38.
1913, p. 644.

Oppet, A.—Lehrbuch vergl. mikros. Anat. Wirbelthiere. Part IT.

1897.

Owen, R.—-On the Anatomy of the Cape Hyrax. P. Z. 8. 1832,
p. 202.

Id.—Anatomy of Vertebrates. Vol. IJ]. Mammals. London,
1868.

TuLLBERG, TycHo.—Ueber das System der Nagethiere. Nov.
Act. Reg. Soc. Sci. Upsala, 1899.
Witiey, ArrHuR.—Convergence in Evolution. London, 1911

Ce a
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ON LICE FROM THE SOCIETY 8S GARDENS.

7. Studies on the Anoplura and Mallophaga, being a
Report upon a Collection from the Mammals and Birds
in the Society’s Gardens.—Part L., with a Preface,
By Brucr F. Cummines, British Museum (Natural
History) *.

[Received January 5, 1916: Read February 22, 1916. |

(Text-figures 1—24.)

INDEX.

SYSTEMATIC : Page
Wrinognatius pithodes, Sp. We) ..icry-s.22.efs cer and -eeereee-eec cess 200
Prichodectes hemstrags, SP. MW. ......-+-0veeee eve eee vente cece 213
FEI RAGPOSGTDON SjeboiTo:- vetidedeoek Wuecind node GANNencses yous OoddtbeseecebeaEtn | be (5

STRUCTURE.

DEVELOPMENT.

Preface.

The following paper is the first part of a report upon the
Anoplura and Mallophaga collected upon the Mammals and
Birds that have died from time to time in the Society's Gardens,
and kindly submitted to me for identification and study by
the Zoological Society of London. On account of the oppor-
tunities for straggling on to other species of host afforded to
ectoparasites on animals in a menagerie, the correct names
of the hosts in some instances may be open to doubt, while the
labour involved in the identification of the specimens is thereby
much increased, particularly in two such groups as these, which
in greater part remain in a state of considerable systematic
confusion. In the present collection, however, I have come
across very little evidence of straggling, and, in any case, such
a contingency is amply compensated for by the fact that through
the kind offices of the late Proseetor, Dr. F. KE. Beddard, F.BR.S.,
and the Society’s Pathologist, Professor H. G. Plimmer, F.R.5.,
much of the material has been fixed and carefully preserved
in alcohol. It has thus been possible to add to the knowledge
of many of the forms coming under review and to turn the
report into something more than a dull census of names. The
value of the systematist’s routine work would be greatly
enhanced if an endeavour were constantly made to add at least
something, however small, to the pre-existing knowledge of the
morphology of-each species. The correct naming of an animal
is the systematist’s opportunity—too frequently neglected.

In entomology of late, those who observe the Heavens will
have seen certain signs in the sky, one being the growing demand
for spirit-specimens part passu with the growing recognition
that there is no group of insects in which a duplicate series of

* Published by permission of the Trustees and communicated by the SECRETARY,
A J

254 MR. B. F. CUMMINGS ON LICE

spirit-specimens is not essential for systematic study; while in
many groups, such as those here considered and the Ephe-
meride, pinned or carded material is for the purposes of study
almost worthless. Another portent is the obsolescence of the
short, superficial Latin diagnosis in the description of new
species—the persistent remains of the early influence of the
schoolmen on systematic zoology. In some quarters the super-
ficial diagnosis is already swept into limbo with the Sedan
chair—both of them vehicles which could not possibly take us
far, as trafliic runs nowadays.

As précis work the superficial diagnosis is admirable. <A
certain number of characters from a certain restricted area
of an animal’s anatomy (as a rule, the easily accessible external
parts) are selected and then welded into a cast-iron paragraph
as exact and inflexible as a lawyer’s deed-poll. But in ento-
mology, as in politics, the lawyer is a strong but undesirable
element, and the only justification for the superficial diagnosis is
that the descriptive writer sometimes may find it convenient as
a summary to tag on to the end of a long description.

The systematist 1s primarily a morphologist whose task it is
to discover the true phylogenetic relationships, at least between
the small subdivisions such as genera and species. This is a
responsible task, especially in entomology, requiring the dis-
section, careful morphological examination, and description of
each species. Many characters which have to be dissected to be
seen are already in use among systematic entomologists, ¢. g.,
the male genitalia, the spermatheca, the mouth-parts, and endo-
skeleton. It is not suggested, of course, that internal or con-
cealed characters are “necessarily netter than external ones.
What are the best characters for systematic use is a question
which has to be investigated and settled for each individual
group of the Animal Kingdom. Quite recently Dr. P. Chalmers
Mitchell wrote * :—“ The more experience I gain of avian
anatomy, the more I am convineed that systematists are well
advised when they rely, at least with regard to the discrimination
of species and genera, more upon those superficial characters
that they can observe in the series of museum collections than
on the uncertain indications afforded by the presence or absence
of this or that muscle.”

Yet, how many systematists have conscientiously gone through
the anatomy of their group with a view to settling this fundamental
question of the best combination of characters for the purposes
of classification (for I imagine that single-character classification,
such as Huxley’s palate and Garrod’s ambiens among birds is now
discredited). The descriptive writer is an opportunist who, as a
rule, seizes upon the most convenient characters that offer them-
selves. Certainly, it is in entomological opportunism that we
must seek the cause of the present systematic disorder among the

= PLS. 19s, p. 43;

BS

FROM THE SOCIETY'S GARDENS. - a)
Anoplura and Mallophaga, for recent authors alone have agreed
to treat these two groups from the morphological standpoint *.

Lest in the following essay it be urged, perhaps, that I have
recke | not my own rede, it must be explained that the amount
of anatomieal work which a systematist is able to accomplish on
any one species is governed by the nature of the material—its
quantity and quality. No attempt has been made to dissect
species of which only one or two specimens were available, and
in no case has it been possible to attempt the anatomy of the
soft parts, as Part I. deals mainly with those specimens—in a
poor state of preservation—which formed the nucleus of the
collection handed over to me in the beginning from another
worker who was prevented from carrying out the work.

ANOPLURA.
Genus Prepicutus Linné.

PrpicuLus capitis De Geer (1).

Piaget (2, p. 626) described as a new species of Pediculus a
form taken on Ateles pentadactylus Is. Geoff. from the collection
of the Museum of Leiden. P. consobrinws, as it was named, is so
close to P. capitis that one searches Piaget’s text and ficures in
vain for reliable distinguishing characters, and Neumann
(3, p. 440) concludes that P. consobrinus cannot be clearly
separated from P. capitis. Without Piaget’s types before me,
it is impossible to be certain about this, but that P. capitis does
on ogcasion in menageries pass from man to monkeys is shown
by specimens captured in the Gardens on the Red-faced Spider
Monkey (Ateles paniscus Linn., Family Cebide).

PrprcuLus AFFINIS Mjoberg f (4, p. 169).

In large numbers on Ateles paniscws Linn. (Family Cebidee)
together with larve. Ina lot distinct from the preceding.

Fahrenholz (5, p. 8) has described the larve of Baaliondlons
capitis, with which the larve of P. affinis agree very closely, not
only in the 3-segmented antenna, but in details of abdominal

* The whole question of the methods of research employed by the systematic
zoologist is discussed im an anonymous paper published in ‘The American
Naturalist’ for May 1914 (p. 369) entitled “Taxonomy and Evolution.”

+ This series of Pediculus, which undoubtedly belongs to the form named
P. affinis by Mjoberg, proves on examination to be a very inconsiderable variety
of P. capitis. Every one of the characters given by Mjéberg is inconstant and
occurs occasionally, I find, on varieties of P. capitis from savage races. P. affinis
is probably a straggler from human beings, establishing itself on Afteles on account
of a certain similarity in the blood and hair between Afeles and the Anthropoids as
adduced by Friedenthal. On this assumption the family Pediculide is, as far as
known at present, confined to the Old World: the Pediculinee on Man and Apes,
the Pedicinine on the lower monkeys. If it be held that P. affinis is a true
parasite and at no time a straggler from Man, then, in view of the very generat
opinion that Platyrhine and Catarhine monkeys dive reed very early from one another,
this Cebid Pediculus would have to be regarded as a very remarkable case of con-
vergence. The question will be fully dealt with in the “Annals and Magazine of
Natural History.’ 4

256 MR. B. F. CUMMINGS ON LICE

chetotaxy. Fahrenholz omits all reference to the larval chto-
taxy of the head and thorax, and it is therefore worthy of
remark that, while the head and thoracic chetotaxy (dorsal
surface) of larvee and adults are almost identical, the arrangement
of the hairs on the ventral surface of these parts in the larve of
P. affinis differs considerably, especially that of Stage I., from
that of the adult in the same regions. On the dorsal surface the
differences are negligible. But on the ventral surface of the head
in Stage J. there are but two hairs (or bristles); in Stage IT.
there are four (two in front and two behind). and two minute
hairs on each side in the preantennal area. The lower surface
of the head of Stage IIT., as regards chetotaxy, closely resembles
the adult, except that the small group of hairs behind each
antenna contains fewer hairs than in the group of the adult.
On the ventral surface of the thorax there are fewer of the
minute seattered hairs.

These facts are of some interest, being contrary to what has
heen found to obtain in the larve of three species of Polyplax
(6, p. 271), where the chetotaxy of the head and thorax is fully
adult in Stage I., while that of the abdomen in the same stage
agrees with Pediculus in presenting great differences.

Genus Prpicints Gervais.

Several species of Pedicinine, including one undoubtedly new
form from Colobws caudatus Thos. (Family Cercopithecide), are
contained in the collection. But this group is difficult and ir
some confusion, and the Society's material is insufficient to
justify any attempt to straighten things out. Fahrenholz’s
paper (5, p. 12) clears the way for a revision, which can best
be effected by a study of the male copulatory apparatus.

Only the two following species have been definitely named :—

PEDICINUS RHESI Fahrenholz (5, p. 16).

A few specimens from J/acacus inwas (now known as Pithecus
amuus Linn.) Fahrenholz’s specimens were obtained upon a
“* WVacacus rhesus.” Both hests belong to the Cercopithecide.

Genus Puruirpepicinus Fabrenholz (5, p. 22).
PHTHIRPEDICINUS MicROPS (Nitzsch) * (5, p. 25).

A single female from an unrecorded host.

Genus Potypuax Enderlein.
POLYPLAX SPINULOSA (Burm.) (7).
Numbers of specimens of this cosmopolitan species caught on
the cosrnopolitan rat (Zpimys norvegicus Erx].), The larve and

* |The parentheses around the names of authors placed after scientific names in

this paper are used in accordance with Article 28 of the International Rules of
Nomenclature (Proc. 7th Int. Cong. Boston, 1907, p. 44 (1912))—Ep110R. |

FROM THE SOCIETY’S GARDENS. 257

the male copulatory apparatus are described and figured in the
P.Z.S. 1915, pp. 256, 268, text-figs. 7, 15, 16).

Huplanation of the Terminology used in describing the Male
Copulatory Apparatus of Anoplura and Mallophaga.

In almost all Anoplura and Mallophaga, it is easy to recognise
at once the basal plate and the parameres. The basal plate—
probably double in origin as two longitudinal apodemes—is a
chitinous lamina usually, if not always, longer than broad, to
the posterior lateral angles of which are articulated the two
chitinous appendages known as parameres. Between the para-
meres is the mesosome, the parts of which are not so readily
made out unless a specimen be carefully dissected. Fundamen-
tally, the mesosome is a sac—the enlarged and extrusible end
continuous with the ductus ejaculatorius. This sac—called
by Mjoberg “the preputial sac ”—presents two regions of
chitinisation—a distal and a proximal. At the distal end is the
rod of the penis or virga, with frequently a splint on each side
called the telomere, and one below—the hypomere*. At the
proximal end are the endomeres, usually strongly chitinised
bands or rods, one on each side, supporting the membrane of
the sac, of which they are only local thickenings. The whole
of the genitalia exhibit enormous variety in form, and the
mesosomatic parts in particular are occasionally so much modified
that it becomes difficult to recognise their conformation to the
general plan just sketched out above. For example, in many
Philopterids, such as Docophorus, no sacular portion of the
apparatus is recognisable, and the distal chitinisations lie well
back within the proximal, the whole forming a solid and compact
mesosome. The above terms are, therefore, adopted solely for
‘convenience of description.

Genus LinocnatHus Enderlein.

LINOGNATHUS CAVI#-CAPENSIS (Pallas) (8, p. 37). (Text-fig. 1.)

The ample material of this species submitted provided the
opportunity for a dissection of the copulatory apparatus of the
male, which hitherto has been neither described nor figured.
The accompanying text-illustration (text-fig. 1) makes it easy
to dispense with a good deal of circumlocutionary description.

Basal plate: A fairly long and narrow rod, the posterior end
of which, for the purpose of affording articulation to the para-
meres, is widened, and each posterior lateral angle drawn out
into a short process, leaving the lower margin of the plate
concave between.

Parameres: Kach paramere at the base is fairly broad and fits
in around the concave lower margin of the basal plate. Halfway

* For these terms, first applied to specialised Philopterid forms, see Waterston,
Annals of the 8. African Museum, vol. x. pt. 9, 1914, p. 279.

Proc. Zoou. Soc.—1916, No. XVII. 17

258 MR. B. F. CUMMINGS ON LICE

down on the inner side, each paramere shows a very pronounced
thickening or nodule of chitin, At the extreme distal end the
small acute termination of the paramere bends in sharply to face
its fellow.

Text-figure 1.

Par End

P

Linognathus cavie-capensis. Male copulatory apparatus. X 378.

BP. Basal plate; Hnd. endomere; P. penis; Par. paramere.

Endomeres : These are fused at the base into a single piece.
The two limbs support the “ preputial sac,” on the lower side of
which, in the retracted condition, may be observed six large
pustular alveoli, probably containing minute directive hairs or
other form of sense-organ.

Penis: Atthe tip is an aperture, surrounded by a noose of
chitin. This is probably the opening for the seminal fluid.

Telomeres: Not separately distinguishable.

FROM THE SOCIETY’S GARDENS. 259

The hypomere is in the form of a thin flat band, running back
from the tip of the penis along the lower surface of the sac.
LINoGNATHUS LIMNOTRAGI Cummings (8, p. 36). (Text-fig. 2.)

Several specimens collected on Limnotragus gratus Scl. (Congo)
and also on Tauwrotragus oryx.

Text-figure 2.

Linognathus limnotragi. Male copulatory apparatus. X 232.
BP. Basal plate; H. endomere; P. penis; Par. paramere ;
X. cross-section along the line a—b.

The parameres are dissected apart.

The following is a description of the male copulatory apparatus,
unfortunately omitted from the original description :—
Basal plate: Rod-like, with an expanded posterior endéjust
; oe

260 _ MR. B. F, CUMMINGS ON LICE

as in the preceding species. The posterior lateral processes,
however, are less developed, and the posterior margin of the
expanded posterior end is only slightly concave.

Parameres: These are of a remarkable type (text-fig. 2).
Proximally they are broad blade-like pieces which meet each
other (but do not fuse) beneath the mesosome in a fairly long
median groove, then dorsally wrap themselves around the
mesosome lying between them, forming a kind of sheath, from
the end of which the penis projects, and, like the somewhat
narrower distal ends of the parameres, curls up dorsalwards.

Penis: Asin L. cavice-capensis, this ends m a loop-like aperture,
of much smaller size, relatively. On each side of it may be
discerned, under a high power, a small telomere, the three
together forming a complex, running backwards into a broader
basal part t between the parameres.

The part “ E” T regard as the endomeral part of the copulatory
tube. These parts are reconsidered on p. 266.

LINOGNATHUS TIBIALIS (Piaget) (2, p. 646).

6 2 Q from an unrecorded host.

LInoGNATHUS GAZELLA Mjoéberg (4, p. 157).

@ 2 from an unrecorded host.

LINOGNATHUS PITHODES, sp. n. (Text-figs. 3-5 )

23 gd and 12 Q 9 from the Indian Antelope Antelope cervi-
capra Linn. — Lucas described a variety of L. tibialis trom the
same host (Ann. Soc. Ent. France, 1847, p. 534).

The species about to be described may at once be distinguished
from all other members of the genus at present known by the
presence on the abdomen of the male of a segmental series of
tergites, each possessing a row of closely ‘placed spines or
“thorns,” short and very thick, and set in a perfectly straight
alignment (see text-fig. 3). The trefoil-shaped appearance of
the posterior end of the abdomen in the male is also an obvious
character for the species.

The genus Linognathus has hitherto been in part defined by
the absence of abdominal sclerites; but, as in other respects the
new species is typically linognathoid, it is better to expand the
diagnosis of the old genus than to create a new one.

Katernal Form, Maty.—A stout insect with a large tun-
shaped abdomen. Head: Short and broad, almost as broad as
long, extending only a little beyond the antenne. In front the
head is truncate and broad, the lateral angles rounded. Mouth
opens ventrally in the middle of a circle of broad-banded chitin,
the anterior semicircle of which runs across the dorsal surface
of the truncate front of the head and then down on each
side, the posterior semicircle being composed of thinner chitin.
The post-antennal region of the head is a little broader than the

FROM THE SOCIETY'S GARDENS. 26]

pre-antennal, and the two margins are parallel to one another.
Antenne: Elongate, graduated in width from the base to the
tip (see measurements, p. 266). Cephalic apodemes as usual in
Linognathus absent, the head fitting into a deep V-shaped cleft

Text-figure 3.

At : :
i GP ;

P / A o—a i O Dy owe
P >| ye ee \a <
‘ Bw SS Bag, ; . :
ol Naat tala EW a\ Na
ft) a rrr |, eo :
) i \| 'f \ | if TS i = | Ke,

I AMAANGES Smee

| ‘ | hy Shit rte eo,
wi

i
WW
\ 2\

sera ee

=

!

vi

Wee K

| |

Linognathus pithodes, 8. X 68.

of the prothorax. Thorax: Broader than the head. Legs
powerful, particularly the coxe. The first pair of coxe are large
enough to reach forward on each side nearly as far as the

262 MR. B. F. CUMMINGS ON LICE

antenne, The cox on the same side are contiguous, but widely
separated from those of the other side. Femora short and very
convex in the postaxial margin. Abdomen: Broad, globose.
The tip of the copulatory apparatus projects from the tip of a
long, stout, genital papilla shaped like a finger and terminating
the abdomen. It lies between two smaller processes which
project only a little, and are formed by the pleurites of the last
segment. Dorsally there are five transverse strips of light brown
chitin forming the tergite on each of segments 8 to 7, the anterior
ones most clearly delimited.

Text-figure 4.

i=
“Ls = s

a

—
— Se a
a) a

T=

au S SS &
SS ass —) 4

{ TERZT I

Linognathus pithodes, 2. Ventral surface, showing gonopods. X 46°5.

External Form. FrmaitE.—Abdomen: <A little longer and
narrower than in the male. Tergites absent. As in the female
of L. tibialis, L. fahrenholzi and others, the end of the abdomen
is drawn out into a pair of processes, which in this species are
quite short and project only a little beyond the terminal tergum.

FROM THE SOCIETY'S GARDENS. 263

The gonopods measure °2 of a millimetre in length and -09 wide,
each with a strong chitinous external margin (text-fig. 4).

Cheetotaxy. Mate.—Remarkable for the presence along the
abdominal tergites of a segmental series of perfectly straight
transverse rows of closely placed spines—short, thick, and dark
brown in colour. Head: Several well-spaced hairs on the
chitinous circle around the mouth. Dorsal surface: hairs
arranged as in the text-figure. Ventral surface: a stout hair
on each side at about the level of the postaxial margin of the
antenna. Thorax: A stout bristle on each side of the pronotum.
The usual mesothoracic bristle; between it and the spiracle on
each side three small spiny hairs. Two spines on the hind
margin of the metanotum. Abdomen: The rows of modified
Short spines in straight alignment on the dorsal surface are
given in the following table :—

SISMAINGII I Us We aesand srabieenenes 2
wo Piri ccs SSeS trek NO 2
5

DAWA all ite |) OU ee
8

ba Toe oN Bas UR { :
Sle ROR en ees ie
18

Pa nolo eas [1s
ice SR a, tte Meare 16

7)

On the tergum of segment 8 are two bristles, while across
the tergum of the terminal segment is a row rather difficult to
observe of eight or nine small hairs which connect up with two
small groups of hairs, one on each pleurite. On the pleure of
each segment, dorsally, numerous scattered lanceolate spines. A
pair of small hairs at the tip of the genital papilla. Other
hairs as in text-figure 3. On the ventral surface the spines are
mostly lanceolate in shape, more widely spaced and fewer in
number, arranged in six rows. In the first and second there
are four; in the third, eight; in the fourth, nine (the two
middle ones being longest); in the fifth, twelve (the four middle
ones being longest); in the sixth, ten (the four middle ones being
longest and the lateral ones irregularly arranged). On the
sternum of segment 8 there are two long bristles and a lanceolate
spine on the inside of each pleurumn.

Chetotaxy. Kumaue.—Head: As in the male; but the hairs
on the dorsal surface are stouter and longer. Thorax: As in
the male. Abdomen: Both dorsal and ventral surfaces covered
with a large number of stout lanceolate spines arranged in no
definite pattern or order. On the tergum of the eighth segment,
somewhat apart from the others, are two extra long hairs. On
the eighth sternum, in the middle, just before the gonopods, are
four elongate bristles in two pairs. Between the two couples is

264. MR. B. F. CUMMINGS ON LICE

a patch of minute hairs, ten in number. Tips of gonopods with
several elongate bristles and a patch (situated chiefly on the:
dorsal surface of the gonopod) of smallish thin hairs. Pleura
of segments 3 and 4 with a single long bristle each. Those of
segments 5, 6, 7, and 8 each with two.

Text-figure 5.

ha

Ps.P

Linognathus pithodes. Male copulatory apparatus. X 178.
The parameres have been pulled apart.

BP. Basal plate; Hnd. endomere; P. penis; Paz. paramere;
Ps.P. pseudo-penis.

Coloration.—The thicker parts of the exoskeleton are of a
warm brown colour, as, for example, the band on the anterior:

FROM THE SOCIETY’S GARDENS, 265

margin of the head, the “ thumbs” and claws of the second and
third pairs of legs, and also at the extremity of the abdomen
in the bays, one on each side of the base of the finger-shaped
genital papilla in the male. The tergites are of a light brown
colour, and each antennal segment is similarly banded. Across
the distal broad part of the tibie of the front legs is a com-
paratively narrow, transverse, brown band. Coxe of a uniform
deep brown colour.

Mate Copulatory Apparatus (text-fig. 5).— Basal plate : Broader
than the plate in the two preceding species of Linognathus. At
the base it is slightly bifid, each short limb being a process for
the articulation of a paramere. A suture is observable in the
median line of the plate, extending from the cleft between the
two basal processes up to just bey ond half the length of the plate.
This groove or suture, which indicates the originally double
nature of the plate probably throughout the Order, les in a
narrow thickening in the mid-line of the plate, on each side of
which the chitin is flattened and thinner. At the upper end the
plate is of a uniform thinness and isa little broader. Parameres:
These are long, narrow, and tapering to an acute apex. Towards
the base each paramere develops on its inner side a kind of
supporting ledge or shelf which runs in below the mesosome.
The outer side becomes thickened and turns in almost at right
angles on arriving at the basal plate, with the limb of which
on each side of the cleft it is connected. Penis: Although it is.
impossible to speak definitely when the apparatus is in the
retracted state, I think it is fairly safe to assume that the loop-
like apertures marked P in text-figs. 1, 2 & 5 are homologous.
Now, in L. limnotragi this aperture obviously lies at the tip of
the penis, whereas in the present new form it is seen lying
within the chitinous bars marked Hnd. It may therefore
be supposed that the distal end of the penis-tube with the
aperture is here telescoped into the firm chitinous base, which
may be either endomeral or the base of the penis. The pseudo-
penis (Ps.P.) is also probably an endomeral chitinisation.

Measurements (millimetre scale).

3. 1 oe
| é | |p Heres Bh we
Length. Breadth. | Length. Breadth.
IBUSGE Ges emeamenee 29 20 35 | 17
(median) (median line) |
ANINORERS peonos cae0n0 *22 "30 H “30 | 35
| (lateral margin) (at base)
Abdomen .. ......| 80 64 | "85 65
|
|

266 MR. B. F. CUMMINGS ON LICE

Antenna of male.

Length. Breadth.

Segment 1 ... ........| "064. “080
oe Crate 076 056 |
USS Maree -050 040 |

Pe eePe hued a ata -040 040

‘038

es “044 Gatiese)

(at tip)

otal eeccs: 274

Comparison with the Male Copulatory Apparatus in
Linognathus from Hland.
Subjoined, with a ‘discussion, is a description of the male
copulatory apparatus of a new variety of Linognathus tibialis

Text-figure 6.

Linognathus tibialis. Male copulatory apparatus. X 140.
BP. Basal plate; H. endomere; P. paramere; Ps.P. pseudo-penis.

FROM THE SOCIETY’S GARDENS. 267

from an Eland in the Zoological Gardens at Edinburgh. It is
interesting to include a description and figure of the Eland
Linognathus here for the purpose of comparison.

Basal plate: This closely resembles the plate of L. pithodes in
the bifid lower end, the median suture, and median raised strip.
Parameres: Proximally broad and fitting around the mesosome
closely, a feature in which they resemble those of L. limnotragi
(which wrap in around the mesosome extensively) and L. pithodes,
in which there is a broad leaf-like shelf (see p. 264). They
resemble Z. cavice-capensis in the possession on the inner concave
surface about halfway down of an area of thickened chitin,
longer than the so-called “nodule” in the Procavian Linognathus.
‘The appearance suggests, in both instances, a strengthening of
the middle part for clasping the mesosome. MJesosome: As in
L. pithodes, a pseudo-penis is present and fused with the rest of
the endomeral chitinisation (text-fig. 6). A thin ledge of thin
chitin runs along the outside of each of the two parallel bars.
Between them lies the aperture and what I interpret as the
true penis.

Genus Hysopuruirus Enderlein.

HyYBOPHTHIRUS NOTOPHALLUS (Neumann) (8, p. 44, and 9).
(Text-figs. 7, 8.)

Eggs, larve, and adults, male and female, from Orycteropus

afer (Pall.).

Text-figure 7.

Hybophthirus notophallus. Egg.
C. Cement; Z. lid. X 36.

Both Enderlein and Neumann (see 8, p. 44) have described
this isolated louse-form; but Neumann's description is much
the fuller, and includes a figure and description of the male

268 MR. B. F. CUMMINGS ON LICE

copulatory apparatus. The Society’s specimens agree in toto with
those described by Neumann. In Enderlein’s figure there is a.
discrepancy in the metanotum which I do not understand.

I have re-examined the male copulatory apparatus and the
following description supplements Neumann’s and brings the
parts under recently devised terminology. Basal plate: Fairly
long and broad, broader in front than behind. The broader
anterior end is somewhat spatulate, the concave side facing

Text-figure 8.

eC ! \ 2
TERZI J \ i} \ |
b T r ay \\ ]

Hybophthirus notophallus. Larva, Stage I. X 39.

dorsally. Parameres : Strong, fairly straight rods. They bulge
a little and then immediately narrow before the tip. The tip is.
short and turned outwards. Mesosome: The parts between the
parameres are merged into a single structure, in which the
following parts may be distinguished :—Two bars, one on each
side (probably the endomeres), and apically a triangular plate—
the pseudo-penis. On the dorsal surface, projecting from about.
the level of the distal ends of the endomeres, is a small median

FROM THE SOCIETY'S GARDENS. 269

chitinous papilla, which is probably the penis. The larger distal
end of this papilla is supported on each side by a chitinous
margin—these latter parts, lower down, running into one another
and forming a narrow neck.

The egg is shown in outline in text-fig. 7. The egg is large
and fastened to the hair by a very strong thick cement along a
straight inner surface, equal almost to two-thirds its total length.
The shell and the lid are quite smooth. The micropyle apparatus
on the lid consists of a number of very minute canals, which do
not, aS in many Anoplura, project as papillz on the lid. Length
15 mm.; greatest width -72 mm.

Larva, Stage I. (text-fig. 8).—Head is very rounded. Abdomen
without sclerites, ovate, smooth margin all the way round.
Thorax with straight hind margin. Almost bald, except for
two mesonotal hairs and two long hairs on each side of the last
segment of abdomen. Other minute hairs as shown in the text-
figure. Described from a single specimen.

Measurements (millimetre scale).

Length. Breadth.

JEISEIGUE oh, Bcgrte yor, Sian aes 10) 38

(behind antenne)

SVOTAU eee eer oe 40 “72,
(at base)

JNOGKORTNEN 3, one aoe nay oo. 1:20 | 1:02

Rotaleeen 2°60
Length of antenna ......... BS} wulial,
MALLOPHAGA.

Genus BoopiA Piaget.

Boorra TARSATA Piaget (2, p. 599). (Text-fig. 9.)

Twenty-two specimens from Phascolomys mitchelli Owen.
Piaget’s specimens were taken on a P. fossor.

In all the males the copulatory apparatus was unfortunately
retracted. It is practically impossible to draw out these parts in
a dead specimen, and in our present state of knowledge very
unsafe to describe such a complex structure as they present so
long as they are withdrawn within the abdomen.

Mouth-parts.—A brief reference to the esophageal sclerite in
Boopia is made in my paper in the P. Z.S8. for 1913, p. 138. The
following description confirms the statement there made and
extends it, all parts of the mouth being passed in review.

The text-figure includes the antenna (text-fig. 9). . Between

270 MR. B. F. CUMMINGS ON LICE

the so-called paraglosse the front margin of the labiwm is a little
concave in the middle and somewhat swollen on each side at.
the base of each paraglossa. Each swollen patch carries several
small spiny hairs. irst maxille: For the palpi, see text-
figure. The lobes are twice as long as broad. The inner surface
densely studded with small curved hooks, those at the distal
end longest. Mandibles: The right mandible is roughly quadri-
lateral, the two distal lateral angles being formed by two distinct

Text-figure 9.

Sy

Ss

PC

Boopia tarsata. Mouth-parts. XX 300.

OG. Ciliated groove; FP. fringed plate; IC. median cornu; MP. main plate
of lyriform organ; Mayp..maxillary palpus; P. paraglossa; PC. post. cornu;
PB. post. band; R. right mandible. X. Antenna.

apices—-one acuminate, the other broad and rounded—separated
from one another by a bay. The lefé is less powerful, with two
acuminate apices—one alongside the other, and one a little
longer than the other. Jsopogometric apparatus: The text-
figure shows the peculiarly complex framework on which this.
apparatus is held together. The main plate of the pharyngeal
sclerite bears a curious resemblance to the sternum of such a

FROM THE SOCIETY'S GARDENS. Daal

Ratite bird as Dinornis or Apteryx. The posterior lateral pieces.
are produced some way backwards and join on each side a trans-
verse band. From each end of the transverse band a posterior
cornu coils upwards on each side of the pharynx. The plate also.
has a posterior median process and two anterior lateral processes.
The latter run forward and each gives articulation to a chitinous
strip which bends round at the mouth and there forms the double
fringed plate. Forwards, from the pharyngeal plate between the
two anterior processes, two strong narrow cords run, forming
between them the “‘ciliated groove,” but diverging and even-
tually curling around posterior to the fringed plate on each side,
so as to turn back into a broad, somewhat indefinite sheet of
rather thin opaque chitin set with teeth.

Genus TricHopEctrses Nitzsch.

TRICHODECTES BREVICEPS Rudow (11).
8 92 from Lama glama lL. (Family Camelide). Rudow’s

descriptions are notoriously bad, and are, as a rule, insufticient
even for identification purposes. I prefer, however, to refer
these female specimens to his species for the present, rather than
describe them as new.

TRICHODECTES PARUMPILOSUS Piaget (2, p. 397).

Two immature specimens from Cervus «xanthopygus M.-
Kdwards, I refer very doubtfully to this species.

TRICHODECTES LATUS Nitzsch (10, p. 53). (Text-fig. 10.)

Twelve specimens, including males, from Canis latrans Linn.

Male Copulatory Apparatus (text-fig. 10)— Basal plate: The out
line is shown in the figure. The lateral margins are strongly
developed rods, the main body of the plate between being concave,
so that the whole plate is of a trough-like form. At its anterior
end the plate is rounded and the chitin becomes opaque and the
outline indefinite. Parameres: At the base these are broad and
leaflike, folding around the mesosome so as to form a sheath.
The margins of the parameres almost meet each other over the
dorsal surface at the base and similarly over the ventral surface..
The sac is shot out between the parameres and is covered with
denticles. One-half of the distance from its extremity is a
transverse row of about eighteen, triangular denticles, larger
than the rest, and functioning probably as retinacula. On the
dorsal surface the base of this sac is strengthened by a flat
lanceolate splint of chitin, with a broader base (at the lower:
margin of the basal plate) and a much narrower distal end
towards the tips of the parameres. When the apparatus is
retracted, this lanceolate band at about halfway is bent down-
wards and back upon itself, so that in side view it appears as an
enigmatic loop. Between the two lateral margins, in the clear

272, MR, B. F. CUMMINGS ON LICE

space at the base of the plate, may be seen a small forked rod,
the fork pointing forwards. I am unable to say definitely
whether this is the penis-rod attached to the end of the extru-
sible sac, or whether it is merely a median chitinisation of the
basal plate.

Text-figure 10.

Trichodectes latus. Male copulatory apparatus. > 200.

BP. Basal plate; FR. forked rod; ZB. lanceolate band (endomeral) ;
P. paramere; PS. preputial sac.

TRICHODECTES CRASSUS Nitzsch (10, p. 53).

Eighteen specimens from the Common Badger (J/eéles meles
Linn.).
The male copulatory apparatus of this species agrees very

FROM THE SOCIETY'S GARDENS. ike,

closely with that of the preceding species. 7. pinguis N. also
presents a close resemblance in the male genitalia to 7’. latus.
In a revision and splitting up of this large and unwieldy genus,
these are facts which should be borne in mind. (Concerning this
subject, see also p. 283.)

TRICHODECTES CORNUTUS Gervais (12).

One female and two larve from Gazella euchore, now: known as
Antidorcas euchore Zimm. Gervais’s specimens were collected on
Antilope dorcas. 'Taschenberg (18; p. 220) identifies the species
with Rudow’s 7. longiceps (11, p. 110), taken on A. arabica.
Neumann (14, p. 626) records it from Hippotragus equinus.
But, as Piaget remarks, the species requires to be examined
again and described with more care.

TRICHODECTES HEMITRAGI, sp. n. (Text-figs. 11, 12.)

The material on which the following description is based
consists of 13 females from the Tahr (Hemitragus jemiaicus
Ham. Smith).

This new parasite is of considerable interest, on account of
certain features in the anatomy of the mouth-parts, which are
figured and detailed below. It is sufficient here to say that the
pharyngeal sclerite or lyriform organ, upon superficial inspection
apparently absent, proves on dissection to be present, but in so
highly modified a form, that it must be considered unique in the
Trichodectide so far examined, while it diverges greatly from
the typical form of the organ in the Mallophaga as a whole
(compare text-fig. 12 with text-fig. 16). This is the more
interesting, as 7’. hemitragi 9 is unmistakably a Trichodect, and
presents, with this exception, no particularly novel characters.
‘The male, however, is yet to be discovered, and may prove to
rank as a distinet genus.

In examples of Docophorus bisignatus, from the Storks and
Tbises, | have pointed out an instance (20, p. 134), very similar
to the present one, of an abrupt deviation in the form of the
esophageal sclerite from that of the rest of the Mallophaga,
where, particularly in Docophorus and Trichodectes and in the
Ischnocera generally, it presents a fairly uniform appearance.

It would be premature to discuss the reason for this funda-
mental change in the character of this organ, occurring so
abruptly among forms not otherwise anomalous, until our know-
ledge of the function of the isopogometric apparatus is more
exact and fuller. But it is, in any event, a very remarkable fact.

It may be pointed out that the pharyngeal sclerite is an
internal organ, and in both the cases mentioned above is
invisible without dissection. In these instances, therefore, a
purely superficial. diagnosis could only have resulted in the
omission of an important and deep-rooted morphological differ-
ence—a character which, if external, would probably entitle the

Proc. Zoou. Soc.—1916, Nc. X VILL. 18

274 MR, B. F, CUMMINGS ON LICE

species to generic rank, in the opinion of most of the systematic
workers in this Order.

The following is a description :—

External Form (text-fig. 11). Famare—Head: Preantennal
area quite short. No frontal sinus, front margin straight or

Text-figure 11.

Trichodectes hemitragi, @. X 55°5.

very slightly concave, with a narrow marginal band. Inferior:
‘“hair-canal’’ absent or obsolete. Temples very rounded and .
swell outwards behind the eye, making the head a little broader
behind the antenne than in front. Dorsally, two occipital

FROM THE SOCIETY'S GARDENS. 275

bands as rafters of the skull run forward towards the mandibles,
and are straight and parallel to one another. In front of each
antenna 1s a fairly large incrassation, rounded in form and con-
_ nected by a narrower neck with the margin. Thorav: Narrower
_than the head, with straight metanotal margin and convex lateral
margin. Abdomen: Broader than the head, elliptical. A brown
band on the dorsum of the segments, those on 5, 6 and 7 being
the deepest in colour and in length. Gonopods (see text-fig. 11).

Chetotaxy. Frmate.—Head: Dorsal surface covered with a

great many fairly short hairs. Antenne also set with a great
many hairs, long and short, including one long one preaxially
and a row of four long ones dorso-postaxially in segment 2, and
on segment 3, along the postaxial margin, a straight row of six
fairly long hairs. Ventrally, postantennal area appears to be
quite bare. On the preantennal area there are numerous long
bristles between the antenna and the frontal margin. Thorax:
Dorsal surface set with small bristles, arranged as shown in the
figure. Abdomen: Dorsal surface covered with bristles, long
and short, which it is possible roughly to analyse into three
transverse rows on each segment. There is, however, a small
bare area inside each pleurite. Each pleurite carries numerous
small hairs and two long ones, which are particularly long in
segments 6 and 7. Dorsum of last segment almost bare, except
for four or five long hairs in a widely-spaced transverse row.
‘Ventral surface thickly covered with hairs, there being a par-’
ticularly dense patch between the gonopods. At the extreme
end of the abdomen there are two long bristles dorsally and two
ventrally.

Mouth-parts.— Althoughthe hair-canal in front of the mandibles
may be said to be absent, it is indicated on the ventral surface
by a difference in the thickness of the chitin of the margin.
Mandibles are large and strongly ridged, the right one in almost
its whole breadth lying behind the left, far forward near the
front margin of the head. On the right one are three distinct
apices, the middle one being the longest. At the base it runs in
as a stout quadrangular process (text-fig. 12). Opposite this and
on the dorsal surface is a large knob of dense chitin. The left
mandible has three apices, two of which are very small and close
together, and the usual narrow basal process. The transverse
ridges are particularly strong, prominent, and downwardly
directed at the base of the ventral tooth. Ridges are continuous
in both mandibles across the surface of the mandible. First
Macxille: These do not call for particular remark. Labiwm :
Front margin straight with a small, short, squat paraglosse
at each lateral angle. Jsopogometric Apparatus: 'The pharyngeal
sclerite or lyriform organ is a slender and delicate piece of
chitin, consisting of two large sprawling posterior cornua, a

‘prominent median cornu between these, a smal] and insignificant
‘“nucleus” or main body, and two anterior cornua rather broad

and long. The chitinous chord or duct, as usual, runs forward
ilfsh=

276 MR. B. F. CUMMINGS ON LICE

and bifureates, each branch entering a “gland” or basal piece.
The latter has a short posterior tendon attached to it.

Text-figure 12.

G
D
AC
a LO
MC
PC

Trichodectes hemitragi. Mouth-parts. 166. Labium not shown.
L. Left mandible. R. Right mandible. X 120.

BP. Basal process; D. duct; G. “gland”; K. knob; ZO. lyriform organ ;
PC., MC.. AC. posterior, median, and anterior cornua; QP. quadrangular
process; 1, 2, 3. apices.

Measurements (millimetre scale), 2.

|

| |

| Length. | Breadth.
Head a. cheers) 50 52

| (behind antennee)
UMNO Eb Ghetecca nse astinsa snl 28 eh)

|
NDA OMEN esac e| 1:10 "84,

Totaliywee el 1:88

Length of Antenna :—

Segment 1 ............... 070
AN a dei A ah a BERIT
aan iy Oe 120

iO taller 290

TRICHODECTES HARRISONI, sp. n. (Text-figs. 13-16.)

Several specimens, male and female, collected on the White-
tailed Gnu (Connochetes gnu Zimm.). In the shape of the
abdomen of the male, more particularly of the last segment
which is produced and has a deep median bay, this new form
recalls Damalinia and 7’. forficula. There is, however, no frontal
sinus, the head being semicircular in front, and I consider its
nearest allies, therefore, to be 7. forficula P. (from Cervus

FROM THE SOCIETY’S. GARDENS. OT

porcinus) and 7. climax N. (from Capra hircus), more especially
the former.

The species is named after Mr. Launcelot Harrison, B.Sc., of
the University of Sydney.

Text-figure 13.

Trichodectes harrisoni, 6. XX about 55.

a. Tip of antenna, much enlarged.

External. Form. Maue (text-fig. 13).—Head: Front margin
semicircular. Marginal band narrow at the sides, broader across.
the front. Frontal sinus absent. Antenne arise about midway,

278 MR. B. F. CUMMINGS ON LICE

one on each side. Postantennal area as broad as the base of the
preantennal semicircle. Temples rounded. Occipital margin
broad; the two dorsal rafters of the skull are parallel and
widely separated. There are also two ventral rafters pursuing
the same course. At the occiput the dorsal and the ventral
rafter of each side are united one to the other by a concave
broad band forming the sides of the occipital hole. Forwards
the dorsal rafters become evanescent in the chitin of the roof,
just about the level of the mandibles. Each ventral rafter in
front splits into two branches, the outer curling around into the
posterior part of the antennary socket and the inner one appa-
rently becoming absorbed in the thick chitin, which gives the
mandibles articulation. Antenna: First segment large. swollen,
all three segments about equal in length. Hair-canal absent or,
at any rate, very shallow; two somewhat convergent chitinous
bands run from the clypeus to the front margin of the head and
indicate the sides of the canal. As in 7’. climax and others
the frontal marginal band is thickened, with a narrow, median,
longitudinal, white cleft. In a greatly developed hair-canal,
such as 7’. subrostratus N. possesses, there is no band crossing
the frontal sinus, the lateral marginal bands, one on each side,
running down the sides of the hair-canal. Thorax: Narrower
than the head. ‘here are two distinct parts—pro-+ mesothorax
and metathorax or prothorax and meso+metathorax. The latter
is a little the broader. Lateral margins of both are rounded.
Coxee of first pair of legs lie close to one another in the centre.
First pair of legs short, the tibiz of the second and third pairs
remarkably long. Abdomen: Tapers elegantly to the anal
extremity. The tip is bifid. The material at my disposal is
insufficient to determine the precise morphology of the bifid
tip. In Damalinia, according to Mjoberg, it is the produced
sternite of the last segment. There are two other species of
Trichodectes with bifid tips to the abdomen, viz., 7’. forficula P.
and 7’. appendiculatus P., but I have not been able to examine
either of these for comparison. A tergite on each of the first
three segments; but each tergite gives a suggestion of being
double, and in the following three segments each tergite is
plainly divided in half by a transverse light-coloured band.
It is possible that these divisions are only colour-differences.
On the penultimate segment, a small tergite. The termination
of the abdomen is of a clear whitish chitin. A single sternite in
each segment. Hach sternite, even in the posterior segment, is
an integral whole, so that a genital plate may be said to be
absent, although the last two sternites are, on each side,
bracketed together by a lateral band of brown chitin.

External Form. Frmaue (text-fig. 14).—The usual sexual
differences in the antenne. Abdomen: Ovate. A single tergite
and sternite on each segment, dark brown in colour. Tergite 1
fits the whole space between the pleurites. In the tergites that
follow, there is a clear space laterally between tergite and

FROM THE SOCIETY'S GARDENS. 279

pleurite. The gonopods are fairly broad, ear-like plates pro-
jecting beyond the end of the abdomen. They cover the sternal
surface and lie transversely across the end of the abdomen, their
concave surfaces uppermost, adpressed against the sternal abdo-
minal surface.

Text-figure 14.

BOZAw ET
= )
LA

Wes =

S77 Ii. Wwe
Z Val \ Calan \*

; yh re Ss

, <a ¥
L A ipgemaneniil! (OSS

rH a Ti ) ; \
[XA Af WTAUAN AAO TAUAANANINUAU ANI AY) a)
@) i WATT rT i "Ny N\

]

° @ ay fe)
Gil aa iN J

‘fo

Hy\

Trichodectes harrisoni, 2. XX about 56.

Chetotaxy. Mauts,—Head: A great many scattered hairs over
ppreantennal area dorsally, and also over dorsal surface of the
temples. Median dorsal postantennal area, between the two
longitudinal bands, bare except for a transverse row of short hairs.
Ventrally, numerous short hairs in preantennal area on each side

280 MR. B. F. CUMMINGS ON LICE ”

-of the hair-canal; postantennal area apparently bare. - Thoram:
‘Small scattered hairs on pronotum and a row along posterior
margin. A transverse row of short hairs on metanotum. Other
hairs as in the figure. Abdomen: A transverse row of quite short,
closely-placed hairs along the posterior margin of each tergite ; in
those segments where two tergites are present, the row runs across
the front margin of the second. A small semicircle of short hairs.

Text-figure 15.

Trichodectes harrisoni. Male copulatory apparatus. X 118°5.

BP. Basal plate; H. endomere; P. paramere; Pen. penis rod; PS. preputial sac..

on anterior lip of genital opening. The whole of the dorsal surface
of the apex of the abdomen studded with short hairs. Ventrally,
a row of hairs along posterior margin of each sternite. Fewer
hairs on ventral surface of the end of the abdomen than on the-
dorsal. Other hairs as in the figure. .

“U

FROM THE SOCIETY'S GARDENS. 28}

~ Chetotaxy. Femaue.—As in the male, except in the region of

the genital opening. Here, a fringe of hairs runs around the
margin of each gonopod, and there is a patch of short ones
between the gonopods at the base.

Male Copulatory Apparatus.—This resembles that of 7’. forficula
according to Piaget’s rather obscure figure. Basal plate: Consists.
of two straight parallel-sided marginal bands, parallel to one
another, with a transparent and apparently membranous median
strip between. At its base for the articulation of the paramere
each band is obliquely truncate. Parameres: Much shorter than
the endomeres. Hach paramere in the mid-part of its length isa
narrow rod; towards the base its inner margin runs out to form
an inner trochanter, as broad as the band of the basal plate to
which it is attached. Distally, the paramere broadens dorso-
ventrally and forms a concave flange, the concave surface on the

Text-figure 16.

Trichodectes harrisoni. Male mouth-parts (maxillz not shown). X 200.
L. Left mandible. R. Right mandible. X 168.

G. Gland; K. knob; ZO. lyriform organ; P. paraglossa; 1, 2, 3. apices.

inside. Hndomeres: Large pieces, the tips reaching almost to
the genital opening. They are fused into one piece at the base
and form a single forked sclerite, concave dorsally, so as to hold
the sac. Hach limb of the fork is quite broad, but has an acute
apex and carries a small nodular tooth subapically. The sac is
long, covered with small denticles. Penis and telomeres are
developed on the distal end, but are very delicate and trans-
parent, and hard to make out (text-fig. 15).

Mouth-parts (text-fig. 16).—Mandibles: These resemble those
of 7. hemitragi, than which, however, they are relatively much
smaller and less powerful. When in repose, moreover, only the
distal ends overlap, that of the right being a little behind and
above (on the inside of) the left. In each there are three apices ;
on the right there is the same stout quadrangular process, with

i
282 MR. B. F. CUMMINGS ON LICE

the large knob opposite. On the left there is the usual sharp
basal process (present in the left mandible of most Mallophaga).
On both mandibles are transverse ridges fewer than in 7’. hemi-
tragi. First Maxille: Small lobes that call for no particular
remark. JLahbiwm: “ Paraglosse ” short and columnar, rounded
at the tips, firm outer surface. Five inwardly directed spines on
each distal end. No lobes, labial margin straight. Ten minute
hairs, set in large alveoli on the labium, and further back two
short hairs, one on each side. Lsopogometric Apparatus: No
posterior cornua on the lyriform organ. Anterior cornua broad,
and almost as long as the “ nucleus.”

Measurements (millimetre scale).

Length. Breadth.
3 P- 3. 2.
FELGAG). sssancaseocis 5 MA coil ‘56 (in| “Bd
front of antennz)
TORE so5 suo one non 32 22 “40 “4.4.
| (metathorax) (metathorax)
Abdomen ......... 1°35 | 1:20 “75 y
| (segment 3) | (segment 3)
ay ae ais | 22, 2
Total ...... | 212 193i
Length of Antenna. é | 2)
|
Segment 1 ...... TSO os! 076
A ese 135 | 012
deacane) 110 012
Motalleea: 395 | 10
| Length of Legs, 3. | Ist. 2nd. 3rd.
JENSTONON EN a Woes daa boo ues 016 180 "02
Tibia + tarsus ...... 200 “345 028
Claw (curved) ......... 060 120 iy
a SS
Total ...... 276 645 —

FROM THE SOCIETY’S GARDENS. 283

TRICHODECTES Ovis Linn. s

Specimens from Ovis musimon Linn. Also been recorded from
O. aries, O. ornata, O. melanocephala.

TRICHODECTES sp.

12 2 2 from Capreolus capreolus Linn. The specimens
belong to the much confused tibialis-group of Trichodectes from
Deer. It is thoroughly unsafe to identify specimens of this
series from females until the confusion, caused chiefly by the
absence of morphological evidence in previous descriptions and
figures, has been dispelled. These females certainly are not
T. tabralis, but agree most with some unnamed Trichodects from
Reed-Buck (Africa), shown me by Mr. Waterston among some
material belonging to the Imperial Bureau of Entomology.

Genus Eurricnopuitus Mjoberg.

EUTRICHOPHILUS SETOSUS (Gieb.) (10, p. 56). (Text-fig. 17.)

37 2 2 from Frithizon dorsatum Linn.

I have been able to make a preparation of the male copulatory
apparatus from a male contained in a tube of this species presented
to the British Museum by the Hon. N. C. Rothschild. In many
species of Trichodectide, males are rare and in JZ’. scalaris N.
unknown.

The Family Trichodectidze consists at present of but three
genera—T'richodectes, Damalinia, and Hutrichophilus. Before
Damalinia and Eutrichophilus were split otf in 1910 by Mjoberg,
the old genus Zrichodectes was simply a miscellany, which still
requires breaking up into genera—a by no means easy task, on
account of the difficulty in finding convincing characters. In
view of a future revision of the family, attention is drawn to
the probable value of the male genitalia systematically. I find,
for example, that in Hutrichophilus setosus and in FL. coéndu
Stobbe (15) the male genitalia are of quite the same type. This
is described below. Another type, perfectly distinct, is formed
by Trichodectes latus, 7’. crassus, 7’. pinguis, and probably by
others (see p. 271). Still another type may be seen in the
male genitalia of 7’richodectes gastrodes Cummings (16, p. 99),
T. mephitidis Osborn (17, p. 242), 7’. geomydis Osborn (18, p. 54),
and 1’. interrupto-fasciatus Keil. & Ferris (19, p. 61), which agree
in the fusion of the parameres at their distal ends and in the bifid
form of the endomeres (see 16, text-fig. 4; and 19, pl. vil. fig. 2,
pl. viii. figs. 4 & 6).

Male Copulatory Apparatus (text-fig. 17).— Basal plate : Broad
anteriorly narrowing gradually to the posterior end, where the
plate is constricted into a narrow “ waist,” to which the endo-
meres and parameres are attached. The lateral margins are
marrow, rod-like, the posterior third broader. The anterior part
of the plate, as is frequently the case, is thinned out, composed
of delicate chitin with an almost invisible anterior margin.

284 MR. B. F. CUMMINGS ON LICE

Parameres: Of the appendages at the base. of the plate, I take:
the outside ones to be endomeral and the two elongate inner
ones parameres. The homologies of the other parts figured I do-
not indicate, pending further dissections.

«

Text-figure 17.

Butrichophilus setosus. Male copulatory apparatus. X 182.

BP. Basal plate; #. endomerial chitinisations; P. parameres;
?. problematical parts.

Comparison with the Male Copulatory Apparatus of
E. coéndu Stobe.

This is of exactly the same type. Basal plate; Much broader
in relation to its length than that of ZH. setosus, and much
broader at the base. Between the two lateral margins at the
base lies the same little chitinous piece as in the preceding
species, only it is larger and stretches right across the plate,
almost from one margin to the other, narrowing at each end;
in the middle, pointing upwards and backwards from the hind
margin, is a small sharp-pointed process. The two inner appen-
dages are narrow, and twice as long as the outer ones. These,
probably the endomeres, are very short, broad, stunted pieces,.
articulating with the basal plate.

FROM THE SOCIETY'S .GARDENS. 285

Genus Lamosorurium Nitzsch.

LZMOBOTHRIUM TITAN Piaget (2, p. 578).
62 9, 3 larve from an Accipitrine bird. Host’s name not
given.
Genus Gontocores Burm.

GoNIOcoTES MICROTHORAX Nitzsch (10, p. 184).
1 3,1 @ from the Common Partridge (Perdix perdi Linn.).

Text-figure 18.

Par.

Goniocotes gigas. Male copulatory'apparatus. X 87.

BP. Basal plate; H. endomeral chitinisation; Par. paramere.

‘The parameres possess sensory hairs in large alveoli. Compare with text-fig, 23.

286 MR. B, F. CUMMINGS ON LICE

GONIOCOTES VERRUCOSUS Taschenberg (18, p. 94).

I have ventured to identify with Taschenberg’s species a single
male from Orypturus noctivagus (Wied), The species was de-
seribed by Taschenberg from a single male taken on Crypturus
variegatus (Gmel.).

GONIOCOTES sp.
1 larva from Querquedula flavirostris ( Vieill.).

GoNIocorEs eras Piaget (2, p. 238): (Text-fig. 18.)

1 ¢, 19, and an immature form from Crossoptilon mant-
churicum Swinhoe.

The text-figure of the male copulatory apparatus is drawn
from a specimen in a tube full of this species, presented to the
British Museum by the Hon, N. C. Rothschild.

Male Copulatory Apparatus.—Basal plate: Very long and
narrow ; parallel-sided. Anterior half uniform brown. Posterior
half with brown lateral margins and a pale median area.
Parameres: Relatively short, spear-shaped.

Genus StronecyLocotes Taschenberg.

STRONGYLOCOTES CoNICEPS Tasch. (18, p. 63

1 ¢ from Crypturus noctivagus (Wied).
This species is known from a single male specimen taken on
Crypturus variegatus (Gmel.).

Goniodide from Tinamous.

The collection contains Goniodide from Vothoprocta cinerascens
(Burm.), Rhynchotus rufescens (Temm.), and Orypturus noctivagus
(Wied), belonging to four or five different species, but in each
instance the material, consisting of but one or two specimens,
is insufficient in a confused group such as this to make their
determination satisfactory. For the time being, therefore, I am
reserving these specimens until the time is ripe for a much-
needed revision of Tinamou Goniodide.

Genus Gontopes Nitzsch.

GonIobEs cotcHicus Denny (21).

1 2 from Phasianus versicolor (Vieill.) and 1 imperfect
specimen from P. colchicus.

GonroDEs DisPARr Nitzsch (10, p. 193).
13,1 @ from the Common Partridge (Perdia perdia Linn.).

GonIoDES MINOR Piaget (2, p. 256).
136,22 9,and 1 larva from Leptoptila reichenbachi Pelzeln.

FROM THE SOCIETY'S GARDENS, ° 287

GONIODES MEGACEROS Kell. & Paine (22).

1 3 from Lophophorus refulgens Temm,

The species is based on a single male from the same host.
Concerning the copulatory apparatus, the authors, in passing,
note that it is “‘ prominent with heavily chitinised rods reaching
to the second abdominal segment.” In their figure (pl. xv.
fig. 8) the genitalia are shown as seen through the integument,
but with the parameres in front and the basal plate behind!
The specimen prepared probably had its genitalia exserted and
coiled over its back with parameres pointing backwards, and in
the course of being mounted the apparatus became pressed down
upon the abdomen in the reversed position.

GONIODES FALCICORNIS Nitzsch (10, p. 198). (Text-fig. 19.)

Several specimens from Pavo cristatus var. nigripennis Sclater.
Mouth-parts (text-fig. 19).— Mandibles ; The left (dorsal surface)
is roughly of the shape of an equilateral triangle; at the exterior

Text-figure 19.

Goniodes falcicornis. Mouth-parts. X 113°5.
L. Left mandible. R. Right mandible.

BP. Basal process; C. condyle; G. gland; ZO. lyriform organ; NV. chitinous
nodule; Pav. paraglossa; QP. quadrangular process ; Sel. sclerite.

basal angle is a small cup-shaped socket where the mandible arti-
culates with the head ; at the inner basal angle is the usual tooth-
like process which is here in the form of a long narrow stylet,
curved at the base, and carrying at the tip preaxially a small
recurved tooth, Theapex of the triangle is produced, and carries
two distinct cutting-edges with separate tips. On the ventral
surface the mandible is more of an isosceles triangle in shape, the
base-line being shorter and about midway along its length, running
out into a large rounded condyle of dense chitin. In the right

mandible the basal process as usual is quadrangular, the large.

288 ' MR. B. F, CUMMINGS ON LICE

middle condyle is the same as in the right, and the socket at the
external angle is somewhat deeper. Two very distinct apices
are present, situated laterally one to the other, the longer one
separated from the shorter by a considerable space. Ridges in
both mandibles absent. irst Maxille: These lobes call for no
special mention. Labiwm: ‘‘ Paraglosse” as usual; anterior
margin is concave in the middle with a small convex swelling
on each side carrying four or five spines. On the dorsal surface
of the labium, 7. e. within the oral opening, is visible a sclerite
of the same shape and in the same position as that described in
Trichodectes gastrodes (16, p. 99). It consists of a transverse
band with two limbs at each extremity, one anterior and one
posterior. The posterior one in this case is very long, and runs
back on the inside of the under surface of the labium almost as
far as the hind end of the “gland” or basal piece. The anterior
one is noteworthy, as it runs forward a short way and then near
the base of the paraglossa runs intoa dark brown chitinous nodule,
visible on the other side of the labium. Shipley (24) regards this
nodule described in G. tetraonis as a-labial appendage. Wso-
phageal sclerite: The text-figure shows the form of this organ in
this species.

Male Copulatory Apparatus.—M joberg (4, p. 249, text-fig. 142)
has published a drawing of this apparatus which is so inaccurate
as to require, some time in the future, to be earefully refigured.
The morphology of the parts is more or less clear, there being
basal plate, parameres, endomeres, and penis, but the features
which make this relatively enormous apparatus so remarkable
are the strange and complex forms which the several parts have
assumed,

GoNIODES BICUSPIDATUS Piaget (2, p. 278). (Text-figs. 20-22.)

Several specimens from Z’ragopan caboti (Gould).

A tube of the same species from Ceriornis satyrus, presented
to the Museum by Lord Rothschild, contained several larve of
two stages, the description of which is included below.

Male Copulatory Apparatus (text-fig. 20).—The following
description is drawn up from a specimen preserved in copula
with the female. The parts, therefore, were exserted and their
exact relation easily made out. Sasal plate: Long and fairly
broad. Anterior end thin and colourless. Dark brown marginal
bands along posterior half. Parameres: Rather like a rabbit’s
ear in outline. The distal end is produced into a very narrow,
slender, needle-like apex. The sac is covered with small teeth,
especially at the distal end, but I have been able to discover no
true penis. At the base of the sac the endomeral chitinisations
consist of two parts, a dorsal and a ventral. The dorsal is a
stout, lanceolate, median piece, quite separate from the sac, and
probably functioning asa penis. The ventral is a small, thin,
rectangular plate, lymg between the two lateral bands at the
base of the basal plate, and giving support to the sac. From each

FROM THE SOCIETY'S GARDENS. 289

of the two anterior lateral angles it sends off a long supporting
process down the membranous sac. When functioning, the para-
meres stand off at right angles to the sides of the basal plate and
serve to anchor the apparatus within the female’s genital cavity.

Text-figure 20.

Goniodes bicuspidatus. Male copulatory apparatus. X 60.

BP. Basal plate; End. endomere; Par. paramere ; Prp.S. preputial sac ;
Ps.P. pseudo-penis.

Larve.—The larvee, I believe, were all those which, ultimately,

with one exception (see text-fig. 22, a) would have matured into

Proc. Zoou. Soc.—1916, No. XIX. 19

290 MR. B. F. CUMMINGS ON LICE

females, as in all the specimens the head had assumed, even in
Stage I., the definitive female form. Both the head and the
thorax, not only in form but in chetotaxy, are practically
identical with the adult female, even in the earliest stage in the
collection, which will probably prove to be Stage I. I have also
made one or two dissections of the larval mouth-parts and
can discover no characters in which they differ materially from
those of the adults. The larval abdomen, however, requires
separate treatment.

Text-figure Dale

/

es Pw \leseSss

\

|

Goniodes bicuspidatus. Larva. StageI. X 49.

Stage I. (text-fig. 21). Haxternal Porm.—Abdomen : Small with
an even margin, devoid of all sclerites, Chetotaxy : The arrange-
ment of bristles cannot be safely described from one specimen,
and that given in the figure must be regarded as approximate

FROM THE SOCIELY’S GARDENS. 291

only. The abdomen bears a great many bristles, all relatively
very long and powerful. ’ :

Stage LI. (text-fig. 22). Huternal Porm.—Abdomen: Pleurites
of a simple form developed. As in the adult the pair immedi-
ately behind the metanotum are very large and run in over the

Text-figure 22.

Goniodes bicuspidatus. Larva. Stage II. (or ILI. ?). X 33.

a. Antenna ofjanother larva, probably male, showing a slight
enlargement of segment 3.

dorsum a considerable way. But the rest are simple quadri-
lateral plates lying dorsally on the lateral margin. The small
Leyes

292 MR. B. F. CUMMINGS ON LICE

spiracle lies on the inner margin of each pleurite, about halfway
along its length. Tergites are present in the form of circular or
oval plates, two on each segment, and each tergite is just on the
inside of, but separated by a space from, the pleurite. In the
adult this space is filled in, and the tergite and pleurite unite.

Chetotaxy.—Abdomen : This differs but little from that of the
adult female. But there are apparently some interesting differ-
ences from Stage I. (compare the metanotum in text-figs. 21 &
22). There are fewer hairs in the mid-dorsal region of each
segment, and the chetotaxy around the end of the abdomen and
the genital opening differs considerably, of course, from the adults,
in which the sexual organs are matured.

Genus Ruopaxoceras Taschenberg.

RHOPALOCERAS STYLIFER Nitzsch (10, p. 200). (Text-figs. 23,
24.)

1g. Host not given; probably from Meleagris gallopavo
Linn.

Male Copulatory Apparatus (text-fig. 23).—For the purposes of
- the following description I have been able to make use of speci-
mens of this common Turkey parasite, kindly placed at my
disposal by the Rev. Jas. Waterston, B.Se.

The apparatus in this remarkable-looking insect presents
features of great interest, Inasmuch as it is reduced to a very
small size (although in no sense atrophied), in correspondence no
doubt with the fact that the terminal segments of the abdomen
have become modified so as to take part in the function of
copulation. Similar adaptation of the end of the abdomen to
the function of copulation is not rare in Mallophaga. Basal
plate: Long and narrow, margins slightly thicker than the
median area; the anterior end a little broader and thinner.
Rest of the Apparatus: This is so highly modified that the
attempt here made to bring it into line with the parts in other
Mallophaga is only partial and tentative. Parameres as such are
absent. Instead, articulating with the basal plate is a small
trowel-shaped plate with the concave side uppermost. Along
the lateral margins (dorsal surface) of the distal half of this
plate there are on each side seven minute directive hairs with
large alveoli. Contained within the hollow formed by the
trowel-shaped plate, and coiling upwards and backwards so as
to resemble a crook, lies the penis (or a pseudo-penis?). This
crook-shaped piece at its base is set in a stout horseshoe-shaped
piece of dense chitin, the two arms of the horseshoe being
directed towards the basal plate. The parts are perhaps a
modification of those of G. gigas (see text-fig. 18), the parameres
having coalesced. (Cf. 4, fig. 143.)

Terminal Segments of the Abdomen of the Male (text-fig. 24).
Dorsal surface: The 7th tergite is a narrow transverse band which,

FROM THE SOCIETY’S GARDENS. 293

unlike the preceding tergites, runs right across the dorsal surface
from side to side. This is sueceeded by a large shield-like plate
of chitin forming the fused tergites of the 8th and 9th segments.
There is a deep median bay in its posterior margin, into which

Text- figure 23.

Rhopaloceras stylifer. Male copulatory apparatus. 185.
BP. Basal plate; Par. paramere (fused); Ps.P. pseudo-penis (A. side view).

is closely fitted the broad bases of the terminal stylets. Ventral
surface: The 7th sternite is broad and long and, unlike the
preceding sternite, runs right across the sternum from side
to side. Jn front, the anterior part encroaches somewhat upon

294 MR. B. F. CUMMINGS ON LICE

the sternal area of the 6th segment. Immediately in front of
the base of the appendage, which is hinged on to the 8th
sternite, the 7th shows a noticeable quadrilateral development
of its middle part which carries two bristles. The long, narrow,
finger-shaped appendage is attached in the mid-line of the 8th
and extends to the end of the abdomen, where it curves up
between the two terminal stylets, which are excavated somewhat
on their inner surfaces to allow the appendage to pass and {to

Text-figure 24.

> (
Za

ee

———
ZA

Goniodes stylifer, 6. Terminal segments of the abdomen. Ventral view. X 586

A, Appendage; St. terminal stylet; VIZ., VIII. segments.

project on the dorsal surface. This appendage on its dorsal
surface is concave, convex below. The dorsal channel is formed
by the bending over of the free lateral margins, which meet in
the middle but can be separated by a needle and bent back and
flattened ott as a plate. At its base the chitin is pinched up
to form a hinge. On each side of the middle line the sternite
is developed into a triangular piece, with the apex pointing
inwards. The 9th and 10th sternites apparently not developed.

FROM THE SOCIETY'S GARDENS. 295,

Literature.

(1) mi ao Eist-lns: vole vin pai, ple 1. fis. 6
1778).
(2) Prager, E.—Les Pédiculines. Leide, 1880.
(3) Neumann, L. G.—Arch. de Parasitologie, 1911.
(4) Msopere, E.—Arkiv for Zoologi, Stockholm, Bd. 6, No. 13,
1910.
(5) Fanrennoiz, H.—Jahresber. Naturh. gesell. Hannover,
(niedersiichs., zool. Ver.), 2-4, (1910-12) 1912.
(6) Cummines, B. F.—Proc. Zool. Soc. 1915, p. 245.
(7) BURMEISTER, H.—Gen. Rhynchota, No. 8, 1838.
(8) Cummines, B. F.—Bull. Ent. Res. iv., May 1913.
(9) Neumann, L. G.—Jahrbiicher des Nassauischen Vereins fiir
Naturkunde in Wiesbaden, 62 Jahrg. 1909, p. 2.
(10) GirpeL, C. G.—Insecta Epizoa, Leipzig, 1874.
(11) Rupow, F.—Zeit. f. ges. Naturw. vol. xxvii. 1866, p. 110.
(12) Gervais.—Hist. Ins. Aptéres, vol. iii. p. 315, 1847.
(13) TascuenBerc, O.—Nov. Act. Acad. Caes.-Leop. Carol.,
Bd. 44, No. 1, 1882.
(14) Neumann, L. G.—Arch. de Parasitologie, xv. 1913.
(15) SropsE, R.—Deut. ent. Zeit., Heft v. p. 566, 1913.
(16) Cummines, B. F.—Ann. & Mag. Nat. Hist. 1916, Jan.
(17) Ossorn, H.-—Bull. 5, n.s., Div. of Entomology, U.S. Dept.
of Agricul., Washington, 1896.
(18) Osporn, H.—/bid., Bull. 7, 1891.
(19) Kennoce, V. L., & Ferrris.—Leland Stanford Junior
University Pub., Univ. Ser. 1915.
(20) Cummines, B. F.—Proc. Zool. Soc. 1913, p. 128.
(21) Denny, H.—Monographia Anoplurorum Britanniz, London,
1842, p. 158.
(22) Kexioce, V. L., & Parne.—Records of Indian Museum,
vol. x. pt. 4, No. 12, Aug. 1914, p. 227.
(23) SurpLey, A. E.— Proc. Zool. Soe. 1909, p. 314.

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THE SECRETARY ON ADDITIONS TO THE MENAGERIE. (297

EXHIBITIONS AND NOTICES.
February 8, 1916.

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

The Secrerary read the following report on the Additions
to the Society’s Menagerie during ‘the months of November,
December, and January : ==

NOVEMBER.

The number of registered additions to the Society’s Menagerie
during the month of November was 53. Of these 32 were
acquired by presentation, 16 were received on deposit, 3 in
exchange, | by purchase, and 1 was born in the Gardens.

The number of departures during the same period, by death
and removals, was 150.

Amongst the additions special attention may be directed
iO) =

1 Leopard cub (Felis pardus), from Accra, presented by
Hugh M. Willoughby on November 12th.

1 Caracal (Felis caracal) and 1 Fettered Cat (/. ocreatus),
from Berbera, presented by Dr. R. E. Drake-Brockman, F.Z.8.,
on November 30th.

3 Lund’s Opossums (Didelphys albiventris) and 1 Wied’s
Opossum (D. aurita), from Minas Geraes, both new to the
Collection, presented by Prof. J. P. Hill, F.R.S., F.Z.S., on
November 6th.

1 Yellow-rumped Tanager (Rhamphocelus icteronotus), from
Kceuador, new to the Collection, presented by Alfred Ezra, F.Z.5.,
on N ovember 13th.

DECEMBER.

The number of registered additions to the Society's Menagerie
during the month of December was 107. Of these 68 were
acquired by presentation, 37 were received on deposit, and 2 in-
exchange.

The number of departures during the same period, by death
and removals, was 144.

Amongst the additions special attention may be directed
0) =

1 Kyra Cat (Felis eyra), 1 Salt-Desert Cat (7. salinarum),
and 1 Allamand’s Grison (Grison allamandt), the last two new
to the Collection, from Cordova in the Argentine, presented by
W. A. Smithers, C.M.Z.8., on December 16th.

2 Mongolian Sousliks (Citellus mongolicus) and 2 Sand-Ham-
sters (Cricetulus griseus), the latter new to the Collection, and

298 MR. OLDFIELD THOMAS ON A

3 Great Hagle Owls (Lubo bubo), from Mongolia, presented by
A. L. Hall on December 10th.

JANUARY.

The number of registered additions to the Society’s Menagerie
during the month of January was 57. Of these 43 were acquired
by presentation, 8 were received on deposit, 3 in exchange, and
3 were born in the Gardens.

The number of departures during the same period, by death
and removals, was 123.

Amongst the additions special attention may be directed
to :=—

1 Anoa (Anow depressicornis) 2 from Celebes, and 1 Pere
David’s Deer (Hlaphurus davidianus) 3 from Northern China,
presented by H.G. The Duke of Bedford, K.G., Pres.Z.S., on
January 20th and 26th.

2 Argentine Frogs (Leptodactylus mystacinus) and 6 South-
American Sand-Toads (Lufo arenarun) from Cordova, Argentina,
new to the Collection, presented by Wilfred A. Smithers,
C.M.Z.S., on January 31st.

Mr. R. E. Hoxprxe exhibited the skull of a Roebuck, showing
an unusual deviation in the direction of the suture of the right
frontal bone, which extended considerably beyond the median
line towards the left.

Mr. C. Tare Reean, M.A., F.Z.S8., exhibited, by means of
lantern-slides, a series of drawings of larval Fishes from the
Antarctic.

The development of Myctophum antarcticum was compared
with that of the northern J. glaciale, and larval Nototheniidee
were described.

A new Sable Antelope from Angola*.

Mr. OuprietD Tuomas, F.R.S., F.Z.S., exhibited the scalp and
frontlet, with horns, of a male Sable Antelope from the Luando
River, Angola, which had been presented to the National
Museum, together with a female mask and horns, by Mr. H.
F. Varian.

This magnificent animal differed widely from the ordinary
Sable, both by its immensely finer horns, and by the characters

* [The complete account of the new subspecies described in this communication
appears here; but since the name and a preliminary diagnosis were published in the
‘ Abstract,’ No. 151, 1916, it is distinguished by the name being underlined.—
Eprror.]

(or)
al

NEW SABLE ANTELOPE FROM ANGOLA.

(unripe abr snp.

yoddifp) alopayuy a[qeg urposuy Jo pray

"T One Y-9xo J,

300 ON A NEW SABLE ANTELOPE FROM ANGOLA.

of its face-marking, and Mr. Thomas proposed to distinguish it as
a new subspecies under the name of

HiprorraGus * NIGER VARIANI 7.

Thomas, Abstract P. Z.8. 1916, p. 1 (Feb. 15th).

The horns of the type measured 57 inches (1445 mm.) in
length along the front curve, by 11 inches (280 mm.) in cireum-
ference at the base, and 27 inches from tip to tip. Their trans-
verse ridges, which were extremely well marked, were 47 in
number. Good Rhodesian and Nyasa Sable horns were about
45 to 50 inches in length, while those of the East African Sable
were not known to reach 40 inches. The female horns of
H. n. variant were 35 inches (890 mm.) in length by 7 inches
(179 mm.) in circumference.

A skull of this form, which had been lent to Mr. Thomas for
comparison, measured 480 mm. in length, 170 mm. in breadth,
with an upper tooth-row of 122 mm., these dimensions in a skull
of the ordinary Sable being respectively 435, 160, and 114 mm.

But, apart from its splendid horns, the most striking character
of H. 2. variant was the practically complete obliteration of the
usual prominent white streaks running from the anteorbital
white tufts forwards to the sides of the muzzle, the whole of the
upper side of the face being therefore deep black, with the
exception of the anteorbital tufts themselves, which were white
as usual, Along the ordinary positions of the white streaks a
few lighter hairs were perceptible, these being rather more
numerous in the female.

The dark parts of the head were of the deepest black, the
light parts buffy whitish or cream-colour, except the middle line
of the interramia, which was white. Occiput mixed black and
ferruginous-tawny. Hars, as usual, rich tawny outside, and
white within. The face of the female was blackish brown, and
the crown and occiput tawny.

It was with considerable hesitation that Mr. Thomas had only
distinguished this Sable as a subspecies, and not as a species, so
striking was the difference from ordinary Sables in both horns
and marking; but the presence of light hairs along the usual
pesition of the facial streaks, and the fact that in H.n. kirku
(figured by Matschie as H. n. kauf/mannz), the nearest neighbour
of H. n. variant, the dark stripes were broader and the light
stripes narrower than in H. m. niger, showed that these mark-
ings were variable and plastic, and did not indicate any really

* This generic name was used provisionally pending the decision of the authori-
ties as to the names suggested in 1914 for tixation by Fiat. Should Hippotragus
be rejected, the technical name of the genus would be still in doubt until the
question of the validity of Hgocerus, Desm. 1822, nee Aegoceros, Pall. 1811, was
settled, a very knotty point. A law covering this latter case had been proposed by
the Linnean Society’s Committee on Nomenclature in 1906, and submitted to the
International Congress, who, however, only accepted it as applying to specific
names, a restriction much to be regretted.

+ Type. Face, skin, and frontlet with horns. B.M, 16.2.21.1.

ON THE ANTLERS OF A VIRGINIAN DEER. 301

essential difference, such as to render unlikely the possibility
that intermediates might yet be found.

To this subspecies there presumably belonged the well-known
61-inch horn in the Florence Museum, which had long been a
wonder to all sportsmen, who had only had for comparison with
it the relatively short horns of //. 1. niger, those of the E. African
H. n. roosevelti being still shorter.

Bocage’s Hippotragus niger, in his papers on Angolan Mam-
mals, was of course also H. n. variant, but his only material was
a single pair of horns, 51 inches in length, brought home from
the “interior of Mossamedes ” by Welwitsch.

Mr. Varian had taken great pains to secure specimens of this
animal, and to obtain information about its range, and it was
with much pleasure that Mr. Thomas had named it in his honour.
Mr. Varian had also taken steps to induce the local authorities
to give it protection, which, in view of the considerable sums
given for such horns as it carried, would be much needed to save
it from extermination, now that its existence had become known
to sportsmen and hunters.

Judging by the greater length of the skull, it would, no doubt,
prove that H. n. variant not only carried longer horns, but was
larger in all dimensions than the true Sable. It was hoped
that a complete specimen of this splendid addition to the list
of African Antelopes would soon be obtained for the National
Museum, whose warmest thanks were already due to Mr. Varian
for the donation of the tine trophy now exhibited.

Antlers of a Virginian Deer affected by Cancer.

Mr. R. I. Pocock, F.R.S., F.Z.8., Curator of Mammals,
exhibited the successive Antlers of a Virginian Deer (Odocoileus
americanus) that had died of cancer in the Society’s Gardens, and
made the following remarks :—

“The male Virginian Deer (Odocoileus americanus) referred to
in Prof. Plimmer’s report (see p. 83, 16) as having died of cancer,
was purchased as a young animal on Jan. 17th, 1911, and died
Dee. 27th, 1915. The following accurate records ‘of its successive
antlers were kept during the five years that it lived in the
Gardens :—

1. The first antlers, shed March 15th, 1912, were simple snags
about 4 inch in length.

2. The second antlers, shed March 23rd, 1913, measured
1014 inches along outer curve, were simply forked at the
tip; the supernumerary tine on the back of the beam
measured just under 14 inch, and the two antlers together
weighed 5+ oz.

3. The third antlers, shed March 24th, 1914, measured
9 inches, were simple, carrying no supplementary tine.
The two together weighed 47 oz.

302 ON A SIAMESE FIGHTING-FISH AND A COLOMBIAN CAT-FISH.

4. The fourth antlers, shed March 15th, 1915, measured
7? inches; the left antler was simple, the right was
foaleerl the supplementary tine measuring 1} inches.
The ae antlers together weighed 32 oz.

5. The fifth antlers, veal ourmamnetshed. were taken off the head of

i the dead animal on Dec. 27th, 1915. The longer of the
two measured 83 inches. The right was simple, the left
forked, the supplementary tine measuring ? inch. The
two together weighed 33 oz.

Thus, although there was a great and, so far as I am aware,
normal increase in size of the second antlers as compared with
the first, the third, fourth, and fifth antlers showed no corre-
sponding elaboration, but, on the contrary, degeneration, the
third being shorter rod lighter than the second, and producing
no tines. The fourth algo were lighter and shomer than the
third. Nevertheless, the right one produced a supplementary
tine which, however, was shorter than the supplementary tines
of the second pair. The fifth antlers in the matter of length
showed a slight recovery as compared with the fourth, and the
two together acquired the same weight, but the supplementary
tine was still shorter, and the dried integument adhered to the
antlers instead of peeling off and leaving them normally bur-
nished. Since, in Prof. Plimmer’s opinion, the growth of the
cancer from which this Stag died was probably a gradual process
extending over a few years, it seems justifiable to infer, in the
absence of any other obvious cause to account for the fact, that
degeneration of the antlers was attributable to this disease. It
may be added that the testicles, which Prof. Plimmer particularly
examined at my request, were unaffected by the cancer, and were
normal except for the absence of ripe spermatozoa.”

February 22, 1916.

Dr. A. Surru Woopwarp, F.R.S., Vice-President,

in the Chair.

The Rev. H. N. Hurcuinson, M.A., F.Z.S., exhibited a number
of drawings prepared by Mr. T. W. Par tt of restorations of
various extinct animals.

Mr. C. Tare Reean, M.A., F.Z.S., gave a lantern-exhibition
illustrating the breeding-habits of a Siamese Fighting- Fish (Betta
splendens Regan) and the climbing-habits of a Cat-fish (Arges

marmoratus Regan) from the Andes of Colombia.

ON THE TYMPANIC BULLA IN HYZNAS. 303

The Tympanie Bulla in Hyenas.

Mr. R. I. Pococg, F.R.S., F.Z.S., Curator of Mammals, gave
an exhibition, illustrated by lantern-slides, to show the presence
of two chambers in the tympanic bulla of the Hyznide, and
remarked :—

“In his paper upon the base of the skull in the Fissipede
Carnivora (P. Z.8. 1869, pp. 4-37), Prof. Flower laid stress
upon the presence or absence of a bony partition dividing the
cavity of the tympanic bulla into two compartments in the
Aluroidea. Although on general grounds he followed Turner *
in classifying the Hyznas with the Felide and Viverride, he
described the bulla of the Hyznas as ‘ perfectly simple within,
without trace of division into compartments’ (p. 26). Subse-
quently, Mivart (P. Z. 5. 1882, p. 199) wrote ‘though there is
no septum, yet I have detected in both species of yena, inside
the auditory bulla, two osseous ridges or lamine, which, if
further developed, would divide off a small anterior chamber
from the much larger and externally more prominent posterior
portion.’ These two papers appear to be the sources whence
subsequent authors, like Weber, Sedgwick, and others, have
derived their information; Weber, following Mivart, described
the partition as low, and Sedgwick, following Flower, recorded it
as absent.

Both Flower and Mivart were quite mistaken ; the bulla in all
Hyeenas is divided by a strong partition into a larger outer or
anterior and a smaller inner or posterior chamber.

It may be recalled that in the Felide and Viverride the
septum rises from the floor of the bulla and typically extends
upwards till it touches the periotic (petrous) bone. This par-
tition may arise just below the lower rim of the external auditory
meatus, or it may arise far away from that point. In the former
case the antero-external chamber is small, in the latter it is large
as compared with the postero-internal or posterior chamber ; but
the free edge of the partition always reaches, or is situated
close to, the same portion of the periotic, namely, the portion
which is pierced by the fenestra rotunda of the inner ear, and it
is always just at this point that there is a passage or orifice
between the two chambers.

The outer chamber is itself partially divided from the external
auditory meatus by a horseshoe-shaped ridge or crest, the
tympanic ring, which is well shown in Flower’s figure of the
section of the bulla of the Tiger (text-fig. 1, B, t7.).

When the bulla of the inverted skull of the Hyena is opened

* P. Z. 8. 1848, pp. 68-88. Flower’s paper is little more than an amplification of
this valuable paper by Turner, so far as the Carnivora are concerned. It does not
seem, however, that Turner was acquainted with this partitior, his mention of the
division of the bulla into two parts referring to the superficial groove marking the
position of the partition.

304 MR. R. I. POCOCK ON THE

anywhere between its anterior extremity and the paroccipital
process, it presents the appearances which misled Flower and

Text-figure 1.

(Copied from Flower’s figures, P. Z. 8. 1869, pp. 16-17.)

A. Right half of the base of the skull of the Tiger (Felis tigris) with the bulla laid
open to expose the inner chamber (?.c.), with the septum or partition (s.)
ascending to the periotic (pev.), and the orifice (o.) leading from the inner to
the outer chamber between the septum and the periotic; e., internal orifice
of eustachian tube; ov., foramen ovale; jfp., foramen lacerum posticum ;
/po., paroccipital process ; 7., mastoid ; eam., external auditory meatus.

B. Section of the auditory bulla of the Tiger. ic., inner chamber; oc., outer
chamber with the orifice (0.) between the two and the septum (s.) dividing
them; ¢., half the tympanic ring in the outer chamber; eam., external
auditory meatus ; per., periotic.

Mivart. The greater part of the space is occupied by a single
large cavity, which opens by a wide cleft in front into a smaller

Description of Text-figure 2 (continued).

C. Left bulla enlarged, with the posterior portion of the septum, marked s in fig. A,
cut away to show the cavity of the inner chamber and the antero-internal
portion of the bulla also cut away, and the carotid canal (c.) laid open ;. the
passage leading from the outer to the inner chamber between | the periotic
(per.) and the septum marked by an arrow. (f., foramen piercing sphenoid
and corresponding with the auterior carotid foramen of Mongooses ; fp., fora-
men lacerum posticum. Other lettering as m figs. A & B.

D. Part of the right side of the skull viewed from the occipital aspect, with the
bulla laid open from behind to show the inner chamber with the periotic
(per.), carrying the fenestra rotunda, partially blocking the orifice between
the two chambers divided by the septum (s.). Other lettering as in fig. A.

E. Right bulla of the Spotted Hyzna (Crocuta crocuta). ; A line drawn between
the paroccipital process (po.) and the stylomastoid foramen (s¢.) would
mark the edge of the partition between the two chambers.

TYMPANIC BULLA IN HYZENAS. 305

Text-figure 2.

D

A. Base of the skull of the Striped Hyena (Hyena hyena) with the left tympanic
bulla opened. 6., right tympanic bulla; c¢., carotid canal, its course shown by
an arrow; é., inner orifice of eustachian tube, the course of which is shown
by an arrow ; eam., external auditory meatus; s., partition or septum between
the two chambers; ¢., tympanic ring; st., stylomastoid foramen ; po., par-
occip:tal process ; m., mastoid; ov., foramen ovale.

B. Anterior portion of left bulla enlarged and viewed obliquely from the inner side
to show that the crest mistaken by Mivart for a low septum is the tympanic
ring (¢r.), with the internal auditory meatus (iam.) ; per., periotic; fm., fora-
men lacerum medium, exaggerated in size.

Proc. Zoou. Soc.—1916, No. XX. 20

306 MR. R. I. POCOCK ON THE

cavity continuous with the eustachian tube and the external
auditory meatus. This cleft deeply notches the floor of the large
cavity, and the floor slopes backwards and upwards from the
cleft, through which a portion of the periotic is visible, to the par-
occipital process. The edges of the cleft are no doubt the ‘two
osseous ridges or laminze which, if further developed, would
divide off a small anterior chamber from the much larger... .
posterior portion,’ described by Mivart. That is true; but the
two chambers would not correspond to the two present in the
Tiger, for the outer of the two lamine is the tympanic ring, and
is therefore not the homologue of the partition dividing the
Tiger’s bulla, as Mivart supposed. The ‘small anterior chamber’
of the bulla, which is exceptionally large in the Hyezenas, is
merely the anterior part of the tympanic chamber.

Whether Flower correctly interpreted this lamina as the
tympanic ring or not, does not appear; but he may be given the
benefit of the doubt. Nevertheless, both he and Mivart failed
to detect that where the apparent floor—or roof, if the skull be
held in its normal position—of the bulla abuts against the
periotic, there is quite a distinct orifice through which a probe
can be passed backwards into a second chamber lying behind and
below the apparent floor of the bulla. This chamber can be laid
open by cutting away the paroccipital bone externally to the
occipital condyle. It will then be seen quite clearly that the
bony plate, regarded by Flower and Mivart as the floor (or roof)
of the bulla, is, in reality, a partition dividing the bulla into two
chambers, and passing from the periphery of the cavity of the
bulla to the periotic, exactly as is the case in the Tiger, allowance
being made for the origin of the partition much farther back
than in that Feline. It is not, however, much farther back than
in some other Aluroids, e. g., Cynictis.

Nevertheless, it is not certain, in my opinion, that the par-
tition in the Hyznas is the exact homologue of that of the Cats.
The inner wall of the posterior chamber of the bulla in Afluroidea
is often strengthened by bony crests or ridges of varying height,
and one such crest, curving round the back of the chamber and
occupying the position of the partition, where it rises from the
bulla near the paroccipital, in Hycna, is present in two immature
skulls of Proteles, in addition to the normal vertical partition
which in these specimens is thin and imperfectly ossified or fene-
strated. The interest of this fact lies in the circumstance that
Proteles in several of its cranial characters occupies a position
midway between Hyena and the Mungotine. Hence it is possible
that in Hyena the normal partition has been replaced by a
secondary partition of stronger. growth. However that may be,
it is quite clear that the bulla of Hyena can no longer be
described as undivided.

Two other points of systematic importance may be alluded to:
the bulla in Hyznas is fused anteriorly to the basisphenoid, as

TYMPANIC BULLA IN HYANAS. 307

in the Cats, and beneath its anterior end, and concealed by it,
there is a foramen piercing the sphenoid and remote from
the foramen lacerum medium. This sphenoidal foramen seems
to correspond exactly with the exposed foramen by which the
internal carotid artery in the Mongooses enters the skull,
after traversing the bulla. In the Hyenas, nevertheless, this
artery is said by Mivart to enter the skull by the foramen
jacerum medium, the existence of a carotid foramen in the
sphenoid being denied by that author. The foramen, never-
theless, persists, as it does in Proteles, whether the carotid enters
it or not.”

March 7, 1916.

The Marquess oF SuiGo, Vice-President,
in the Chair.

Mr. Harry K. Kusrace gave a bioscope exhibition of films
illustrating his experiences as a big-game hunter and cinemato-
grapher in Kast Africa, showing the natives and the characteristic
animals of that country in their natural state.

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abs dhe iti ae if Aa he aN Bai ‘# yg Wis aie

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No. 151.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON,.*
February 8th, 1916.

Prof, E. W. MacBripz, D.Sc., F.R.S., Vice-President,
in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

The Secrerary read a Report on the Additions to the Society’s
Menagerie during the months of November and December, 1915,

and January 1916.

Mr. R. E. Hope exhibited the skull of a Roebuck, showing
an unusual deviation in the direction of the suture of the right
frontal bone, which extended considerably beyond the median

line towards the left.

Mr. Ouprietp Tuomas, F.R.S., F.Z.S8., exhibited the scalp and
frontlet with horns of a male Sable Antelope from the Luando
River, Angola, which had been presented to the National
Museum by Mr. H. F. Varian, together with a female mask and
horns.

This magnificent animal differed widely from the ordinary
Sable both by its far finer horns and by the character of its face-
markings, and Mr. Thomas proposed to distinguish it as a new
subspecies under the name of Hippotragus niger variant.

The horns of the type measured 57 inches in length along the
front curve, by 11 inches in circumference at the base, and had

* This Abstract is published by the Society at its offices, Zoological Gardens,
Regent's Park, N.W., on the Tuesday following the date of Meeting to which
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,
to all Fellows who subscribe to the Publications ; but it may be obtained on the
day of publication at the price of Sixpence, or, if desired, sent post-free for
the sum of Ste Shilings per annum, payable in advance.

2

47 transverse ridges upon them. Good South African and
Nyasa Sable horns were ordinarily about 45 to 50 inches in
length, while those of the East African Sable were not known
to reach 40 inches. The female horns of H. n. variant were
39 inches long.

In colour the face of the new form differed conspicuously by
the almost complete obliteration of the usual prominent white
streaks running from the anteorbital white tufts forwards to the
sides of the muzzle, the whole of the upper side of the face being
therefore deep black, with the exception of the anteorbital tufts
themselves, which were white as usual. In both male and
female a few odd light hairs alone indicated the ordinary position
of the streaks.

The type-specimen would be registered as B.M. No. 16.2.21.1.

Mr. C. Tare Reean, M:A., F.Z.S., exhibited, by means of
lantern-slides, a series of drawings of larval Fishes from the

Antarctic.
The development of Myctophum antarcticwm was compared
with that of the northern J. glaciale and larval Notothentide

were described.

Mr. R. I. Pocock, F.R.S., F.Z.S., Curator of Mammals,
exhibited the successive Antlers of a Virginian Deer (Odocoileus
americanus) that had died of cancer in the Society’s Gardens, and
attributed the degeneration shown to this disease.

Prof. H. G. Puuwer, F.RS., F.Z.S., Pathologist to the Society,
read his Report on the Deaths which occurred in the Society’s
Gardens during 1915, and on the Blood-parasites found during

the same period.

Prof. E. B. Poutron, M.A., F.R.S., F.Z.S., presented a paper
on a collection of Moths made in Somaliland by Mr. W. Feather,
containing descriptions of ten new genera and a large number of
new species by Sir George F, Hampson, Bt., F.Z.8., and others.

The next Meeting of the Society for Scientific Business will
be held on Tuesday, February 22nd, 1916, at half-past Five
o'clock p.M., when the following communications will be made :—

3
The Rev. H. N. Hurcutnson, B.A., F.Z.S.

Exhibition of drawings of Extinct Animals.

R. I. Pocock, F.R.8., F.Z.S.
Exhibition to illustrate the structure of the Tympanic Bulla
in Hyenas.
C. Tate Recan, M.A., F.Z.8.

Lantern exhibition of the nest of a Fighting Fish and the
climbing habits of a Catfish.

Bruce F, Cummines.
Studies on the Anoplura and Mallophaga, being a Report
upon a Collection from the Society’s Gardens.—Part I.
P. CHAatmers Mitcuett, M.A., D.Sc., F.R.S., F.Z.8.

Further Observations on the Intestinal Tract of Mammals.

The following Papers have been received :—

G. A. Boutenesr, F.R.S., F.Z.S.

1. On the Lizards allied to Lacerta muralis, with an Account
of Lacerta agilis and L. parva.

2. On Specimens of the Bolti, Tilapia nilotica, a Teleostean
Fish with increased number of Anal Spines.

Major R. MEINERTZHAGEN, F.Z.S8.

Notes on the Sitatunga or Marsh-Antelope of the Sesse
Islands.

T. Goopry, D.Sc.

Observations on the Cytology of Flagellates and Ameebe
obtained from old stored Soil.

Ropert GuRNeEY, M.A., F.Z.8.

On some fresh-water Entomostraca from Ceylon.

The Publication Committee desire to call the attention of
those who propose to offer Papers to the Society, to the great
increase in the cost of paper and printing. This will render it
necessary for the present that papers should be condensed, and
be limited so far as possible to the description of new results.

4

Communications intended for the Scientific Meetings should
be addressed to

P. CHALMERS MITCHELL,
Secretary.
ZOOLOGICAL Society oF Lonpon,
Recent’s Park, Lonpon, N.W.
february ldth, 1914.

NOH 152:

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

February 22nd, 1916.

Dr. A. Smirax Woopwarp, F.R.S., Vice-President,

in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

The Rev. H. N. Hurcuinson, B.A., F.Z.S., exhibited a number
of drawings prepared by Mr. T. W. Parfitt of restorations of
various extinct animals.

Mr. C. Tare Reean, M.A., F.Z.S., gave a lantern-exhibition
illustrating the breeding-habits of a Siamese Fighting-Fish
(Betta splendens Regan) and the climbing-habits of a Cat-fish
(Arges marmoratus Regan) from the Andes of Colombia.

Mr. R. I. Pococn, F.R.S., F.Z.S., Curator of Mammals, gave
an exhibition, illustrated by lantern-slides, to show the structure
of the tympanic bulla in the Hyznas, and pointed out that both
Flower and Mivart were wrong in stating that the Hyznas
differ from the Felide, Viverride, and Protelidz, in having the
eavity of the bulla undivided by a bony septum. ‘The septum,
which those authors mistook for the roof of the bulla, runs from
the back of the bulla, where it abuts against the paroccipital,
obliquely upwards and forwards to the periotic, and divides the
cavity of the bulla into an anterior larger and a posterior smaller
chamber. The Hyznas thus agree with the other families of
Hluroid Carnivores in having the bulla divided.

* This Abstract is published by the Society at its offices, Zovlogical Gardens,
Regent's Park, N.W., on the Tuesday following the date of Meeting to which
it refers. It will be issued, along with the ‘Proceedings,’ free of extra charge,
to all Fellows who subscribe to the Publications; but it may be obtained on the
day of publication at the price of Sixpence, or, if desired, sent post-free for
the sum of Six Shillings per annum, payable in advance.

6

Mr. Bruce F. Cummrnes read a paper containing the first part
of a report on a collection of Anoplura and Mallophaga obtained
from animals in the Society’s Gardens. He dealt with the
structure and development of the various species, and gave
descriptions of three new forms.

Dr. P. CHALMERS Mircnett, M.A., F.R.S., F.Z.8., Secretary to
the Society, gave an account of his paper entitled “ Further
Observations on the Intestinal Tract of Mammals,” illustrating
his remarks with a large series of lantern-slides.

The next Meeting of the Society for Scientific Business will be
held on Tuesday, March 7th, 1916, at half-past Five o’clock P.M.,
when the following communication will be made :—

CINEMATOGRAPH EXHIBITION.

The Srcrerary has been able to arrange with Mr. Harry K.
Eustace, Big Game Hunter and Cinematographer, to give an
exhibition of his films of African Animals at this Meeting.

The Papers previously announced for reading at this Meeting
are postponed.

The following Papers have been received :—
Re i. Pocock, HIR:S., Halcs:, EeZcs:

Lantern-exhibition to show structure of the Alisphenoid
Canal in some Civets and Hyzenas.

Major R. MrInertzHacEn, F.Z.S.

Notes on the Sitatunga or Marsh-Antelope of the Sesse
Islands.

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

1. On Specimens of the Perciform Fish, Tilapia nilotica,
with Increased Number of Anal Spines.

2. On the Lizards allied to Lacerta muralis, with an Account
of Lacerta agilis and L. parva.

“I

T. Goopry, D.Sc.

Observations on the Cytology of Flagellates and Am«be
obtained from old stored Soil.

Rosert Gurney, M.A., F.Z.8.

On some Fresh-water Entomostraca from Ceylon.

Ref Pocock, HRS: WaU:S., E.Z.S:
On the External Characters of the Mongooses (Mungotide).

The Publication Committee desire to call the attention of
those who propose to offer Papers to the Society, to the great
increase in the cost of paper and printing. This will render it
necessary for the present that papers should be condensed, and
be limited so far as possible to the description of new results.

Communications intended for the Scientific Meetings should
be addressed to

P. CHALMERS MITCHELL,
Secretary.
ZOOLOGICAL Soctety or Lonpon,
RecGent’s Park, Lonpon, N.W.
February 29th, 1916.

ee

No. 153.

ABSTRACT OF THE PROCEEDINGS

ZOOLOGICAL SOCIETY OF LONDON
March 7th, 1916.

The Marquess or Stico, Vice-President,
in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

Mr. Harry K. Eustace gave a bioscope exhibition of his films
illustrating his experiences as a big-game hunter and cinemato-
grapher in East Africa, showing the natives and the characteristic
animals of that country in their natural state.

The next Meeting of the Society for Scientific Business will be
held on Tuesday, March 21st, 1916, at half-past Five o'clock P.m.,
when the following communications will be made :—

* This Abstract is published by the Society at its offices, Zoological Gardens,
Regent’s Park, N.W., on the Tuesday following the date of Meeting to which
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,
to all Fellows who subscribe to the Publications; but it may be obtained on the
day of publication at the price of Sixpence, or, if desired, sent post-free for
the sum of Sia Skillings per annum, payable in advance.

10
J.T. Cunnincuam, M.A., F.Z.S.

Exhibition of skins illustrating results of Mendelian Cross
in Fowls.
IR. I. Pocock, FURS: ELGS., F-Z:s.:

Lantern-exhibition to show structure of the Alisphenoid
Canal in some Civets and Hyzenas.

T. Goopry, D.Sc.
Observations on the Cytology of Flagellates and Amcebe
obtained from old stored Soil.
Major R. MrInertzyaceEn, F.Z.S.

Notes on the Sitatunga or Marsh-Antelope of the Sesse
Islands.

The following papers have been received :—

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

1. On Specimens of the Perciform Fish, Tilapia nilotica,
with Increased Number of Anal Spines.

2. On the Lizards allied to Lacerta muralis.

Rosert Gurney, M.A., F.Z.8.

On some Fresh-water Entomostraca from Ceylon.

R. I. Pococg, F.R.S., F.L.8., F.Z.S.
On the External Characters of the Mongooses (Mungotide).

Dr. H. Murr Evans.
The Poison-Organ of the Sting-Ray (Trygon pastinaca).

Dr. J. C. Nomar

1. Methods of estimating the Size of Fish from the Size of
their Scales.

2. Experimental Determination of the Factors which cause
Patterns to appear conspicuous in Nature.

et

The Publication Committee desire to call the attention of
those who propose to offer Papers. to the Society, to the great
inerease in the cost of paper and printing. This will render it
necessary for the present that papers should be condensed, and
be limited so far as possible to the description of new results.

Communications intended for the Scientific Meetings should
be addressed to

P. CHALMERS MITCHELL,
Secretary.
ZOOLOGICAL SOCIETY OF LONDON,

Recent’s Park, Lonpon, N.W.
March \Ath, 1916.

No. 154.

ABSTRACT OF THE PROCEEDINGS

ZOOLOGICAL SOCIETY OF LONDON.*
: March 21st, 1916.

Dr. 8. F. Harmer, F.R.8., Vice-President,

in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

THE SECRETARY read a Report on the Additions made to the
Society’s Menagerie during the month of February 1916.

Mr. K. T. Newron, F.R.S./F.Z.S., exhibited the pelt and bones
of a Black Hare, for which he was under ohligation to Mr. G. F.
Brooke of Leadenhall Market, who had received it with a large
consignment of Brown Hares from Siberia; but, unfortunately,
the locality was not known. ‘This hare is of small size and with
short rabbit-like ears. The head and back are black excepting
only a small white spot on the forehead ; and towards the sides
there are numerous long hairs with white tips. Lower down
upon the sides the fur becomes tawny and passes into white
underneath. All the feet, but especially the hinder ones, have
light brown hair up the upper parts.

The skull and limb bones show characters agreeing with those
of the hare; but in size the animal was intermediate between
our common hare and the rabbit.

Mr. D. M.S. Watson, F.Z.8., gave an account of some obser-
vations he had made on the habits and life-history of Platypus
and Hechidna.

* This Abstract is published by the Society at its offices, Zoological Gardens,
Regent’s Park, N.W., on the Tuesday following the date of Meeting to which
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,
to all Fellows who subscribe to the Publications ; but it may be obtained on
the day of publication at the price of Sixpence, or, if desired, sent post-free
for the sum of Siv Shillings per annum, payable in adyance.

14

Mr. R. I. Pococng, F.R.S., F.Z.S., Curator of Mammals, gave
an exhibition, illustrated by lantern-slides, to show some points
connected with the alisphenoid canal in the Viverride and
Hyzenide, and demonstrated : (1) that the canal is always present
in Viverricula, its occasional apparent absence in that animal
being due to the closure of its posterior orifice near the foramen
rotundum, which opens into the posterior end of the canal ;
(2) that the canal may be complete in Crocuta, as Cuvier stated,
or may be closed at its posterior end.

Dr. T. Goopry read a paper entitled “Observations on the
Cytology of Flagellates and Amcebe obtained from old stored
Soil.” This paper deals with the eytology and nuclear changes
during division of three species of Flagellates and two species of
Ameebe obtained from soil stored in bottles at the Rothamsted
Laboratory for practically fifty years. One of the Flagellates and
the two Ameebee are new to science.

The next Meeting of the Society for Scientific Business will be
held on Tuesday, April 4th, 1916, at half-past Five o’clock p.M.,
when the following communications will be made :—

D. Seru-Smira, F.Z.S.

Exhibition of a small Intensive Poultry House.

Prof. J. P. Hint, D.Se., F.R.S., F.Z.S.

Exhibition of living Cecilians from South America.

G. A. Boutenesr, F.R.S., F.Z.S.

1. On Specimens of the Perciform Fish, Zilapia nilotica,
with Increased Number of Anal Spines.

2. On the Lizards allied to Lacerta muralis, with an Account
of Lacerta agilis and L. parva.

15

Ropert Gurney, M.A., F.Z.8.

On some Fresh-water Entomostraca from Ceylon.

Major R. MEInertzHacen, F.Z.S.
a SEE Eee

Notes on the Sitatunga or Marsh-Antelope of the Sesse
Islands.

The following Papers have been received :—

R. I. Pocock, F.R.S., FLS., F.Z.8.
On the External Characters of the Mongooses (Mungotide).

Major H. Murr Evans, M.D., R.A.M.C.
The Poison-Organ of the Sting-Ray (7rygon pastinaca).

Dr. J. C. Morrram.

An Experimental Determination of the Factors which cause
Patterns to appear conspicuous in Nature.

The Publication Committee desire to call the attention of
those who propose to offer Papers to the Society, to the great
increase in the cost of paper and printing. This will render it
necessary for the present that papers should be condensed, and
be limited so far as possible to the description of new results.

Communications intended for the Scientific Meetings should
be addressed to
P. CHALMERS MITCHELL,
Secretary.
- ZOOLOGICAL SocrnTY OF Lonpon,
Regent's Park, Lonpon, N.W.
March 28th, 1916.

PAPERS.

he Morphology of the Cyprinodont Fishes of the Subfamily Phallostethine, with
- Descriptions of a new Genus and two new Species. By C. Tarn Ruaan, M.A., F.Z.S.
(Plates LL-IV., and Text-figures 1-15.) .... 2... sees se eee eee e tee eee eee eee

On a Collection of Mammals from the Coast and Islands of South-East Siam. By
©. Boven Kuoss, F.Z.8., F.R.G.S. With an Account of the Fruit-Bats, by Dr. Knup
Anpmrsen, F.Z.8. (Plate I., and Pope faures Tie yb crenata acre riaress ave cath ctevatey atm

3. Report on the Deaths which occurred in the Zoological Gardens during 1915,
together with a List of the Blood-Parasites found during the Year. By H. G.
Puer, F.R.S., F.Z.8., Professor of Compar ative Pathology in the Imperial Sue

of Science and Technology, London, and Pathologist to the Society HiGoe

ag

k A Frog with Serameeically Abnormal Hind Feet. By R. W. Haroup Row, B.S8c.,
be F.LS., F.Z.8., Assistant Lecturer and Demonstrator in Zoology, University of
London, King’s College. (Yext-figure 1.) -. i... ce eee ee creer eee eee ee eens

6. On peolecicn of Moths made in Somaliland by Mr. W. Feather. By Professor E
B. Pouuroy, M.A., F.R.S8., F.Z.S. With Descriptions of New Species by Sir G. F.
Hamrson, Bart., L. B. Provr, J. H. Durrant, and Dr. Karu Jorpay. (Plates

UD eee

6. Further Observations on the Intestinal Tract of Mammals. By P. Cmarmers
~ Mrrcunct, M.A., D.Se., L.D., F.B.S., Secretary to the Society. (Text-figures 1-80.).

7. Studies on the Anoplura and Mallophaga, being a Report upon a Collection from the
Mammals and Birds in the Society's Gardens. —Part I., with a Preface. ve Bruce
F. Cummrnes, British Museum (Natural History). (Text-figures 1-24.) ..

Page

27

77

87

91

LIST OF PLATES.

1916, Parr I. (pp. 1-307).

REGAN: Ply le AS Phallostethus... Be Neostetnusucneenie ieeeenee
Il. Neostethus lankestert ......--..0. Lice Rajah a 1
Tit. A. Neostethus. B. Phallostethus ......-........-
IV. A, B. Phallostethus. C. Neostethus ..........00
Koss : Plo. “Sketch Map of SH Siam. .eec nce seme eter ae veie RRR
2 GPA Se 2
BOwEtON - HE Ir \ Somaliland Hepidoptera js cpiss- cic case ron eee 91

NOTICE.

The ‘ Proceedings’ for the year are issued in four parts, paged consecutively,
so that the complete reference is now P. Z. 8. 1916, p.... The Distribution

is as follows:—
: Part I, issued in March,

ye ee June.
Aes Hui abaamrss September,
Pritiane Viaw tee December.

‘ Proceedings,’ 1915, Part IV. (pp. 541-712), were published on
December 24th, 1915.

The Abstracts of the ‘Proceedings,’ Nos, 151-154, are
contained in this Part.

se Fee

PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

AQVOLOGICAL SOCIETY
OF LONDON.

1916.

PART II.

CONTAINING Paczs 309 To 448, witH 7 PLATES
AND 45 Text-ricures, TrtLepace, INDEX, ETC.

JUNE 1916.

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

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

[Price Twelve Shillings.]

LIST OF CONTENTS.

1916, Parr II. (pp. 309-448).

EXHIBITIONS AND NOTICES.

The Szcrrrary, Report on the Additions to the Society’s Menagerie during the month

ofsMebranry 1916 (0). .e hse chee coke a pate Sh Wana tas A a Boe ee eae oes A
Mr. E.T. Newron, F.R.S., F.Z.S. Exhibition of the skin of a Siberian Black Hare .... 441 a

Mr. D. M. 8. Warson, F.Z.S. Notice of remarks on the habits of Platypus and Echidna. 441 |

Mr. R. I. Pocock, F.R.S., F.L.S8., F.Z.S., Curator of Mammals. Lantern-exhibition to
show the structure of the Alisphenoid Canal in some Civets and Hyxnas. (Text- }
Tt R ich ida IP hia ears ICME e ey baie: An mee ac vich teed es tele eee eee oo ae

Mr. J. T. Cunnincuam, M.A., F.Z.S. Exhibition of skins illustrating results of
Mendelian Cross in Fowls ...... be teen sence teeeee distaoaieoetions seen el see 446

Mr. D. Seru-Smurrn, F.Z.S., Curator of Birds, Exhibition of a small Intensive Poultry-

LOUISE yc: Rese ea cust clos OS riots serene seaeee a eect roto eras tenet SS Dia hee On eS ese AAG

Prof. J. P. Hisu, D.Se., F.R.S., F.Z.S. Exhibition of living Ceeilians from South

WALI ETE Caly See hare cate hcl Epevenels etary kate eats etme ete Hee MS Dogma MCSE emptied: St
Mr. G. A. Boutencrr, F.R.S., F.Z.5 “7° > % © sapc> entitled ‘On the Lizards allied

to Lacerta muralis, with an ac: j Nd Tn eaeh Be SAaoddon dc — 447
The Secrerary. Report on the Acc: Menagerie during the month ats

On Mino NOG: Geng ha6 5 on bo ee EIA OF to od ap bkuOob tot oo cons 447

The Secrerary. Notice of a letter received from Lt.-Col. R. T. Lairmr, D.Se., F.Z.S.,

AL MEGs smrelerencesto bil arzilOsis seas roteycrer ctece lone ovale ierenelale a aceree tren ete teeters eer nae 4473

Mr. C. Tarn Ruean, M.A., F.Z.S. Exhibition of lantern-slides illustrating parental care
Din TTS VO Ra Ce EE a A ete BRACE Pou ane aed un pA ee Tra sO

The Secretary. Report cn the Additions to the Society’s Menagerie during the month
GR PA TALON ya oe is) a ciete tere ere peoetcte ik ane Shel eee Be eres Sramau ol o . 448

Mr. R. H. Burne, M.A., F.Z.S. Exhibition of preparations of connections between the

Swim-bladder and Ear in Fishes ......0..c+.ce ee eeceuccecs pare neuer ze LSS

Correction to Prof. E. B. Pouzron’s paper on Moths from Somaliland ................ 448

Contents continued on page 3 of Wrapper,

2

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1 Sa

ZOOLOGICAL SOCIETY OF LONDON.

Tuts Society was founded in 1826 by Sir Sraurorp Rarruzs,
Mr. J. Sasinz, Mr. N. A. Vieors, and other eminent Naturalists,
for the advancement of Zoology and Animal Physiology, and for the
introduction of new and curious subjects of the Animal Kingdom,

an wasincorporated by Royal Charter in 1829,

Patron.

HIS MAJESTY THE KING,

COUNCIL.
HIS GRACH THE DUKE OF BEDFORD, K.G., F.R.S., President,

Tar Hon. Cacit Barine, M.A. Pror, Ervesr W. MacBrinz,

AuFrrep H. Cocks, Hse., M.A.

Tue Rr. Hon. Tue Hart or
Cromer, P.C.,G.C.B.,G.C.M.G.,
F.R.S., Vice-Presedent.

Cuartes Drummonpn, KEse.,
Treasurer,
Aurrep Ezra, Ese.

Carrain Hues 8. Guapstonn,
M.A.

Sipney Freprric Harmer, Ese.,
MEAG Sco e ChenNeO a Vace-
President. —

Cox. Srr Watrer R. Lawrence,
With, (CiOetled

Str Epuunp Gites Loner, Br.,
Vice-President.

M.A, D.Sc., F.R.S., Vice-
President.

Guy A. K. Marswatn, Esa., D.Sc.

E.G. B. Mrapr-Watpo, Ese.

P. Caatmers Mircnett, Ese.,
Whale IDYSGh, IG ID,. 8 3b Si
Secretary.

ALBERT Pam, Esq.

Tur Kart or Porrsmourn.
OxrprizLp Tuomas, Esa., F.R.S,

AUBYN
M.A.

AntHony H. Wrnerietp, Ese.

Trevor-Barrys, Ese.,

Artnur Surra Woopwarp, Kse.,
LL.D.,F.R.S., Vice-President.

Henry Woopwarp, Esa., LL.D.,
F.R.S., Vice-President.

2

The Society consists of Fellows, and Honorary, Foreign, and
Corresponding Members, elected according to the By-Laws. It
carries out the objects of its foundation by means of its collection
of living animals, by its Library, and by its Scientific Publications.

The Office of the Society, Regent’s Park, N.W., where all com-
munications should be sent, addressed to “‘ The Secretary,” is open
from Ten till Five, except on Saturdays, when it closes at OnE P.M.

The Library, under the superintendence of Mr. Henry G. J. Peavot,
is open daily (except Sunday) from Ten a.m. till Five p.m.; on
Saturdays, Ten a.m. till Two p.m.

The Library is closed from Good Friday to Easter Monday, and
upon all other Bank Holidays. It is also closed annually for
cleaning purposes during the whole month of September.

The Meetings of the Society for General Business are held in the
Meeting Room at the Society's Office on the third Wednesday in
every month of the year, except in September and October, at half-
past Four o’clock p.m.

The Meetings for Scientifie Business are held in the Meeting
Room at the Society’s Office fortnightly on Tuesdays, except in
July, August, September, and December and January, at half-past
Five o’clock p.m.

The Anniversary Meeting is held on the 29th. of April, or the
nearest convenient day, at Four p.m.

The Society’s Gardens are open daily from Nine o’clock until
Sunset. Mr. R. I. Pocock, F.R.S., F.L.S., is the resident Super-
intendent: and Curator of Mammals, Mr. D. Seth-Smith is Curator
of Birds and Inspector of Works, Mr. E. G. Boulenger is Curator
of Reptiles, Prof. H. M. Lefroy is Curator of Insects, and
Professor H. G. Pliimmer, F.R.S., M.R.C.S., is Pathologist. Appli-
cations for anatomical material or facilities for work in the
Prosectorium should be addressed to the Secretary of the Society.

TERMS FOR THE ADMISSION OF FELLOWS.

Frriows pay an Admission Fee of £5, and an Annual Contri-
bution of £3, due on the Ist. of January, and payable in advance,
or a Composition of £45 in leu thereof; the whole payment,
including the Admission Fee, being £50.

No person can become a Frriow until the Admission Fee and
first Annual Subscription have been paid, or the annual payments
have been compounded for.

Fettows elected in November and December are not liable for
the Subscription for the year in which they are elected.

3

PRIVILEGES OF FELLOWS.

Frttows haye Personal Admission to the Gardens upon signing
their names in the book at the entrance gate, and may introduce
Two Companions daily.

The Wire or Houssanp of a Futtow can exercise these privileges
in the absence of the Fellow.

Until further notice, Fxtnows will receive 40 undated Green
Cards, available on any Sunday or week-day up to the end of
February of the year following the year of issue, and 20 White
Cards available on any week-day up to the same date. Twenty
of the Green Cards may be exchanged for a book containing two
Orders for each Sunday in the year. Twenty White Cards may
be exchanged for a book of dated Week-day Orders, each Order
available for any day during the week except Sunday. Special
children’s tickets are no longer issued, but the Green and White
Cards are perforated, and each half is valid for a Child under twelve
years of age. It is particularly requested that Fellows will sign
every ticket before it goes out of their possession. Unsigned tickets
are not available.

Frttows are not allowed to pass in friends on their written
order or on presentation of their visiting ecards.

Frttows have the privilege of receiving the Society’s ordinary
Publications issued during the year upon payment of the additional
Subscription of One Guinea. This Subscription is due upon the
1st. of January, and must be paid before the day of the Anniversary
Meeting, after which the privilege lapses. FxEttows are likewise
entitled to purchase these Publications at 25 per cent. less than
the price charged to the public. <A further reduction of 25 per
cent. is also made upon all purchases of Publications issued prior
to 1881, if above the value of Five Pounds.

Fettows also have the privilege of subscribing to the Annual
Volume of ‘ The Zoological Record, which gives a list of the Works
and Publications relating to Zoology in each year, for the sum of
One Pound Ten Shillings. Separate divisions of volumes 39 to
42 can also be supplied. Full particulars of these publications can
be had on application to the Secretary.

Frettows may obtain a TransFeRaBLE Ivory Ticker admitting
two persons, available throughout the whole period of Fellowship,
on payment of Ten Pounds in one sum. A second similar ticket
may be obtained on payment of a further sum of Twenty Pounds.

A

Any Frttow who intends to be absent from the United Kingdom
during the space of at least one year, may, upon giving to the
Secretary notice in writing, have his or her name placed upon the
“dormant list,” and will then be called upon to pay an annual
subscription of £1 only during such absence, but after three years
must make a further application to be retained on that; list.

Any Frtrow, having paid all fees due to the Society, is at liberty
to withdraw his or her name upon giving notice in writing to the
Secretary.

Ladies or Gentlemen wishing to become Fellows of the Society
are requested to communicate with ‘The Secretary.”

P. CHALMERS MITCHELL,

Secretary.
Regent’s Park, London, N.W.
June, 1916.
MEETINGS
Z OF THE
ZOOLOGICAL SOCIETY OF LONDON
FOR

SCIENTIFIC BUSINESS.

1916,
TuEsDAY, OcTOBER ...... 24,
5 NOVEMBER = 4-4) avaand, Jie

The Chair will be taken at half-past Five o'clock precisely.

ZOOLOGICAL SOCIETY OF LONDON.

THE ZOOLOGICAL RECORD.

ipa ZootogicaL Record gives, by means of an annual Volume,
complete lists of the Works and Publications relating to

Zoology that have appeared during the year preceding the issue

of the Volume, together with a Subject and a Systematic Index.

Since 1906 the Zootocicat Rucorp has been amalgamated with
the Zoology volume of the International Catalogue of Scientific
Literature, and has appeared in two forms, different only in title-
page and binding, one in series with the Zoological Record, the
other forming Volume N, Zoology, of the Annual Issue of the
International Catalogue.

On account of difficulties arising from the War, the Executive
Committee of the International Catalogue is at present unable to
undertake the issue of any volumes of the 15th Annual Issue.
The Zoological Society of London, in these special circumstances,
has undertaken to produce the Zoological Record as usual, pre-
cisely in accordance with the form that the volume has assumed
since the amalgamation.

Fellows of the Zoological Society, and Institutions already on its
subscription-list, and any new subscribers whose subscriptions are
received before August “1st, 1916, will receive the volume dealing
with the literature for 1915 (Zoological Record, Vol. 52: Inter-
national Catalogue, loth Annual Issue, N, Zoology) as usual at the
end of the year or early in 1917.

The Society is able to supply complete sets of the Record on the
following terms :—
Vols. 1 to 42, price £16 10s. net. Vol. 48 and onwards at 40s. each.
The prices for separate volumes are as follows :—
Vols. 1 to 42 (except Vols. 4 and 6 which are sold with sets only),
10s. each.
Vols. 43 to 50 (obtainable separately only from Messrs. Harrison
& Sons), 40s. each.
Vols. 51 and 52. 40s. each.

Index Zoologicus. An alphabetical list of names of genera
and subgenera proposed for use in Zoology, as recorded in the
‘Zoological Record,’ 1880-1900; together with other names not
included in the ‘ Nomenclator Zoologicus’ of 8. H. Scudder. Com-
piled (for the Zoological Society of London) by Cuaruus Owsn
Warernouss and edited by Davip Suarp, Kditor of the ‘ Zoological
Record.’ London, 1902. Price to Fellows, 18s.; price to the
public, 20s., or if sold with a set of the ‘ Zoological Record,’ 10s.

Index Zoologicus, No. II, An alphabetical list of names of
genera and subgenera proposed for use in Zoology, as recorded in
the ‘ Zoological Record,’ Vols. 38-47 inclusive (1901-1910), and
the Zoology volumes of the ‘ International Catalogue of Scientific

2

Literature,’ Annual Issues 1-10. Compiled (for the Zoological
Society of London) by Cuartes Ownn Warteruovse, I1.8.0., and
edited by Davip Suarp, M.A., F.R.S., Editor of the ‘Zoological
Record.’ London, 1912. Price to Fellows, 12s. 6d. net; price
to the public, 15s. net., or if sold with a set of the ‘ Zoological
Record,’ 10s.

SEPARATE DIVISIONS OF THE ZOOLOGICAL RECORD.

Divisions of Vols. 39 to 42 and Vols. 51 and 52 of the ‘ Zoological
Record’ can be supplied by the Society, but those of Vols. 43 to 50
can be had only from Messrs. Harrison & Sons, 46 St. Martin’s
Lane, W.C.

2

S.

S
=}
(a)
or

List of abbreviations of journals, ete. .. .. 2
Special Records, viz. :—
I. General Subjects ..
Il. Mammalia :
ASTOR AA Vie SAP Rie ite Sait tee Mk
IV. Reptilia and Batrachia..
V. Pisces Ae Seu ak SS
VI. Tunicata
VII. Mollusca
VIII. Brachiopoda . .
IX. Bryozoa
X. Crustacea
XI. Arachnida
XII. Myriopoda
XIII. Insecta -. :
XIY. Echinoderma
XV. Vermes .. :
XVI. Coelenterata ..
XVII. Spongiz ae
XVIII. Protozoa 0 Tonal cnet tect cme
Index of new names of genera and subgenera. 2

P. CHALMERS MITCHELL,
Secretary.

et
WporWwWwWNMrRwWNRR Bre bDNW @bb

SPODODBDOAaGTDaDGWD®SGOAAAGADS

Recent’s Park, Lonpoy, N.W.
June, 1916.

ZOOLOGICAL SOCIETY OF LONDON...

LIST OF PUBLICATIONS.

THE scientific publications of the Zoological Society of London
are of two kinds—“ Proceedings,” published in an octavo
form, and “ Transactions,” in quarto.

According to the present arrangements, the “‘ Proceedings”’
contain not only notices of all business transacted at the scien-
tific meetings, but also all the papers read at such meetings
and recommended to be published in the ‘‘ Proceedings ” by
the Committee of Publication. A large number of coloured
plates and engravings are issued in the volumes of the
“‘ Proceedings,”’ to illustrate the new or otherwise remark-
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IP Zo So IIB, GOODIE, Ril, I,

T. Goodey, del.

SOIL PROTOZOA.

PZ So IMIG, GOODIE, Als Ml,

T. Goodey, del.

SOIL PROTOZOA.

P. Zo S. IGG, GOODEY, Pl. Il.

SOIL PROTOZOA.

cr oe sins tcomscontnantemre ainsi
<a

T. Goodey, del

ie, ZnS) Wi; (COODIE, JL iy.

56. 57

67
66

69

72

T. Goodey, del.

SOIL PROTOZOA.

ites)

ON SOIL PROTOZOA. 30

PAPERS.

8. Observations on the Cytology of Flagellates and
Amoebe obtained from old Stored Soil. Byvahlie
GoovEy, D.Sc.*, Protozoologist, Research Laboratory
in Agricultural Zoology, University of Birmingham.

[Received February 3, 1916; Read March 21, 1916.]

(Plates I.-IV. and Text-figure 1.)

INDEX. Page
inbroduc:o nee mere NRE ne: ere idea Sa ae ai oo BO
BAY eH OVS USI ey seein Se lle Oe a ne a a PE S10)

A, FLAGELLATA,
(1) Prowazekia (Bodo) saltans.

(a) Structure ......... PRR oe aad nec bcs Mecoctos, MOLUL

(2) Less) OROVELTVGINOIN. oreo eponeaccoeencnodcabdosusccasce © BILE

(@eSystematiesPositrone see eee lS
(2) Tetramitus spiralis, sp. n.

(QuStrnctiives meses eerie eee ee ee OL

()y IE DROGINCHOM... coovoococcesouncontenseockescocneace BIL

(e)M System atvcePositiony wees cee ss eeeneeee eee OLS
(3) Spironema multiciliatum.

(a) Structure ....... OER ae ce rod ose e ols)

(6) Systematic Osos, iets op ate ea Mmeson)

B. Rwizopopa.
(1) Amceba lawesiana, sp. n.

(Q)aStruchavet te nesctes. Mecsas eee OL
@)ekeproductron eee eee eee eT Oo
(Op Encystatio ny soy eee ieee 324,
(GypRemarksi ste crenata nee ate cena em OD
(2) Ameba agricola, sp. n.
(QP Structure’ ait neers eee eee OG
(O)eReproductionie eee eee eter e ee eee eet 6
(o) Remarks eee se- sce eee ee OTD
Literature... a So sarenneee Seti ae nanan Oe eas eel ta ie RAR
Explanation of ihe Blas a fale Ses RSS Oe ga ils Aa 330
INTRODUCTION.

Within the last few years a good deal of attention has been
paid to soil-protozoa, owing to the important function ascribed to
them by Russell and Hutchinson? in their hypothesis advanced
to account for the changes observed on partially sterilising soil.
According to these investigators, soil-protozoa act as a factor
limiting bacterial activity, and so prevent a normal soil from
attaining its full fertility.

It is of importance, therefore, that we should ascertain what
kinds of protozoa are present in the soil and as much as possible

* Communicated by Prof. F. W. GamsBtez, F.R.S., F.Z.S.

y+ Russell and Hutchinson, Journ. Agric. Sci. vol. in. pt. xi. (1909), and vol. v.
pt. xi. (1913).

Proc. Zoou. Soc.—1916, No. XXI. Zak

310 DR. 'T. GOODEY ON

of their life-histories ; also whether they are forms capable of
consuming bacteria, and thus able to function as a limiting factor
on soil- bacteria.

One of the chief methods by which soil-protozoa can be
studied is examination in cultures made in suitable media; and
although it has recently been claimed * that the cultural forms
are not necessarily those occurring in a tr ophic condition in
the soil, and may not therefore be. concerned in the biological
changes of the soil, yet I have quite recently obtained some experi-
mental results which point most positively to the cultural protozoa,
especially ameebe, acting as a check to the increase of bacter ‘al
numbers. | hope to publish an account of this work shortly. In
the present paper an account is given of a few forms of flagellates
and amcebe which were obtained culturally from some old soils
stored at Kothamsted Experimental Station, together with some
observations on their cytology and methods of division. The
protozoan fauna of these soils was a limited one when compared
with that of an ordinary garden or field-soil, and for this reason
presented a suitable field for wor king out the different or e@anisms
in detail.

In a recent number of ‘ The Annals of Applied Biology ’ I have
recorded the culture of amcebe and flagellates from soil bottled
so far back as 1865 and left untouched since then ; thus proving
the survival of protozoa, no doubt in an encysted condition, for a
period of 49 years. It was my intention at the outset of the work
met ely to obtain an idea of the character of the protozoan fauna
surviving in the different soils examined. In order to do this,
and to determine as nearly as possible the different species winoh
cropped up, it was necessary to make a number of permanent
stained preparations and to study these in considerable detail.
Asa result of thesé observations, | have obtained a number of
interesting facts on the cytology, mode of nuclear division, ete., 1n
several of the forms examined.

The samples of soil tested for protozoa were taken from bottles
of soil obtained originally from five of the fields under experi-
mental cultivation at Rothamsted. These were: Broadbalk soil
bottled in 1865, Geescroft soil bottled in 1865, Agdell soil bottled
in 1867, Hoosfield soil bottled in 1868, and Barnfield soil bottled
in 1870.

MeruHops.

As a culture medium, saline egg-albumen was used and found
very serviceable. A small quantity of soil was placed in this
medium, contained in a circular flat-bottomed glass dish furnished
with a close-fitting flat lid. This was then put into an incubator
at 22°C, or left at room-temperature. After a few days, micro-
scopic examination of the culture revealed active protozoa. These
were frequently found on the surface or at the bottom of the

** Russell, E. J., “ Soil Protozoa and Soil Bacteria,” Proc. Roy. Soc. B, vol. Ixxxix.
p. 76 (1915).

SOIL PROTOZOA. 311

liquid ; and in order to obtain these forms, coverslips were floated
on the surface and placed on the bottom. In other cases the vast
majority of the organisms occurred about midway in the depth of
the cuiture, and in order to obtain coverslip-preparations of these
I drew out small quantities of the culture where the protozoa were
thickest by means of a fine pipette, and then made smears with
the liquid. As a rule, three cultures were made from each soil,
in order to obtain a representative fauna, and the cultures were
examined for several days to note any succession of forms. Cover-
slip-preparations were fixed in Maier’s solution or in Bouin’s
fixative,and iron-hematoxylin was used as the stain throughout;
occasionally preparations were counterstained with lichtgriin-
picric.

I propose in the following pages to deal with the protozoa
encountered under the heading of the group to which they helong,
mentioning the particular soil or soils from which they were
taken.

A. FLAGELLATA.

i) PROWAZEKIA (Bono) sattans Ebrbg. (PI. L.)

In one of my cultures of Barnfield 1870 soil a very small
jumping flagellate occurred in abundance at the bottom of the
liquid. It appeared somewhat bean-shaped when seen under a
low power, and I at once concluded that it was Bodo saltans. A
coverslip- preparation was made, and on it I obtained many
dividing organisms, from a study of which I have been able to
work out fairly completely the process of nuclear division. J will
describe the structure of a normal organism, and then deal with
the question of identification and nomenclature.

(a) Structure.

The body is somewhat bean-shaped and is oval or round in
cross-section. Seen from the ventral aspect—z. e. the side on which
the flagella arise (fig. 1)—the anterior end appears flattened and
is turned towards the left, where it terminates in an almost
straight edge, This anterior portion of the organism is really an
extension of the dorsal region, and is separated on the ventral
surface from the main part of the body by a considerable depres-
sion, in which the mouth is situated. Fig. 2 shows a side-view
and the relations of the anterior end to the depression, ete.

The trophonucleus is generally found towards the left, side of
the body when the organisin is viewed from the ventral surface.
It consists of a central deeply staining karyosome, which is con-
nected with the nuclear membrane by means of strands which
stretch across the extra-karyosomic zone, and at their insertion
on the membrane produce slight thickenings.

The kinetonucleus is an irregular granular mass, often some-

what triangular in outline, and, on the whole, stains less intensely
91 :

33 2 DR. T. GOODEY ON

than the karyosome of the trophonucleus. It has no nuclear
membrane and is situated towards the ventral surface, close to
the upper end of the mouth-depression. Its position can readily
be made out in figs. 1 & 2

The two flagella arise close to the anterior surface of the
kinetonucleus from very indistinct blepharoplasts. The anterior
flagellum is the shorter of the two, the posterior one being two
or two anda half timesits length. ‘he organism ingests bacteria,
which no doubt serve as a source of food, for the protoplast is
often packed with cocci and other small forms of bacteria.

(b) Reproduction.

The first indication of approaching division which I have been
able to find is the doubling of the anterior flagellum (fig. 4). At
this stage no change is visible in the appearance of the nuclei. I
am unable to say how the flagella become duplicated, but, judging
from the fact that 1 have found no organisms showing protrusions
like flagellar buds, and also that when the flagella become doubled
the members of each pair are equal in length, I am inclined to
the view that the original flagella split longitudinally.

The posterior flagellum becomes doubled later than the anterior
one, and at this time the tr ophonucleus shows a marked change
in appearance. The karyosome becomes much reduced in volume
and divides into two equal parts, whilst at the same time the con-
necting strands between it and the nuclear membrane disappear.
The nucleus elongates a little and stains rather more deeply,
doubtless owing to the liberation of a chromatinic substance from
the karyosome (figs. 5 5&6). Irregular granules of chromatin now
appear at the periphery of the nucleus as the result of further
fragmentation of the karyosome, and become concentrated towards
the equatorial region, This particular stage is very difficult to
determine, and T cannot say whether an equatorial plate is pro-
duced. §o far as I can make out, there is merely an aggregation
of chromatin granules on the surface of the nucleus in this region.
These granules finally concentrate into four principal larger ones,
which are arranged in two pairs. The stage figured on fig. 9 shows
them apparently connected by two crossing strands. The nucleus
now elongates, each end being somewhat pointed, and each pair

of granules becomes drawn towards eg ends of the nucleus.
hese stages are shown in figs. 10, 11, & 12. Soon after this a
constriction appears in the middle of the caneleus, which now
becomes rather dumb-bell-shaped (fig. 15), At about the same
time, or even earlier, the four granules of chromatin begin to
show signs of breaking down, and also stain less intensely
(figs. 13-16). The constriction at the centre of the nucleus
becomes carried still further, until two triangular daughter-
nuclei are formed, each of which contains rather faintly staming
chromatin granules. I have found it impossible to trace the
later stages in the reorganisation of the daughter-nucle1, owing

SOIL PROTOZOA. 313

to the fact that the flagellates are almost always crowded with
deeply staining bacteria which obscure the nuclear elements.
The next stage of the trophonucleus which I have certainly made
out, is that in the daughter-organisms where the granules have
become concentrated again into a central karyosome separated
from the nuclear membrane by a clear zone (fig. 21). Whilst
the above changes are proceeding, the kinetonucleus undergoes
certain alterations. It increases in volume, and may either be
triangular or rhomboidal in outline. One pair of flagella moves
to either side of it, and at these points the kinetonucleus
becomes somewhat drawn out. Soon after the formation of
daughter-trophouuclei has taken place, the kinetonucleus elon-
gates considerably, so that the pairs of flagella are carried farther
apart. This elongation is carried on until a fairly long band of
kinetonuclear material is produced, which finally separates into
two portions—the daughter-kinetonuclei. [I have not found the
stages showing the constriction and division of the kinetonucleus,
but there can be no doubt, I think, that the process is simple and
direct.

Concurrently with the elongation of the kinetonucleus, the
body of the flagellate becomes oval and then grows laterally, So
that the longer axis of the body i is that running from side to side,
not antero- posteriorly. Division of the body is initiated by the
formation of a constriction on the now shorter axis of the body,
and becomes more and more pronounced until the two daughter-
organisms are connected only by a short narrow strand of proto-
plasm. This finally breaks, and the two small organisms are
produced. Division thus takes place along the antero-posterior
axis of the body, and is therefore longitudinal.

(c) Systematic Position.

Because of the rapid spasmodic jumping motion exhibited by
this organism in life, the name sa/éans is eminently applicable to
Tlic Frc om its gener al shape also, and the presence of two flagella,
the posterior one being longer than the anterior, 1t easily fits into
the genus Bodo. The difficult point to determine, however, 1s
whether it should be classified as a Bodo or as a Prowazekia, for it
possesses a kinetonucleus.

Alexeieff (11 & 712) is of the opinion that all the species of
Bodo possessa kinetonucleus, and holds that the genus Prowazekia
isuntenable. According to hs author, my organism should go in
the genus Bodo. Iam a convinced, eweved that his assertion
concerning the presence of a kinetonucleus in Bodo is correct, for
T have Sicaimed a bimastigote form having the anterior flagellum.
shorter than the posterior one in which there is certainly no lemeto-
nucleus, and which undoubtedly belongs to the genus Lodo. i
therefore propose to place my organism in the genus Prowazekia.
At the same time, however, I incexk the name Bodo in brackets,
because I think this form is Ailewsieall with Bodo saltans. Alexeieft

‘

314 DR. T. GOODEY ON

(11) gives the dimensions of Lodo saltans as 6-10 p in length by
3-5 w in breadth. My organism is much smaller than this,
measuring from 5-6 p in length by 2°5-4 «in breadth. In this
respect it comes nearest to Bodo minimus Klebs, which is from
4-5 » in length and is considered to be one of the smallest
flagellates. The latter form, according to its original describer,
is changeable in shape and has a creeping movement. My
organism is very constant in shape, and always moves in rapid
jumps followed by intervals of rest. There can be no doubt, then,
that its specific name should he saltans.

Alexeieff (711, p.508) says that, without a doubt, Bodo saltans is
the same organism as Pr owazeleia parva descr ibed by Nagler (10).
The latter organism differs considerably from that described above
in several respects. Its protoplast is labile and takes on a great

variety of shapes, whereas my organism is constantly bean- shaped.
The method of division in both the tropho- and kinetonucleus is
quite different from that described above. There is no frag-
mentation of the karyosome of the trophonucleus with the
ultimate formation of four chromatin granules or* chromosomes,
but merely an equal division of the karyosome by promitosis, a
centriole and a centrodesmose taking part in the process. The
kinetonucleus divides in a similar manner. It is obvious, there-
fore, that Nigler’s organism is a species distinct from mine, and
his name Prowazekia parva should be allowed to remain.

Among the descriptions of members of the genus Prowazekia
by: Hartmann & Chagas (10), Niigler (11), Mathis & Léger (710),
Withmore (’11), Mart tin (13), aint Bela (14), the division of the
trophonucleus of Prowazekia asiatica by Withmore comes nearest
to that which I have given in the foregoing account. His figs. 18
and 19 show the presence of four principal chromosomes in the
dividing nucleus, which, however, differs considerably in its
earlier division stages from those observed in my organism.

(2) TErRAMITUS SPIRALIS, sp. n. (Pls. II., II., figs. 23-45.)
(a) Structure.

This interesting organism occurred in cultures from three
different soils, viz.: Agdell 1867, Broadbalk 1865, and Barn-
field 1870. I have been fortunate in obtaining it in large
numbers, and have been able to make out most of the details
of its structure and mode of division. JI have not, however,
observed it in the encysted condition, although I kept my
cultures for a long time and had the organisms under close
observation in hanging drops. I will first describe its structure
and movements, and then deal with the process of reproduction.

In the majority of cases the body is pyriform in shape, having
its extreme anterior end somewhat pointed. It may, however,
become much elongated, and then appears more cylindrical.

A groove runs spirally from the anterior to the posterior end

SOIL PROTOZOA. SS)

of the body. Seen from the ventral aspect—i. e., from the side
on which the flagella arise (text-fig. 1)—the groove proceeds from
right to left, and then curves round to the dorsal surface and
reaches the posterior end of the body. If the body is of normal
length there is only this one turn in the spiral, but if the
organism is elongated there may be two turns. The accom-
panying text-figure, drawn from a specimen immediately after
killing with osmic-acid vapour, shows the shape of the organism,
the position of the groove, and the disposition of the flagella.
The latter are arranged in two pairs—a shorter anterior pair and
a longer posterior pair. The anterior pair is direded forwards
during motion and the posterior pair is trailed backwards.

Text-figure 1.

DS

Tetramitus spiralis.

Outline drawing as seen in ventral view, showing the relations of the groove (g.).

The members of each pair are frequently very closely applied,
and often appear as one thick flagellum. When in active
movement, which is caused by the lashing of the anterior pair
of flagella, the organism progresses very rapidly and rotates on
its long axis. The posterior pair of flagella appears to he
within the groove and extends beyond the posterior end of the
organism, which swings from side to side of the line of motion
owing to the rapidity of progression. The mouth is very
difficult to locate, being ill-defined, but in certain examples I
have made out its position a short distance from the anterior
end as a depression in the groove, Bacteria are ingested and no

Bs DR. T. GOODEY ON

doubt serve as a source of food. There does not appear to be a
contractile vacuole. The body measures from 8-12 y in length.

Fixed and stained material shows that the protoplasm is
alveolar in structure, and that the nucleus is situated at the
anterior end of the body, as in Trichomonas and Trichomastix and
similar forms. The nucleus is a vesicular structure of variable
size, but often of quite large dimensions. There is no central
karyosome, but the chromatin is disposed irregularly in two or
more masses. Very frequently there are two semilunar blocks
of chromatin situated on either side or on the anterior and
posterior borders of the nucleus. It will be seen from the
figures in what an irregular fashion the chromatin is arranged.

“The flagella arise from blepharoplasts placed just anter iorly to
the nucleus. They are four in number and, as stated above, are
disposed in two pairs. They take their origin in four blepharo-
plasts, which when seen in side view (fig. 25) appear as two large
granules in contact with each other, but when seen in face view
are easily recognisable as being a group of four distinct granules
in intimate contact (figs. 26 & 27). The posterior pair of granules
is connected with the nucleus by means of two rhizoplasts,
which appear as one short rod in fig. 25 but are well shown
in the ventral view obtained in fig. 26. JI made a very careful
examination of this region, in order to determine if rhizoplasts
were present connecting the anterior pair of blepharoplasts
with the nucleus, and I am satisfied that the pair connecting the
posterior blepharoplasts is the only one.

(b) Reproduction.

I will deal first with the multiplication of the flagella, because
this always occurs prier to the division of the nucleus. The new
flagella are produced by outgrowths from the body of the organism,
and not by splitting of the old flagella. In this it resembles Copro-
monas, Trichomonas, and many other flagellates. The process is
initiated by the growth in an anterior direction of each pair of
blepharoplasts : a point very difficult to make out in many of the
organisms, but well shown in other cases, one of which is repre-
sented in fig. 27. The flagellar buds arise as delicate hair-like
outgrowths from the developing buds of the blepharoplasts. Each
pair of flagella has thus a new pair of flagella produced imme-
diately anterior to it. The new ones do not stain so deeply as
the old flagella, as will be seen from the figures representing
different stages of the division of the nucleus. In this way the
original posterior pair acquires a new anterior pair, and the original
anterior pair grows longer and becomes the posterior pair of one of
the daughter-organisms, at the same time acquiring a new anterior
pair. The four pairs of flagella thus produced gradually separate
into two sets of two pairs, which finally come to take up positions
at either end of the anterior face of the nucleus; but this migra-
tion takes place at different periods during the progress of nuclear

SOIL PROVOZOA. SIL

division. For example, fig. 28 shows an organism in which the
flagella are widely separated, although the nucleus shows no sign
of approaching division. Iam unable to state the fate of the pair
of rhizoplasts.

In the earliest stages in the division of the nucleus the
chromatin seems to undergo some process of dissolution and re-
organisation whereby certain parts of it, which stain less intensely
than the rest, gradually come towards the centre of the nucleus.
These take the form of roundish or irregular granules which,
at the stage shown in fig. 31, appear to be arranged in a fairly
regular manner on a kind of reticulum or network of linin
threads. Whilst these changes are taking place, the remainder
of the chromatin, having deeply staining properties, becomes
arranged in the form of a ring round the periphery of the
nucleus. ‘This does not always happen in the very early stages,
however, as is shown in fig. 33, where the deeply staining
chromatin is still present as three blocks, whilst at the centre of
the nucleus there is a group of lighter granules. The changes
which occur in the lighter staiming granules are very difficult to
make out, but it seems as though they gradually concentrate
towards the centre of the nucleus, and there become arranged in
an irregular manner on a kind of plate. .In figs. 32 and 33 there
appear to be six principal granules disposed in two bands upon
what seems to be a spindle formation. I do not wish, however,
to lay stress on the presence of a spindle within the nucleus, for
I have failed to make it out with any degree of distinctness, and
even in those examples which present the spindle appearance,
there are always ivregularly-branching linin strands running in
various directions, as shown in figs. 35 and 36.

In the stages represented in figs. 34-36, four principal round
granules are present. These represent the nearest approach to
chromosome formation in the whole series of changes. It would
appear from these figures that the nucleus produces division
centres from within, and is not dependent on the migration
of the blepharoplasts to their antero-lateral positions for the
formation of its poles of division. J have not been able to trace
further the movements of the four internal chromosomes. In
the succeeding stages the nucleus elongates somewhat laterally,
and the peripheral ring of chromatin begins to break up and
travel towards the lateral poles, at the same time advancing on
to the dorsal and ventral surfaces of the nuclear membrane. In
fig. 38 the connections between the blepharoplasts and the
nucleus which ultimately become the rhizoplasts are fairly
clearly seen. The lateral elongation of the nucleus now becomes
more pronounced, and the chromatin, travelling along the linin
threads of the nuclear network, becomes arranged in small
granular masses towards the lateral poles. At this time the
threads stretching across the centre of the nucleus can be made
out fairly easily (figs. 39 & 40). These linin threads are doubt-
less absorbed, and two laterally situated daughter-nuclei are thus

\

318 DR. T. GOODEY ON

produced, The chromatin now becomes re-arranged in granules
of varving size and shape, as shown in figs. 41-44,

During the later stages of nuclear division, the anterior sur-
face of the body becomes much drawn out and flattened. A
depression now appears on this surface of the body and gradually
travels backwards, and at the same time, in some cases, the
protoplast extends laterally (fig. 44). In other cases the body
becomes triangular in outline, and large vacuoles appear towards
the centre of the body and, by rupturing, assist in the pro-
duction of the daughter-organisms (figs. 42 & 43). Division of
the body is thus longitudinal in direction. Fig. 45 shows two
newly-formed organisms which have recently separated, their
drawn-out tail-ends overlapping slightly.

(c) Systematic Position.

The possession of four flagella places my organism undoubtedly
in the genus 7etramitus Perty, and though this to-day is a very
mixed assemblage of forms, comprising, as it does, the free-living
organisms described by Perty and by Klebs (92), and also the
parasitic forms Vetramitus (Macrostoma) mesnili (Wenyon, ’10 a)
and Tetramitus (Macrostoma) caulleryi (Alexeieff, 11 a), there is
no reason why I should create a fresh genus for its reception.
As, however, I have been unable to discover any description or
figures of any free-living member of the genus which fits my
organism, I have decided to make a new species of it, namely,
spiralis.

(3) SPIRONEMA MuULYICcILIATUM Klebs. (Pl. III. figs. 46-48.)
(a) Structure.

This highly interesting organism occurred in one culture made
from Broadbalk 1865 soil. It appeared both on the surface and
at the bottom of the culture. My attention was first attracted
to it by reason of its great length and its peculiar method of
locomotion. It moved slowly in a very hesitating jerky manner
for the most part, but would suddenly exhibit rapid and violent
spiral twists commencing at its anterior end and travelling down
the body, at which times it was propelled at a reasonably fast
pace. It was obvious that the organelle causing the slow jerky
motion were situated at or towards the anterior end, though
they could not be distinguished under a low power of the micro-
scope. ‘Towards the posterior end a contractile vacuole could be
seen in diastole and systole.

I was able to obtain film preparations which, when fixed and
stained, revealed the structure of the organism very clearly.
The body is extremely long in comparison with the width, and
is dorso-ventrally flattened. It measures anything from 20-50 pu
in length, and averages about 4 win width. The middle region
is generally the widest part of the body. The anterior end is
either rounded or has a lateral knob-like portion on either side.

SOIL PROTOZOA. 319

The posterior end is drawn out into a long and exceedingly fine
tapering tail, and the contractile vacuole occurs just where the
body begins to narrow down.

The flagella are numerous and comparatively short. They
vary in number from seven to eighteen, and the smaller the
organism the fewer the flagella. They are situated in most cases
in two lateral rows towards the anterior end of the body, one
row being dorsally and the other ventrally placed. I have care-
fully noted the disposition and number of the flagella, and find
that they are not equally distributed on either side, but exhibit
a considerable amount of variation in this respect. Klebs, on
his Pl. xvi. fig. 9¢, shows a row of flagella extending backwards
on one side as far as the beginning of the tail. JI have not
found anything like this in my organisms.

A few of the organisms were fixed just as they were twisting
spirally, and one of these is shown in fig. 48. It will be seen
from this that the edges of the body are curved, and that the
flagella have their origin close to the edges. Each flagellum
arises from a small basal granule or blepharoplast distinctly seen
in the stained material. J cannot say whether there is a mouth,
and although I watched the creatures in life for a long time, I
never saw them take in food. There are numerous large granular
bodies, however, in the cytoplasm in many of the forms which
appear to be ingested bacteria, and because of this, I am of
the opinion that a mouth is present. I believe it is situated
towards the anterior end, for I have made out, in some cases, a
somewhat lighter area here which might be considered as the
mouth. In the greater part of the body the cytoplasm is very
finely granular and evenly distributed, but towards the posterior
end, in the region of the contractile vacuole, it is frequently much
vacuolated.

The nucleus is a very interesting structure. It is, in most
cases, of considerable length, and is situated about half-way
down the body. It consists of a long narrow rod of granular
material, frequently one-quarter to one-third of the body in
length. Towards the middle of it is placed a circular karyosome
of deeply staining chromatin. The extra-karyosomic portion of
the nucleus appears to be very little different from the general
cytoplasm in staining reactions, and is separated from the latter
on all sides by a very narrow clear space. ‘There does not appear
to be any nuclear membrane. At all events, I have not made
out anything comparable with the nuclear membrane of other
flagellates and ameebe. From the appearance of the stained
examples, it seems that all the chromatin is concentrated in the
deeply staining karyosome, there being only small scattered
granules in the rest of the nucleus.

The nearest approach to this nuclear apparatus which I have
been able to find, is that which occurs in certain Huglenoidea,
for example, in Huglena viridis (Keuten, 95), in which there
is a fairly large nucleus consisting of a central karyosome

320 DR. T. GOODEY ON

surrounded by a granular portion, the bulk of the chromatin being
located in the karyosome, and the rest disposed in fairly large
granules in the extra-karyosomic part. At first I was inclined
to regard the extra-karyosomic portion of the nucleus as a
macronucleus, and the karyosome as a micronucleus, taking the
organism to be a ciliate. But the comparatively large size of the
karyosome, and the fact that it occurs embedded in the centre of
the rest of the nucleus, seems to rule out its micronuclear
homology. Unfortunately, none of my preparations shows the
organism dividing, so that I am unable to indicate the behaviour
of the nucleus during these most important phases.

(b) Systematic Position.

After studying Klebs’s (92, p. 350) description and figures
of the organism Spironema multiciliatum, there can he no doubt
that my organism belongs to the same genus. It agrees in
possessing about the same number of flagella, which are similarly
situated, a posteriorly placed contractile vacuole, and in general
appearance is the same. Klebs’s organisms measured 14-18 p in
length by 2-34 in width, whilst none of my organisms is less
than 20 w in length, and they are about the same as his in width.
This difference in length does not appear to me to be sufticiently
important to warrant the creation of a new species for the
reception of my organism. Klebs describes and figures two
lateral spiral grooves commencing at the anterior end of the
body and extending backwards as far as the beginning of the
tail, and says that the flagella are inserted on the edge of each
groove. These grooves correspond, I believe, to the dorsal and
ventral surfaces of my organism when exhibiting its spiral
twist, for there are really no true lateral grooves, the body being
so thin. He made out nothing of the nuclear apparatus in his
organisms, but from their appearance, and the number and dis-
position of the comparatively short flagella, he looked upon the
creatures as probably forming a connecting-link between the
Cihata and Flagellata, as the following quotation shows :—“ Ich
halte es fiir sehr wahrscheinlich, dass diese vielgeisseligen
Flagellaten einen Ubergang zu den Ciliaten bilden und m%chte
speciell die Aufmerksamkeit auf diese noch so wenig bekannten |
Formen lenken.”

From the appearance and structure of the nucleus, 1 think it
is best to include Spironema in the Flagellata, though there is
something to be said for Klebs’s suggestion of it being a connect-
ing-link between the Ciliata and Flagellata. The posterior
position of the contractile vacuole is a ciliate characteristic,
whilst the organelle at the anterior end might equally be re-
garded as long cilia or short fiagella. The forms which Klebs
examined were obtained from ditch-water, so that there is
nothing very remarkable in my obtaining the same organism in
cultures of soil.

SOIL PROTOZOA. Sek

B. Rurzopopa.

Each of the soils yielded small ameebe of the imax type, and
I have been fortunate in obtaining a number of stages in the
division of one or two of the forms.

(1) AmMa:BA LAWESIANA, sp. n. (Pls. III., IV., figs. 49-65.)

I propose this name * for a small ameceba which occurred in a
culture of Broadbalk 1865 soil. I put up the culture in the
hope of obtaining another kind of protozoon, but instead of
getting this particular organism, I obtained an almost pure
culture of the amceba in question. Unfortunately, I was unable
to devote much attention at the time to observing the living
organisms, and for this reason I am not able to state definitely
whether a contractile vacuole is present or not. The conditions
prevailing at the surface of the culture were very favourable to
active life, for my permanent preparations show that the amcebee
were ingesting large numbers of bacteria and dividing forms are
fairly abundant.

I have been fortunate in obtaining an almost complete series
of dividing organisms, and from the appearance presented by the
nucleus during these phases there can be no doubt that this
amoeba is very closely related to Amaba glebe, which Dobell
(14) has recently described in great detail. It is also similar in
its nuclear changes to dmaba lamellipodia (Gliiser, 12), and the
large ameceba from liver-abscesses, described by Liston and
Martin (11), and also to Ameba cucumis and Ameba gobanni-
ensis (Martin & Lewin, 714).

Nevertheless, it differs from all these in certain important
details, which are dealt with later on, and for this reason I
propose to create a fresh species for its reception.

It is rather smaller than dimwba glebe, and the following are
some of its principal measurements :—

Diameter of rounded forms ......... 12-15 p.
Diameter of nucleus .................. A)
Diameter of karyosome............... 2 pi.
Diameter of ripe cysts .............:. 10-11 p.

(a) Structure.

When in motion, the body becomes extended in the typical
limax shape and presents a blunt advancing pseudopodium. The
protoplast is composed of an almost hyaline ectoplasm and a much
vacuolated endoplasm. In fig. 49 the alveoli of the endoplasm
are very irregular in shape and distribution, but in the almost
spherical forms assumed during nuclear division the alveoli are
fairly regularly distributed throughout the endoplasm and are
more equal in size. The body is often crowded with ingested

* T have named this amoeba after Sir John Lawes, the founder of the Rothamsted
Experiments.

Bp DR. T. GOODEY ON

bacteria, and in those forms exhibiting the slug-like appearance
the posterior end is frequently covered by an adherent mass of
bacteria (fig. 49).

The nucleus consists of a large karyosome, which is separated
from the nuclear membrane by a clear zone and an outer ring of
faintly staining granules. The latter may apparently occur as
very small discrete particles, as in fig. 49, or as a single ring of
small blocks, as in fig. 50. I have not succeeded in making out
any connecting strands between the karyosome and the nuclear
membrane. It resembles the nuclei of Amwba glebe and Ameba

lomellipodia in possessing the ring of faintly staining granules.

(b) Reproduction.

The animal ceases to wander about and comes to rest, at
the same time becoming spherical. JI have not made out
pseudopodia in any of these globular dividing forms. The
earliest stage in the division of the nucleus which I have dis-
covered is shown in fig. 50, where the karyosome has broken
down into four principal masses. This fragmentation of the
karyosome is continued until the central part of the nucleus
originally occupied by the karyosome, or an area slightly larger
than this, becomes filled with a mass which appears to be made
up of very faintly staining particles, amongst which are lodged
the rather more deeply staining granules produced by the dis-
integration of the karyosome. I cannot say whether the ring of
faintly staining granules occurring in the “resting” nucleus
takes any part in the division or whether they disappear.
Dobell says that in Ameaba glebe they entirely disappear, and it
may be the same in my organism. The fine particles produced
by the fragmentation of the karyosome stain much less intensely
than the original karyosome. They gradually aggregate and
produce somewhat larger granules, which become connected up
into a sort of chain formation, which lies in an irregular manner
among the mass of linin particles. The nuclear membrane does
not disappear, and does not seem to become any less distinct than.
during the ‘“‘ resting” condition of the nucleus.

The chain of chromatin granules or chromosomes, as they may
perhaps be called, approaches the equatorial region of the nucleus,
where it ultimately becomes disposed in the form of a ring
(fig. 53), At this stage the first indications of a spindle make
their appearance, becoming elaborated out of the linin matrix in
which the ring of chromatin granules has been lying. The ends
of the spindle are at first broad and rather flattened, but later
on they become very sharply pointed. The plane in which the
long axis of the spindle lies is slightly oblique to the horizontal
plane of the nucleus. This is well seen in fig. 54, where the two
ends of the spindle extend beyond the limits of the nuclear
membrane which is represented in optical section, one end being
over and the other under the nuclear membrane.

SOIL PROLOZOA. 323

In the equatorial ring of chromatin granules I have not been
able to distinguish at all clearly the separate constituent chromo-
somes. ‘This may be due to the fact that they become very
closely packed together. They are most distinctly seen in
figs. 53 & 54. The equatorial ring becomes divided into two in
the plane at right angles to the axis of the spindle. I have not
discovered any organism showing the actual constriction of the
chromosomes, but have obtained a stage where the two daughter-
rings are very closely apposed (fig. 56). In the succeeding stages.
of division the two rings of chromatin gradually become sepa-
rated from each other by a wider interval, owing to the elonga-
tion of the spindle, the fibres of which become quite distinct:
across the centre of the animal.

The poles of the spindle remain sharply pointed until a late
stage in the separation of the new chromatin bands (figs. 59 & 60).

After the stage which is depicted in fig. 55, the word ** band ”
more accurately describes the appearance presented by the
daughter chromatin elements, for I have not been able to make
out any ring-like structure after carefully focussmg on these
parts. Neither have I been able to distinguish separate chromo-
somes, for each band appears to be composed of numerous fine
granules. The nuclear membrane appears to remain intact up
to the stage shown in fig. 56, after which, however, it is not dis-
tinguishable, and I suppose it disappears entirely.

The animal now elongates in the direction of the long axis of
the spindle and becomes ellipsoidal in outline (fig. 59). This
figure shows an interesting condition of the spindle-fibres be-
tween the chromatin bands, in that a twist in them seems to
have been produced as though one of the bands had rotated
through an angle of 180°. A constriction now appears round
the animal, and. the first stage in the process of fission is brought
about. Fig. 60 represents Phi stage, and it is easily seen that
the poles of the spindle are sharply pointed at this time.

The process of the re-formation of the daughter-nuclei now
begins. The pointed poles of the spindle disappear, and the
chromatin granules become scattered irregularly in a mass of
faintly staining linin particles which are apparently formed by
the break-up of the outer portion of the spindle-fibres. In
fic. 61 one daughter-nucleus is seen to consist of a crescentic
area of linin particles in which the small granules of chromatin
are scattered, whilst at the other end of the spindle the daughter-
nucleus consists of a small though well-developed central evanule,
no doubt formed by a fusion of smaller granules, surrounded by
a ving of linin particles, from which it is separated by a clear
zone, I think there can be no doubt that the crescentic daughter-
nucleus represents an earlier stage in the process of reorganisa-
tion than the round form in the other part of the constricted
amceba.

In fig. 62 the constriction of the parent ameba has been
carried a little further, and the spindle-fibres between the

324 DR. T. GOODEY ON

re-forming nuclei could be made out on focussing very carefully.
It can be seen from this figure that the reorganisation process
takes place earlier on the outer side of each nucleus than on the
inner side. JI have not been able to distinguish any reticulate
arrangement in the linin particles which are laid down as the
process of reorganisation commences. The fission of the body
is now carried a little further, and the two daughter-organisms
are produced. There does not appear to be any connecting
strand of protoplasm between the two products of fission, though
in all these stages this region is extremely difficult to make out,
owing to the presence of large masses of adherent bacteria,
inte! I have purposely omitted from the drawings.

The new karyosome now increases in size by the absorption of
the remaining fine granules of chromatin. It is no longer
possible to distinguish any spindle-fibres, and each nucleus
becomes rounded off. ‘The new nuclear membrane is apparently
formed from the zone of linin surrounding the new karyosomes,
and from this zone also the peripheral ring of feebly staining
granules is also produced. The only difference between the
nucleus of the stage represented in fig. 63 and that of a full-.
grown animal is merely one of size.

(c) Eneystation.

On the same preparations which showed dividing animals, I
obtained a few stages revealing the process of encystation. The
first indication of this is the production of intensely staining
small round granules in the endoplasm, as shown in fig. 64. In
this animal I could discover very few ingested bacteria, and it 1s
evident that the normal process of digestion becomes suspended
with the beginning of encystation. There is practically no
difference in the appearance of the nucleus during the process of
encystation, and even when the cyst-membrane has become well
defined, as in fig. 65, it was still possible to distinguish all the
principal structures of the nucleus. The karyosome in the
encysting animals is rather smaller than in normal active forms.
As encystation proceeds, there is a gradual contraction of the
endoplasm round the nucleus, so that the ectoplasm is left asa
distinct region free from granules. This is particularly well
shown in fig. 64, where the line of separation between the two
constituents of the protoplast is especially marked. The animal
diminishes somewhat in bulk, and the cyst-membrane is laid
down around it. This later on becomes much corrugated and
indented, as shown in fig. 65. It is quite well defined at this
stage, but becomes somewhat thicker at a later period; a point
which I have determined by the examination of empty cysts.
There does not appear to be an endocyst. In possessing deeply
stainable granules, the cysts differ from those of Ameba glebe,
in which Dobell describes non-stainable extremely refractile
granules. Ido not know what the real nature of the granules

SOIL PROTOZOA. B45)

produced in the endoplasm is, but they are of fairly common
occurrence in the cysts of other forms of limaxamebe. At all
events they are not particles of chromatin extruded from the
karyosome, for this does not diminish in bulk to any great
extent, and, moreover, there is a sufficient volume of granular
material produced in the endoplasm to make several karyosomes
if it were fused together. Probably they are of a reserve food
character. I cannot, however, throw any light on their presence
or absence in newly excysted organisms, for I did not make any
observations on the excystation of this amezba.

(d) Remarks.

Dobell has gone very thoroughly into the differences and
similarities between his Amwba glebe and its nearly related
forms, so that it is quite unnecessary for me to go into this
question in detail. I will merely point out, therefore, in what
respects my organism differs from or resembles dmeba gleba.

It is obvious, from a comparison of the figures illustrating this
account and that of Dobell, that the amcebe to which they refer
are very closely related in their method of nuclear division.
The type of division is the same in each, and it is merely in
details that differences are presented. The most important are
the following :—

1. Ameba lawesiana is a somewhat smaller organism than
Ameba glebe.

2. The nuclear membrane persists to a much later stage of
division in Ameba lawesiane than in Ameba glebe.

3. The spindle formed in the division of the nucleus is
sharply pointed at each end in Amceba lawestana and
is rounded or barrel-shaped in Amba glebe.

4, The resting-cyst of Ameba lawesiana is irregular in out-
line, whereas that of Ama@ba glebe is perfectly round
and has a smooth outer wall.

5. Within the endoplasm of the cyst of Ameba lawesiana
large numbers of deeply staining granules are produced,
whereas in the cyst of Amaba glebe highly refractive
granules occur.

(2) Ama@Ba AGRICOLA, sp.n. (PI. LV. figs. 66-74.)

I propose to describe under this specific name a small amceba
which occurred in one of the cultures made from Hoosfield 1868
soil. It exhibits some rather remarkable appearances during
the division of the nucleus, which seem to differ from any of the
already described nuclear divisions in amcebe ; and it is on the
strength of this fact that I propose the creation of a new species
for its reception. J made no special observations on the live
animals, so that, in this respect, what I have to say about them
is, unfortunately, incomplete. My notes merely record the
presence of numerous imax amcebe in this particular culture.

Proc. Zoot. Soc.—1916, No. XXII. 22

326 DR T. GOODEY ON

I made one or two film preparations, which, after fixation and
staining, showed the presence of large numbers of amebie,
together with the flagellate Cercomonas longicauda. In gowg
carefully over one of these preparations, | discovered a number
of interesting stages of dividing nuclei; and though I have not
obtained a very complete series of these, I have made out
sufficient to show that I am dealing with an organism hitherto
undescribed.

(a) Structure.

There is nothing remarkable in the appearance of the ordinary
individuals. The. body presents an endless variety of shapes,
and the pseudopodia are very irregular and lobose, whilst the
distinction between ectoplasm and ‘endoplasm i is not at all clear.
So far as I can ascertain, the endoplasm is not alveolate in
structure. At any rate, if alveoli are present they are not large
and distinct like those in Amwba lawesiana and Ameba glebe,
for in only one of the animals could I make out anything at all
approaching alveoli. I do not wish to emphasize this point, how-
ever, for the preparation was slightly over-difterentiated and the
cytoplasm in all the amcebee was only very faintly stained. On
the whole, the endoplasm appears very finely granular in
structure, with somewhat denser masses scattered about in it ;
and the figures which illustrate this account represent it fairly
accurately.

‘he resting nucleus consists of a central deeply staining
karyosome, separated by a clear zone from the nuclear mem-
brane, with which it appears to be connected by very feebly
staining strands stretching across the zone at various points.
if The not been able to "make out the presence of a ring of
granules just within the nuclear membrane as in Amaba
lawesianc. The principal measurements are as follows :—

Length of body 12-15 yw, though this measurement is not of
“much value because of the ver y regular shape of the
organism.

Diameter of nucleus, 2°75--3 pu.

Diameter of karyosome, 1°8-

It is thus rather smaller than Ameba lawesiana.

(b) Reproduction.

The body does not become globular during nuclear division
as in Ameba glebe and Amaba lawesiana, but retains its very
irregular appearance. The karyosome in the earliest stages of
division loses its rounded shape and increases in size. At the
same time it begins to break up into a number of ill-defined
granules, which appear to rest on a matrix which stains only
feebly. I have only encountered a few of these early stages, and
therefore cannot give much information concerning the changes
which go on at this period.

The final result of the break-up of the karyosome is the

SOIL PROTOZOA. 327

formation of irregular chromosomes, some of which appear rod-
like and others rather rounded. While these changes are taking
place the whole nucleus increases in size and becomes barrel-
shaped, attaining a length of 6-7 u. A few spindle-fibres make
their appearance within the nucleus, but no. definite spindle
comparable with that found in Amaba lawesiana is produced.
Moreover, the fibres seem to lie on the nuclear membrane rather
than within the cavity of the nucleus. The chromosomes now
become arranged on the fibres, but I have not discovered any
examples which show all the chromosomes arranged in an
equatorial ring or band. All the stages of this phase of division
show two principal groups of chromatin granules or chromosomes
at each end of the jong axis and other chromosomes irregularly
disposed in the equatorial region. In the latter region the
individual chromosomes are extremely difficult to distinguish,
and it is therefore practically impossible to count them. There
appear, however, to be about eight chromosomes or ehromatin
masses produced within each nucleus, four of which travel to
each end. A description of these stages of division is very
difficult to make owing to the fact that no two stages exactly
agree, as will be seen on referring to figs. 68-70. All of them
are, however, of the same general type, and a detailed description
of each is unnecessary. The chromosomes ultimately become
drawn to the two poles of the nucleus, a stage which is well.
shown in fig. 71.

Shortly after this, or even earlier, the chromosomes begin to
break up and lose their distinct outline. As a result of this,
a granular mass of chromatin, rather triangu'ar in outline, is
produced at each end of the nucleus (figs. 72 & 73). The nucleus
now begins to elongate, and the chromatin is reorganised into
daughter-nuclei, which thus gradually separate further and
further apart. Tn fig. 73 a stage is shown in which a dumb-bell
appearance is presented by the two rounded daughter-nuclei and
the nuclear membrane constricted between them. This is the
latest stage of division that I have obtained. I have failed to
discover any stages showing fission of the animal, and it seems
to be fairly evident that this occurs after nuclear division is
quite complete. In this connection it 1s interesting to note that
I have found a large number of bi-nucleate amcebze on the same
preparation. It is possible that these are forms in which fission
is retarded, or again they may be abnormal individuals, for I
have found one or two tri-nucleate forms as well.

(c) Remarks.

The nuclear division in Ameba agricola differs from that which
occurs in any other amceba. It is obviously a modified mitosis,
but it is not easy to connect it up with any of the numerous
mitotic nuclear divisions which have been described and figured
in other ameebe.

29%

a=

328 DR. T. GOODEY ON

LITERATURE.

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Gén. Expt. 8. 5, T. vi. p. 491.

—— (1911 a).—‘“Sur les Flagellés intestinaux des Poissons
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—— (1912).—‘ Sur la revision du genre Bodo Ehrbg.” Arch.
f. Protist. Bd. xxvi. p. 413.

Brnar, K. (1914).—“ Bau und Vermehrung von Prowazekia
josephi, n. sp.” Arch. f. Protist. Bd. xxxv. p. 103.

Carron, EB. (1910).—“ Protozoaires parasites des branchies des

Labres. Ameba mucicola, ete.” Arch. Zool. Gén. Expt.
Ss), (No We joe Aah .

—— (1910 a).—* Essai sur la structure du noyau et la mitose
chez les Ameebiens. Faits et théories.” Arch. Zool.
Gén. Expt. S. 5, T. v. p. 267.

— (1912). ‘Sur quelques genres d’Amibes libres et parasites.
Synonymies, homonymie, impropriété.” Bull. Soc. Zool.
France, I’. xxxvul. No. 3, p. 109.

Dopett, C. C. (1908).—“The Structure and Life-history of
Copromonas subtilis, n. g. et n. sp.” Quart. Jour. Mic.
Sci. vol. li. p. 75.

— (1909).—* Researches on the intestinal ‘Protozoa of Frogs
and Toads.” Quart Jour. Mic. Sci. vol. li. p. 201.

—— (1914).—“ Cytological Studies on three species of Ameaba :—
A. lacerte Hartmann, A. glebe, n.sp., A. fluvialis, n. sp.”
Arch. f. Protist. Bd. xxxiv. p. 139.

Dorterm, F. (1907).—“‘ Studien zur Naturgeschichte der Proto-

zoen: V., Ambbenstudien.” Arch. f. Protist. Suppl.-Bd.
p. 250.

1911).—‘ Lehrbuch der Protozoenkunde.’ Jena.

Dusarpin, F. (1841).—‘ Histoire Naturelle des Zoophytes In-
fusoires.’ Paris.

Guiser, H. (1912).—“ Untersuchungen uber die Teilung einiger
Amoben.”.. Arch. f. Protist. Bd. xxv. p. 27.

—— (1912 a).—‘‘ Uber Kernteilung, Encystierung und Reifung
von Ameba mira, n. sp.” Arch. f. Protist. Bd. xxvii. -
Be IA.

Goopey, I. (1911).—‘* A Contribution to our Knowledge of the
Protozoa of the Soil.” Proce. Roy. Soc. B, vol. Ixxxiv.
p. 165.

—— (1914).—‘‘ A preliminary Communication on three new
Proteomyxan Rhizopods from Soil.” Arch. f. Protist.
IBals somays Jos 20.

—— (1915). A note on the Remarkable Retention of
Vitality by Protozoa from old stored Soils.” Annals
of Applied Biology, vol. i. Nos. 3 & 4, notes p. 395.

—— (1915).—‘‘ Investigations on Protozoa in relation to the
factor limiting Bacterial Activity in Soil.” Proc. Roy.

. Soe. B, vol. Ixxxvili. p. 437.

SOIL PROTOZOA. 329

Harrmann, M., & Cuaaas, C. (1910).—Flagellaten Studien. Mem.
Inst. Oswaldo Cruz, ii. p. 64.

Hickson, 8. J. (1909).—“ The Mastigophora.” <A Treatise on
Zoology, pt. 1. Fase. i. London.

Kent, W. Savitte (1880-1882)—‘ A Manual of Infusoria.’

London.

Keven, J. (1895).—“ Die Kerntheilung von Euglena viridis
Elrbg.” Zeitschr. f. Wiss. Zool. vol. lx. p. 215.

Kuess, G. (1892).—“ Flagellatenstudien I. and II.” Zeitschr. f.
Wiss. Zool. vol. lv. pp. 265, 353.

Kuczyysxi, M. H. (1914).— ‘Untersuchungen an _ Tricho-
monaden.” Arch. f. Protist. Bd. xxxiii. p. 119.

Liston, W. G., & Martin, C. H. (1911).—‘ Contributions to the
Study of Pathogenic Ameebe from Bombay.” Quart.
Jour. Mic. Sci. vol. lvii. p. 107.

Mackinnon, D. L. (1913).—“Studies on Parasitic Protozoa.
(1) The Flagellate Polymastix and its affinities with
the Trichonymphide.” Quart. Jour. Mic. Sci. vol. lix.
joe “Ue

“Studies on Parasitic Protozoa. (2) A. The En-
eystment of Rhizomastix gracellis Alexeieff. B. Vetra-
trichomastix parisit, n. suabgen. & n. sp.” Quart. Jour.
Mie. Sci. vol. lix. p. 459.

Martin, C. H. (1910).—“ Observations on Vrypanoplasina
congert. Pt. I. Division of Active form.” Quart.
Jour. Mic. Sei. vol. lv. p. 485.

—— (1911).—* A note on the early Stages of Nuclear Division
of the large Ameceba from Liver Abscesses.” Quart.
Jour. Mic. Sci. vol. lvii. p. 279.

—— (1912).—‘‘ A note on the Protozoa from Sick Soils; with
some account of the Life-cycle of a Hlagellate Monad.”
Proc. Roy. Soc. B, vol. Ixxxv. p. 393.

—— (1913).—* Some remarks on the behaviour of the Kineto-
nucleus in the division of Flagellates; with a note on
Prowazekia terricola, a new FElagellate from Sick Soil.”
Zool. Anz. Bd. xli. p. 452.

—— (1913 a).—“ Further observations on the Intestinal Try-
panoplasmas of Fishes, ete.” Quart. Jour. Mic. Sci.
vole alin yp) 17a).

—— & Lewin, K. R. (1914).—“ Some notes on Soil Protozoa.”
Phil. Trans. Roy. Soc. B, vol. cev. p. 77.

——— & Ropserrson, M. (1909).—“ Preliminary note on Trypano-
soma eberthi (Kent), (Spirocheta eberthi Luhe), and
some other Parasitic forms from the Intestine of the
Fowl.” Pree. Roy. Soe. B, vol. 1xxxi. p. 385.

(1911).—* Further observations on the Cxecal Para-
sites of Fowls, etc.” Quart. Jour. Mie. Sci. vol. lvii.
D, OB

MATHIS, C. & Léger, M. (1910).—“ Sur un Flagellé Prowazekia
wetnbergi, n. sp., freqaemment observé dans les selles
de Vhomme.” Bull. Soc. med. chir. de ’ Indochine.

330 DR. T. GOODEY ON

Merciger, L. (1910).—‘ Contribution a l’étude de l’Amibe de la
Blatte (Entameba blatie Biitschli).” Arch. f. Protist.
Bd. xx. p.. 143.

Mincutn, H. A. (1912).—‘ An Introduction to the Study of
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Mororr, Tu. (1904).—“ Beitrag zur Kenntnis einiger Flagel-
laten.” Arch. f. Protist. Bd. ii. p. 143.

Nicter, K. (1909).—“ Entwicklungsgeschichtliche Studien iiber
Amében.” Arch. f. Protist. Bd. xv. p. 1.

—— (1911).—* Prowazekia parva, n. sp., eine weitere freile-
bende Binucleatenform.” Arch. f. Protist. Bd. xxi.
peels

—— (1911 q@).—“Studien iiber Protozoen aus einem Almt-
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id sex xis pL DG:

Pororr, M. (1911).—“ Entwicklungscyclus von Amada minuta,
n. sp.” Arch. f. Protist. Bd. xxii. p. 197.

Prowazex, 8, von. (1903).—“ Flagellatenstudien.” Arch. f.
Protist. Bd. ii. p. 195.

SENN, G. (1900).—“ Flagellata,” in Engler & Prantl’s ‘ Pflanzen-
familien.’ Leipzig.

VauixanerF, K.(1905).—‘“ Beitriige zur Biologie und Entwicklungs-
geschichte von Ameba limax, usw.” Axch. f. Protist.
Bde. paaloie

Wenyon, C. M. (1910).—“ Some observations on a Flagellate of
the genus Cercomonas.” Quart. Jour. Mic. Sci. vol. lv.
p. 241.

-—— (1910 a).—“ A new Flagellate (Macrostoma mesnili, n. sp.)
from the Human Intestine, ete.’ Parasitology, vol. iii.
Tos AL),

Wuerry, W. B. (1913).—‘Studies on the Biology of an Amceba
of the Limax group, Vahlkampfia sp. No. 1.” Arch. f.
Protist. Bd. xxxi. p, 77.

Wirnmore, E. R. (1911).—“ Prowazekia asiatica (syn. Bodo
asiaéicus Castellani & Chalmers).” Arch. f. Protist.
IBele sexs joy BIOs

—— (1911 @).—“‘ Studien itiber Kulturamében aug Manila.”
Arch, f. Protist. Bd. xxiii. p. 81.

EXPLANATION OF THE PLATES.

All the figures are camera lucida drawings, and were made with the aid of
Zeiss 2mm, apochromatie objective and compensating oculars 18 and 12, giving
approximate magnifications of 2786 and 1833 diameters respectively.

Prats TI.
All figures magnified 2786 diameters approximately.
Prowezekia (Bode) saltans,

. Organism seen from the ventral aspect.

. Side view.

. Dorsal view, a small blepharoplast at the base of each flagellum.
. Doubling of the auteyior flagellum.

Hm OS bo

SOIL PROTOZOA. 33a

Figs. 5-8. Early stages in nuclear division.
9-12. Four small chromosomes present in the nucleus.

13-16. Fragmentation of chromosomes and division of the nucleus.

17-20. Hiongation and division of kinetonucleus and constriction of the body
into two daughter-organisms. The new trophonucleiare unrecoguisable
in these stages.

Fig. 21. Two daughter-organisms nearly separated ; new trophonuclei visible.

22. Small recently separated daughter-form.

Prate II.

‘Owing to the exigencies of space in making up the Plate, the full extent of
the two pairs of flagella is shown only in figs. 23 & 26.

Tetramitus spiralis, sp.n. All figures X 2786.

Figs. 23 & 24. Two normal forms, showing the groove.
25 & 26. Showing the relations of flagella, blepharoplasts, rhizoplasts, and
nucleus.
Fig. 27. Showing new flagella arising from anteriorly enlarged blepkaroplasts.
28. The flagella have migrated before the nucleus shows signs of division.
Figs. 29-23. Successive stages in early phases of nuclear division.
31-36. Stages showing four principal chromatin masses within the dividing
nucleus.
37-41. Later stages in nuclear division.

PratE IIT.
Figs. 42-45. Tetramitus spiralis, sp.n. X 2786.

Figs. 42-44. Later stages of division, showing the formation of daughter-organisms.
In fig. 43 there appears to be a production of large vacuoles on the
longitudinal axis of the body.

Fig. 45. The two new organisms have just separated.

Figs. 46-48. Spironema multiciliatum. X 1833.

Fig. 46. A rather small form having § flagella and showing the contractile vacuole
at the beginning of the tail. ‘The extra-karyosomie part of the
nucleus has very small granules on its outer edge.

47. A longer form showing 18 flagella, rather irregularly disposed.
48. A long form showing spiral twist of the anterior part of the body and
contractile vacuole towards posterior end.

Figs. 49-55. Ameda lawesiana, sp.n. X 1833.

Fig. 49. Normal individual of typical limax form, showing feebly staining granules

just inside the nuclear membrane, and a mass of adherent bacteria”
: at posterior end.

Figs. 59-52. Early stages in nuclear division showing disintegration of the karyosome
and the production of a chain of chromatin granules or chromosomes.

Fig. 53. The formation of the spindle at first, having broad ends extending be-
yond the nuclear membrane The chromosomes are arranged in an
equatorial band.

Figs. 54 & 55. The spindle has become pointed at each pole and is placed obliquely
to the horizontal plane; ring of chromosomes.

Prater IV.
Figs. 56-65. Amcoba lawesiana. X 1833.

Fig. 56. Equatorial ring just divided.
Figs. 57 & 58. Elongation of the spindle and separation of the new chromatin
bands.
Fig. 59. Late stage of nuclear division, showing a twisting of the central portion
of the spindle. i
60. Commencement of constriction of the organism.

See ON SOIL PROTOZOA.

Figs. 61-63. Completion of fission and reorganisation of the daughter-nuclei.
64 & 65. Stages in encystation showing the production of deeply staining
granules within the endoplasm. In fig. 64 the endoplasm appears
sharply separated from the ectoplasm.

Figs. 66-74. Amoeba agricola, sp.n. X 2786.

Fig. 66. Normal form.

67. Granulation of the karyosome.

Figs. 68-70. Showing a variety of stages in the arrangement of the chromosomes on
the nuclear spindle. In each case there appear to be two principal
chromosomes at each pole and variously situated chromasomes in the
equatorial region.

Fig. 71. The chromosomes are drawn to each pole.

Figs. 72-74, Granulation of chromosomes and formation of daughter-nuclei.

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ENTOMOSTRACA FROM CEYLON.

ON ENTOMOSTRACA FROM CEYLON. 333

9. On some Fresh-water Entomostraca from Ceylon.

By Rosert Gurnay, M.A., F.Z.S.
[Received February 8, 1916: Read April 4, 1916. ]

(Plates I.-III.* and Text-figure 1.)

INDEX.
GEOGRAPHICAL; Page
An African genus (Oncocypris) in Ceylon ......... 340
SYSTEMATIC :
Diapiomius Widusts; SP. We) aie. -eeeccesy sess eo eee eee RODS
Oncocypris pustulosa, SP. MM. ..cse-cccseeccaees eee -ceees 340
EY SOCY PLC TUOET.GUGs) SP Ms) a. y-)icees sso eeeeeeseeeen OAL

The fresh-water Entomostraca of which an account is given
here were collected by Mr. G. W. Smith during a visit to Ceylon
in September and October, 1907, on his return from Tasmania,
and I desire to express my thanks to him for his kindness in
handing them over to me for examination. The collections
were made in various waters at Colombo, Kandy, Peradeniya,
Mahintele, and Anuradhpura, but I have not thought it
necessary jn most cases to specify precisely in what spot a
particular species was found.

The most complete account that has been published of the
Entomostraca of Ceylon is that given by Daday in 1898; but
there have been several other contributions made to our know-
ledge of them, so that we now know of 94 species or well-marked
varieties inhabiting Ceylon. My. Smith’s collections contain
35 species, of which three are, as I believe, hitherto undescribed,
and several are additions to the list. It is rather remarkable
that, of the four species of Diaptomus here recorded, three have
not before been taken in Ceylon, and it is quite evident that the
fresh-water Entomostraca of Ceylon are very far from being
completely known.

i PAY ELOR@ DAG

CYCLESTHERIA HISLOPI Baird.

Nine specimens, some with developing young in the brood-
pouch, were taken in the Colombo water-supply.

II. CLADOCERA.

DIAPHANOSOMA EXCISUM Sars.

A considerable number of specimens of a species of Diaphano-
soma were found in a tank at Kandy by Lady Horton’s Drive and
also in Colombo Lake, while a few were taken at Anuradhpura.

* For explanation of the Plates see p. 343.

o04 MR. ROBERT GURNEY ON

All belong to the same species, and that, in my opinion, is
D. exsisum, since they agree in all essential respects with Sars’s
description, though somewhat smaller than the size given by him.
Tt is remarkable that the only species of Diaphanosoma recorded
from Ceylon is D. singalensis Daday, which was found by Apstein
to be common in the Colombo Lake from January to September.
All the specimens that I have examined have the ventral shell-
margin markedly reflexed, and cannot possibly be referred to
D. singalensis. It is possible that there is a seasonal alternation
of the two forms. ®

DaPHNIA LUMHOLTZI Sars.
A few young specimens were taken in a tank at Kandy.

CERIODAPHNIA RIGAUDI Richard.

Peradeniya pond; Colombo Lake; Mahintele; Anuradhpura ;
Kandy tank.

CERIODAPHNIA CORNUTA Sars.

Kandy, in a pond by Lady Horton’s Drive and in the tank.

Daday (1898) has expressed, and still maintains, the opinion
that C. cornuta and C. rigaudi ave merely extreme forms of
one species, and a careful examination of the specimens at my
disposal gives much support to such a view. The presence or
absence of a head-spine is, In my specimens, usually correlated
with the possession of a posterior shell-spine, which is slightly
bifurcated or simple respectively; but not only do both rigaudi
and cornuta forms occur together in the same collections, but
also individuals are found which it is almost impossible to assign
to one or the other. The head-spine may be so small as to be
detected with great difficulty, and the shell-spine may show but
the faintest trace of bifurcation. It appears to me that the
rigaudi form may occur alone, but that wherever (in Ceylon)
C. cornuta occurs, there also are found a small number of
individuals which, in the Jack of its distinctive characters,
approach C. rigaudi. One may conclude that the species may
be in fact distinct, but that C. cornuta is very variable and
may approach C. rigaudi in appearance. The cornuta form
described by Stingelin from Java, and having a double head-
spine, must be regarded as merely a variety of the species, since
Daday (1910) has found specimens with double head-spines
together with others of the typical form in Victoria Nyanza.

Mo1na busta Richard.

Kandy ; Mahintele; Anuradhpura. Common.

I take this opportunity of correcting an error in my figure of
this species recently published (1911, pl.u. fig. 1}. In this figure
the reticulations of the ephippium are shown strongly marked
over the egg-space, whereas, though a very faint reticulation can
sometimes be detected, the egg-space usually appears quite

ENTOMOSTRACA FROM CEYLON. 335

unmarked. These faint markings were shown in my drawing
and have become accentuated in the plate.

Macrorurix opiosa Gurney.

Peradeniya pond. Abundant in a plankton collection taken
at night, but much less common in plankton and in weeds during
the day. Also taken at Anuradhpura.

The species bears some resemblance to MV. singalensis, but
differs from it in the form of the upper lip, the arrangement
of cilia, and teeth on the antenna and on the post-abdomen.
While the adult differs considerably from M. triserialis in the
shape of the valves, the young closely resemble it, being of a
pointed pear-shaped form.

MACROTHRIX TRISERIALIS Brady.
Mahintele—Snake’s pool. ‘Two specimens only seen.

LEYDIGIA AUSTRALIS Sars.

Tn a tank at Anuradhpura great numbers of cast shells of a
species of Leydigia were taken, and among these there are two
or three post-abdomina, but none with the terminal claw
attached. However, the shape of the post-abdomen and the
arrangement of the spines, together with the fact that the shell-
valves are not striated, make it certain that the species is
LI. australis, The arrangement of the spines is distinctive
in this species; there is a regular series, decreasing in size
anteriorly, of groups containing one long spine and two very
short ones. and these groups are not replaced by groups of cilia
till the anal depression is reached.

At Mahintele a single specimen of a Leydigia was taken which
agrees most closely with ZL. australis var. ceylonica Daday. In
this specimen the spines of the post-abdomen are more slender
and partly arranged in fours (Pl. I. fig. 1), and the shape of the
post-abdomen recalls that of Z. propingua. My specimen, which
is much decayed, differs from Daday’s form in absence of sculpture
on the shell.

ALONA RECTANGULA Sars.

A few specimens of a small form of this species were taken in
Peradeniya pond. Some females are ephippial, but no males
were seen. In some specimens the upper hp has a minute tooth
on its anterior margin.

ALONA INTERMEDIA Sars.
Old tank at Anuradhpura.

ALONELLA DAvIDI Richard.

One specimen and a moulted shell were included in a collection
from the old tank at Anuradhpura. They agree exactly with
regard to form of post-abdomen and arrangement of spines and

336 MR. ROBERT GURNEY ON

cilia with the specimens described by Stingelin (1904) from Java
and Honolulu, except that here the claws have no cilia. But in
my specimens the shell is distinctly striated, but without reticu-
lations, thus approaching more nearly to Richard’s description.

I have already (1911) given reasons for regarding 4. davidi
and A. punctata Daday as varieties of A. diaphana King, but I
am inclined to think that, though I still believe the three species
to form a gradational series, yet it is perhaps more convenient
and less cumbrous to leave the three names to define the three
varieties.

ALONELLA KARUA King.

Peradeniya and Anuradhpura.

ALONELLA EXcIsA Fischer.
Peradeniya pond.

Cuyborus PARvus Daday. (PI. I. figs. 2, 3.)

Peradeniya pond and Priest’s Well; Kandy; Anuradhpura.

In 1898 Daday described, under the name of C. sphericus var.
pareus, a Chydorus from Colombo Lake, differing chiefly from
O. sphericus in the form of the upper lip (fig. 2), and I have
little hesitation in referring to this species a Chydorus which is
common in some of Mr. Smith’s collections. These specimens,
which vary from 28 to 35 mm. in length, are nearly globular in
shape and usually of an opaque golden-yellow colour. In the
form of the upper lip and of the post-abdomen (fig. 3) they agree
very closely with Daday’s deseription, but they differ in the fact
that I have been unable to detect with certainty any sculpture
on the shell, whereas Daday’s species shows strong reticulation.

Cuyporus BARrorsi Richard. (Pl. I. figs. 4, 5.)

Peradeniya pond ; Anuradhpura.

My specimens unite in a very perplexing way the characters
of C. barroisi and C. poppet Richard. All my specimens agree in
having the upper lip strongly serrated and in the arrangement
of spines on the post-abdomen (fig. 5), and nearly all have the
shell-valves smooth. The majority have also a tooth at the
posterior ventral angle of the shell (fig. 4) and an additional
minute spine at the base of the caudal claws, but I have also
seen specimens which lack the one or the other. In the structure
of the post-abdomen they agree much more closely with C. poppet
than with C. barroist. C. hybridus Daday unites in the same
way the characters of the two species, and it seems to me that
the three are not, in fact, spectfically distinguishable. O. poppet
has been recorded only from South America and from the
Southern United States, but C. barroisi occurs in Syria, Ceylon,
various parts of Malaysia, New Zealand, Hast Africa, and South
America. Stingelin (1913) also expresses the opinion that the
three above-named species should probably be united into one. -

ENTOMOSTRACA FROM CEYLON, Som

IIJ. COPEPODA.

Cyciops pistincrus Richard. (PI. I. fig. 6.).

In a collection from a’ pond by Lady Horton’s Drive at Kandy
a few specimens of a Cyclops were found, which I assign with
some hesitation to this species. The specimens included only a
single adult female, the rest being chiefly adult males.

My specimens differ in some important respects from typical
European specimens with which I have compared them :—

(1) In size. Length: 9, 144 mm.; 3, ‘95mm. Whereas
English specimens measure about 2 mm. ( 2 ).

(2) Furcal rami. Wheveas in typical C. distinctus the rami
are about 24 times longer than they are broad, in
specimens from Kandy they are less than twice the
breadth. Also in the latter the lateral seta is very
long—longer than the ramus. The proportional length
of the remaining sete is the same in both.

Since in other respects—as, for example, in the form of the
fourth pair of legs and their uniting lamella (fig. 6)—the agree-
ment is complete, I do not think the differences are sufficient to
constitute a distinct species.

CYCLoPs HYALINUS Rehberg.
Kandy ; Mahintele; Anuradhpura.

CYCLOPS LEUCKARTI Claus.

Peradeniya; Kandy ; Anuradhpura.

CYCLOPS VARICANS Sars.

Peradeniya; Kandy; Anuradhpura, Rare.

Cyciors varius Lillj., var. proximus Lill}.

A few specimens taken in the Priest’s Well at Peradeniya.

CycnLops PRASINUS Fischer.
Peradeniya—Priest’s Well; Kandy—Lady Horton’s Drive

pond.

CANTHOCAMPTUS GRANDIDIERI Guerne & Richard, var. (Pls. I.
& II. figs. 7-9.)

In collections from Peradeniya and from Anuradhpura a few
specimens of a species of Canthocamptus were found which I find
difticulty in identifying. In most respects they agree very closely
with descriptions given of C. grandidieri, bat with regard to the
furca and to certain details of the fifth pair of legs there are
differences which are constant and considerable. The fureal
rami are more or less quadrangular, bearing at their apex a short
slender seta on the inner angle and a single very long seta with

338 MR. ROBERT GURNEY ON

a rather swollen base (fig. 7); but in place of the usual external
long seta there is, in all adults, merely a small finger-like
outgrowth. On the other hand, in all immature specimens two
sete are present of the usual form. A further difference is that
the ramus is not, as 1s usual in C. grandidiert, distinctly produced
dorsally. There is indeed a slight overhang, but it is scarcely
noticeable (fig. 9). With regard to the fifth pair of legs the
agreement is closer, but here the innermost spine of the basal
joint is very much shorter than the others, and the second joint
is smooth on its inner face (fig. 8). In the proportional length
of the spines on this leg my specimens agree more nearly with
C. laciniatus Van Douwe, which itself seems to me only a variety
of ©. grandidiert. Brady’s description of C. cingalensis is too
incomplete to make any satisfactory comparison possible.

Drapromus DorIArI Richard.

Anuradhpura. Rare.

DIAPTOMUS STRIGILIPES Gurney.
Anuradhpura—Baltring tank; Mahintele. Abundant.

Diapromus ANN Apstein. (PI. IT. fig. 10.)

Peradeniya pond; Kandy ; Colombo Lake.

Brehm has found that specimens from Kandy do not agree in
all respects with the description given by Apstein, the third joint
of the exopodite of ‘the fifth foot in the female being clearly
distinct, and argues that the separation or fusion of this joint
is therefore not of great systematic importance. In all my
specimens this joint is separate, and Apstein has found that
it is, in fact, distinct in his original specimens, The endopodite
of this leg in my specimens, as in Brehm/’s, is considerably shorter
than the first joint of the exopodite, and, in respect of length,
this joint seems to be very variable. I have seen one specimen
(fig. 10) in which this branch was clearly two-jointed, but this is
evidently an abnormality.

DIAProMUS VibDUUS, sp. n. (PI. II. figs. 11-14.) .

In a collection from the Snake’s pool at Mahintele, among
large numbers of D. strigilipes, a single male Diaptomus was
found which appears to belong to an undescribed species. In
spite of very careful search I have found only the single
specimen, but it seems to me so distinct that it is best to
describe and name it.

The body is slender and tapering anteriorly ; last segment of
the thorax with pronounced posterior angles bearing two small
spines on either side (fig. 11). Fourth abdominal segment
asymmetrical, being slightly swollen on the right side. Furcal
rami and sete of normal shape. The left antenna reaches, when
reflexed, to the end of the fourth abdominal segment. ‘The

ENTOMOSTRACA FROM CEYLON. 339

antepenultimate joint of the right antenna has a hyaline mem-
brane and a short outwardly-turned hook (fig. 12). The fifth
leg (fig. 13) on the right side has two small hyaline processes on
the second basal joint ; the endopodite is longer than the first
joint of the exopodite. The first joint of the exopodite is pro-
duced laterally into a pointed process and bears on its posterior
face a large hyaline process (fig. 14). The second joint of the
exopodite has the spine, which is usually lateral and distal in
position, inserted on the posterior face near the base of the joint.
Length 1:75 mm.

IV. OSTRACODA.

Noropromas ocunata Sars. (Pl. IIT. fig. 15.)

Tank by Lady Horton’s Drive.

This species differs very little from WV. entzi Daday, being
somewhat smaller and with different markings. In dorsal view
it is seen that V. enézi is much wider behind than in front,
whereas in JV. oculata the outline is an almost regular oval

(fig. 15).
STRANDESIA (CYPRIS) VITTATA Sars.

Common in Peradeniya pond.

My specimens agree in all respects with Sars’s description,
with the exception that they do not show the coloured bands
mentioned by him, the absence of which is possibly due to the
preservative. The curious “lop-sided” appearance in end view
is very characteristic. Previously recorded from Puching, China.

EvurycyPRIS SUBGLOBOSA Sowevby.
Colombo Lake. Rare.

SLENOCYPRIS MALCOLMSONI Brady.
Colombo Lake.

CYPRICERCUS RETICULATUS Daday ?

A considerable number of specimens of a species of Oypricercus
were taken in Peradeniya pond. Unfortunately all appear to be
immature, not exceeding -7 mm. in length and with the ovary
barely distinguishable. My specimens differ somewhat from
C. reticulatus in shape and also in the complete absence of any
shell-sculpture, but I cannot assign them to any other species
nor safely describe them as new.

Cypripopsis NEwront Brady & Robertson. (PI. IT. fig. 16.)
Syn. C. aldabre Miller.

Great quantities of this species were found in Colombo Lake
and a few specimens at Mahintele. Apstein has recorded Oando-
nella albida Vavra from Colombo Lake, but I have found no

340 ' MR. ROBERT GURNEY ON

specimens which can be referred to that species. On the other
hand, the shape of the shell (fig. 16) points unmistakably to the
closely allied species C. aldabre, with which my specimens agree
also in other respects. Unfortunately I have not been able to
find a single male, so that the comparison is incomplete.

ONCOCYPRIS PUSTULOSA, sp.n. (PI. IIT. figs. 17-21.)

Seen from above the animal is pear-shaped, very broad behind
and. tapering anteriorly, with a constriction in front of the eye
(fig. 17). In quite young individuals the shape is that of an
egg, with the greatest width just behind the middle and tapering
evenly in front and behind. Seen from the side the two valves
are alike, kidney-shaped, broader in front than behind (fig. 18).
The cuticular border is broad anteriorly, scarcely visible ventrally,
and narrow behind. Seen from inside, the structure of the shell
is distinctive. In the left valve the anterior cuticular border is
very broad and springs from the edge of the shell, its point of
origin being marked by the accompanying sete. Beyond this
point is the ‘“ pore-canal” zone, in which are seen a series of
strongly marked semicircular loops which appear to indicate
unbranched pore-canals, but their real nature is not at all clear
(fig. 19). Cutting across them is seen a conspicuous ridge. The
structure is much the same posteriorly and in the right shell.

The surface of the shell is thickly covered with little knobs,
but in very young specimens it is strongly reticulated and pitted.
The eyes are very large and united. The general colour, in
spirit, is golden yellow with a conspicuous greenish pigment-
spot on either side of the eye.

The maxilla has no respiratory plate, but in place of it there
is a single small seta. Mliiller gives the entire absence of setz as
an important character of the genus, and it is possible that the
specimen in which I have seen the seta is abnormal in this
respect. From the small number of specimens at my disposal
T am unable to go further into the matter. In the second
maxilla there are two strong spines on the third lobe, one of
which is toothed and the other smooth. In the first leg the
third and fourth joints are fused; the last joint bears a very
large curved spine and a single short seta (fig. 20). The second
leg has a well-developed terminal joint bearing a long curved
claw, a seta of about the same length, and a small hair (fig. 21).
The furea is a simple flagellum.

- Length -5—58 mm.; width 43-54 mm.; height :3--38 mm.

A very few female specimens of this species were taken ina
tank by Lady Horton's Drive at Kandy. The genus Oncocypris
was established by G. W. Miiller for a species, 0. vodtzkow?, from
Madagascar, which has since been found also in Abyssinia (Daday)
and South Africa (Brady). The only other species of the genus
is O. costata Daday from German Hast Africa. It is therefore of
considerable interest to find that a species of this African genus
occurs also in Ceylon.

ENTOMOSTRACA FROM CEYLON. 34]

PHYSOCYPRIA TUBERATA, Sp.n. (PI. III. fig. 22; text-fig. 1.)

Female. Seen laterally the shell is oval, the greatest height
equal to two-thirds the length and falling behind the middle
(text-fig. lw). The anterior end is much less broad than the
posterior end. The right and left shells are of the same shape,
the left shell slightly the larger and overlapping the right in
front. Both shells have a narrow hyaline border and are fringed

Text-figure 1.

Physocypria tuberata.

a. Female. Left valve, X 66.

6b. Male. Right valve from inside, X 126.

e. Female. Furcal ramus, X 274.

d. Male. Copulatory organ, X 274.

650 P35 Clasping organ of right side, X 274,

‘with long scattered sete. ‘The right shell differs from the left

in having a row of small knobs on the anterior and posterior

margins (text-fig. 1b). Seen from above the shell is much com-

pressed and narrower in front than behind (PI. III. fig. 22).

‘The;surface of the shell is smooth, but marked with small brown
Proc. Zoot. Sec.—1916, No. XXITI. 23

342, MR, ROBERT GURNEY ON

spots. In the second leg the penultimate joint is three and a
half times as long as the last joint and without cilia. The last.
joint bears two subequal claws and a recurved seta longer than
the last three joints of the leg. The furcal rami are slightly
curved and bear two short stout claws and a very short seta at
the apex. The dorsal seta is inserted about the middle, and is
nearly half as long as the furca and about the same length as
the longest claw (text-fig. 1c). Length -45—-5 mm.; height
‘27-28 mm.

Male. The male differs somewhat from the female, the dorsal
margin of the shell being more flattened and the two ends more
equally rounded. The tubercles of the right shell are conspicuous.
The palp of the right second maxilla is slender, broader at the
end, and with a triangular pointed process (text-fig. 1e); the last
joint is in the form of a curved blunt-ended claw. The palp of
the left side is more or less cylindrical, not dilated at the end,
and with a minute tooth in place of the triangular process of the
right side. The distal joint forms a curved sharp-pointed claw.
The copulatory apparatus consists of a large triangular lamella
with pointed recurved end, and a narrow pointed process hinged
to it (text-fig. 1d). The ejaculatory apparatus has six rings of
spines.

A few specimens only of this species were found in a collection
from Colombo Lake, and amongst them was a single male.

The species resembles Cypria crenulata Sars very closely, and,
indeed, may be but a variety of it, but it differs from it in having
knobs on both the anterior and posterior margins of the right
shell, and also in the presence of brown spots on the shell.
Sars gives no particulars of the structure of the male by which
a comparison could be made, but he says that the male is exactly
like the female in shape, and that is not quite the case with
regard to my own specimens.

Literature.

Apsretn, C.—‘‘ Das Plankton im Colombo-See auf Ceylon.”
Zool. Jahrb., Abt. Syst. xxv. 1907, pp. 201-244.

Apstrein, C.—‘* Das Plankton des Gregory-Sees auf Ceylon.”
Zool. Jahrb., Abt. Syst. xxix. 1910, pp. 661-680.

Brapy, G. 8.— Notes on Entomostraca collected by Mr. Haly in
Ceylon.” Journ. Linn. Soce., Zool. xix. 1886, p. 293.

Breumu, V.—‘‘ Ueber die Mikrofauna chinesischer und siidasiat-
ischer Siisswasserbecken.” Arch. f. Hydrobiol. u. Plank-
tonk. iv. 1909, pp. 207-224.

Dapay, E. von.—‘ Mikroskopische Siisswasserthiere aus Ceylon.”
Termés. Fuzetek, Anhangsheft zum xxi. Bd. 1898.
Danvay, E. von.—‘ Untersuchungen iiber die Sitisswasser-Mikro-
fauna Deutsch-Ost-Afrikas.” Zoologica, Bd. xxiii. Heft 59,

1910.

ENTOMOSTRACA FROM CEYLON, 343

GurNEy, R.—‘“ On Two new Entomostraca from Ceylon.”
Spolia Zeylanica, iv. 1907, pp. 126-134.

Gurney, R.—“ On some Fresh-water Entomostraca from Egypt
and the Soudan.” Ann, & Mag. Nat. Hist. (8) vii. 1911,
p. 25.

Popper, 8. A., and Mrizex, A.—‘‘ Die von Herrn Dr. H. Driesch
auf Ceylon gesammelten Siisswasser - Entomostraken.”
Beih. zum Jahrb. d. Hamb. Wiss. Anst. xii. 1895.

StincELIN, TuH.—‘‘ Untersuchungen tiber der Cladoceren-fauna
von Hinterindien, Sumatra und Java.” Zool. Jahrb.,
Abt. Syst. xxi. 1904, pp. 327-370.

STINGELIN, TH.—‘‘ Voyage d’Exploration Scientifique en Colom-
bie.” Cladocera. Mém. Soc. Neuch. Se. nat. v. 1913.

EXPLANATION OF THE PLATES.

PrateE I.

Fig.1. Leydigia australis var. ceylonica. Post-abdomen. X 260.
2. Chydorus parvus. Upper lip. X 445.

3} a Fe Post-abdomen. > 1050.

4, 4 barroisi. Female. X 260.

5. 3 L Post-abdomen. XX 1050.

6. Cyclops distinctus. Uniting lamella of fourth pair of legs of male.
x 445.

7, Canthocamptus grandidieri. Furcal rami. X 445.

8. 53 z Fifth leg of female. XX 445,
Puate II.

Fig. 9. Canthocamptus grandidieri. Furcal ramus, side view. X 445. _
10. Diaptomus anne. Fifth leg of female showing (abnormal) 2-jomted
endopodite. X 445.

if, 5 viduus, sp.n. Male. X 58.
12. 34 5 FA Prehensile antenna. XX 120.
13. i 35 ES Fifth pair of legs. XX 120.
14. 3 - Ss Part of the right leg of the fifth pair.
X 320,
Prate III.

Fig. 15. Notodromas oculata. Dorsal view of female. X 98.
16. Cypridopsis newtoni. Right valve of female. 120.
17. Oncocypris pustulosa, sp. u. Dorsal view of moulted shell. X 150,

18. 3 45 3 Side view. XX 98.

19. 3 # Anterior end of left valve from inside.
< 260.

20. a 3 3 First leg. x 445.

21, Second leg. X 445.

b) 33 39
22. Physocypria tuberata, sp.n. Dorsal view of female. X 98.

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MR. G. A. BOULENGER ON THE BOLTI. 345

10. On Specimens of the Perciform Fish Tilapia nilotica
with increased number of anal spines. By G. A.

Bovtencer, F.R.S., F.Z.8.*
[Received February 1, 1916: Read April 4, 1916.]

INDEX.
VARIATION ; Page

Variation in the number of anal spines in Tilapia
milotica and other Cichlide .......................:.. 845
Taxonomy:
On the value of the number of anal spines for the
division of the Cichlid into genera ............... 345
SYSTEMATIC:
Tilapia nilotica athiensis, var. u., E. Africa

In the Cichlide, as in most Perciform Acanthopterygians,
three is the most frequent number of spines in the anal fin, and
this number may be looked upon asa primitive character. It
has generally been the custom for systematists to attach generic
‘importance to an increase in the number of these spines, even
when unaccompanied by any other character. In this I have
differed, and refused to accept genera based on the number of
anal spines when everything else pointed to close relationship
with species showing the usual number, thus uniting Giinther’s
Oreochromis (4 anal spines) with Tilapia and Pellegrin’s Asta-
toreochromis (4 to 6 anal spines) with Haplochromis. I felt all
the more justified in doing so from the fact that occasionally, as
individual exceptions, the three spines may be increased to four,
as in Zilapia mossambica, variabilis, percivali, Haplochromis
desfontainesti. —There is also the perplexing case of Cyrtocara
moorit, of which only two examples are known, one with three
anal spines, the other with four. My reform in classification
has not met with the approval of Dr. Pellegrin, who has
protested against the suppression of his genus Astatoreochromis,
on the ground that the same character has been used for distin-
guishing American genera— with what regard to natural
affinities appears to me questionable. I think the following fact
disposes once for all of his objection. .

It is with the greatest surprise that, on recently receiving
from Mr.S. lL. Hinde a series of over 30 specimens of a fish
which I identified as the common Tilapia iilotica, a species with
which I am familiar from a study of hundreds of specimens, the
first I took up showed five anal spines, and the others either
four or five. A table showing the variation in 30 of these
specimens is here appended. This series was obtained in the
Makindu and Isavo Rivers, affluents of the Athi River in
British East Africa.

* Published by permission of the Trustees of the British Museum.

MR. G. A. BOULENGER ON

346

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THE BOLTI. 347

Having previously received, after writing the description of
Tilapia nilotica for the ‘Catalogue of African Fresh-water
Fishes,’ two young specimens from another affluent of the same
river, the Simba River, it occurred to me to examine them
carefully, as I should have done before, and I found four to be
the number of spines in both. In their physiognomy, in their
coloration and markings, and in all structural particulars, these
fishes are indistinguishable from Tilapia nilotica; and although,
in view of the constancy of the increased number of anal spines,
the Athi River specimens may be recognised as a new local form,
under the name of var. athiensis, I should not think of proposing
for them a new species.

A further remarkable fact is the presence of four anal spines
in another Tilapia very closely related to, though sufficiently
distinct from, 7’. nilotica, viz. T. (Oreochromis) nigra Gthr., also
from the Athi basin. Why in the Tilapia from this river-system
an increase should have taken place in the number of anal spines
is difficult of explanation, unless it be that an abnormal trans-
formation of a soft ray into a spine, as happens elsewhere, should
have been a peculiarity of the first settlers in that basin of the
widely distributed 7’. nilotica, and, becoming fixed, been passed
on to 7’. nigra, which may well be regarded as derived from that
species. Whatever this explanation be worth, the fact is clear
that, unless our classification of the Cichlide be made still more
artificial than it unfortunately is at present, the number of anal
spines must not be used, as a single character, for the division
into genera, and it affords the best justification that could be
wished for the course I have followed in the past.

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ON THE EXTERNAL CHARACTERS OF MONGOOSES. 349°

11. On the External Characters of the Mongooses (Mun-
goudee): by ik, Te Pococw,. Hho. Pls. :Z.S.,

Curator of Mammals.
[Received February 22, 1916: Read April 18, 1916.1

(Text-figures 1-10.)

INDEX. Pace

The Har, Vibrisss and Rhinarium):..::........-----....= ool

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Introduction.

The facts recorded in this paper are based mainly upon an
examination in the Society’s Prosectorium of examples of the
following genera and species, which have been exhibited during
the past ten years in the Zoological Gardens :—

Mungos mungo, the common Indian Mongoose; many spe-
cimens of both sexes.

- smithii ; one female example from Ceylon.

3 auropunctatus ; two examples from Nepal and Chitta-
gone.

i brachyurus; one example from the Malay Peninsula,
only superficially examined, without drawings being
made.

Myon Hm gracilis ; one unlocalised example of this African form.
5 Helogale undulata; two examples from British Hast Africa
very closely allied to this species, but with less yellow in the
fur.
Ichneumia albicauda ; one female from the White Nile.
Atilawx paludinosus ; one male from South Africa.
Cynictis penicillata ; one example from South Africa.
Ariela fusciata ; one female example from the Sudan, repre-
senting a local race of this species.
Crossarchus obscurus; two examples, male and female, from
West Africa.
Suricata suricatta; two examples, male and female, from
South Africa.

I have also seen, in addition to a few examples of some of the
species above enumerated, a female specimen of Ldeogale puisa,
ticketed Zanzibar (Sir J. Kirk), preserved in alcohol in the
British Museum.

Although. a study of the skulls and teeth has been no part of
my present purpose, I have made use of the characters they

350 MR. R. I. POCOCK ON THE

supply, in conjunction with the external features, in the attempt
to discover the probable affinities of the genera with reference to
a hypothetical archaic type of Mongoose. For this, the fine
series of skulls in the British Museum has been indispensable.

There does not appear to have been any general work on the
classification of Mongooses since the publication of Mr. Thomas’s
and Prof. Mivart’s papers in our ‘ Proceedings’ for 1882. The
classification proposed by Mr. Thomas, and the genera he pre-
served, have been adopted by subsequent authors. It must be
remembered, however, that his paper was written before the
introduction of the more refined and modern methods of distin-
guishing genera. Hence there is no doubt that he would now
agree in assigning generic rank to Jchnewmia, to which at the
time in question he gave subgeneric rank under Herpestes, now
known as JZungos. I am not aware, however, that there has
been published in any faunistic lists a proposal, either tacitly or
definitely expressed, to restore Atilaw and Ariela to the generic
status formerly given them respectively by Cuvier and Gray.
On the available material, Mr. Thomas made Atilaw a synonym of
Herpestes (= ungos) and Ariela a synonym of C' rossarchus. My
reasons for restoring these names to generic rank are given in the
sequel. About A lee there can, I think, be no doubt, assuming
the constancy of the features relied upon; and the only criticism,
it seems to me, that can be made against the severance of Ariela
from Crossarchus is the uncertainty, in the absence of fresh
material, as to the correct generic allocation of all the forms
that in recent years have been described as Crossarchus. This,
however, does not deprive of their force the characters by which
the type-species of Aviela can be distinguished from that of
Crossarchus.

As regards the generic names previously proposed, it must not
be forgotten that Gray and Hodgson divided what is now known
as Mungos into several genera—e. g., Urva, Teniogale, Galerella,
etc.,—the type-species of which were cited by Thomas in 1882.
Tt remains to be seen whether any of these genera will be restored
in the future or not*. I have not sufficient material upon which
to form an opinion of any value; but as at present constituted,
Mungos is the only genus of Viverroid Carnivores common to
the Oriental and Ethiopian Regions.

Two new generic names have been introduced since Thomas’s
paper, namely, Paracynictis for Cynictis selousi, and Galeriscus
based upon G. jacksoni, an alleged Musteline from British East
Africa, which proves to be a species of Bdeogale t.

* The small African Mongoose (Mungos gracilis), the type of Galerella Gray,
differs in several respects from Mongooses of the M. mungo and M. ichnewmon type,
notably in its larger ears, less webbed feet, and in the prominence of the anterior
chamber of the tympanic bulla. In all these respects it approaches the otherwise
very distinct genus Cynictis. Another small African species, IM. pulverulentus,
ye resemble I. gracilis tolerably closely, so far as can be judged from dried
materia

+ Ann. Mag. Nat. Hist. (8) xvii. pp. 176-179, 1916.

EXTERNAL CHARACTERS OF MONGOOSES. 351

The Ear, Vibrisse and Rhinariwm.

The Har.—The only description of the ear of Mongooses with
which I am acquainted is that of Boas (‘ Ohrknorpel und dusseres
Ohr der Siiugetiere,’ p. 140, pl. xxi. fig. 222, Kopenhagen, 1912),
who examined this organ in the common Indian species Herpestes
griseus (= Mungos mungo).

The ear differs in the following particulars from that of all the
genera of Viverride discussed in my previous papers :—(1) The
marginal bursa is absent ; (2) the supratragus or plica principalis
is converted into a large movable laminate flap; (3) above the
supratragus there is a similar but smaller flap; (4) the antero-
internal ridge curves abruptly backwards into the cavity of the
ear, its inferior prominence being set high up and fitting into a
hollow above the antitragus. By the disposition of these ridges
the cavity of the ear is capable of being very completely closed
when the ear is folded. The superior flap closes over the space
above the supratragus, the latter similarly shuts down upon the
antero-internal ridge, and the prominence of the latter fits into
the space above the antitragus, which is itself applied to the ridge
representing the tragus.

In nearly all the genera the ears are set well behind the eye,
are irregularly semicircular in shape, and small, so that the upper
margin hardly projects above the line of the occiput and of the nape
of the neck; but in Cynictis Hina are much larger, project well
above the head, and have the antero-superior rim rising only a
little behind a point above the posterior angle of the eye. In its
structural details the ear in this genus is of the same general
type as that seen in Mungos, Crossarchus, Bdeogale, and others,
with the exception that there is a small shallow pocket behind
the antitragus (text-fig. 3, A, C). I have not observed this in
any other genus, but it is no doubt the homologue of the similarly
situated depression in the Hyznas, which was regarded by Boas
as the representative of the marginal bursa in other Atluroid
Carnivores.

It is perhaps significant that this remnant of the bursa
persists in the genus which of all the Mongooses has the longest
and broadest ears and approaches, in that respect at least, nearest
to the Hyenas. Nevertheless, the ear of the Hyznas, except
for the abnormal position and structure of the bursa, resembles,
broadly speaking, that of other ~Ailuroidea*. Some species of
Mungos, e.g. M. gracilis (text-fig. 1, A), have much larger ears
than species like I. mungo and MW. smithtii. In Ichneumia albi-
cauda (text-fig. 1, D) they are also tolerably large; whereas in
Atilax paludinosus (text-fig. 1, C), Ariela fasciata and Cross-
archus obscurus (text-fig. 2, A, B) they are comparatively small
and rounded. Nevertheless, whatever their size may be, the
ears conform closely to the type described in Mungos nwungo and
M. smithii.

* See Ann. Mag. Nat. Hist. (8) xvii. p. 333, 1916.

352 MR. R. I. POCOCK ON THE

One genus, Suricata, stands quite apart from the others in the
structure of the ear (text-fig. 3, B, D). This organ is small,

Text-figure 1.

Mo NIA wy de 17,
veel

=

eZ

tf:
Os

\
(iT

if

by eee >
aay SS _
HWE TIS SS
eat

Ct

A. Head of Mungos gracilis, from a spirit-specimen, with ear open.
B. 3 Helogale undulata, from a fresh specimen, ,, Ue
Cc. és Atiiax paludinosus, an 4 53 we
D. a Ichnewnia albicauda, a A 3

29

(All 3 natural size.)

EXTERNAL CHARACTERS OF MONGOOSES. 353

semicircular, and set well back and low down on the head, but it
is of a much simpler type than in other genera. There is no
lamina above the supratragus, and the latter isa simple thickened
ridge. In other respects the ear resembles that of the rest of the
genera of Mongooses, differing from the ear of all the Viverride
in the absence of the bursa and the high position of the pro-
minence of the antero-internal ridge above the antitragus.
Amongst the Viverride, the Galidictine* genera are those
which in the structure of the ear come nearest to Suricata.

Text-figure De

A. Head of Ariela fasciata, from a fresh specimen, with ear closed.
B. a Crossarchus obscurus, 4, ss os i

(Both 3 natural size.)

The ear of Suricata, in spite of the absence of the two movable
lamine, is capable of being closed as tightly as in other members
of this group, the supratragal ridge assuming an oblique direction
and being pressed against the antero-internal ridge when the ear
is folded. That being so, it is difficult to understand the reason
for the development of the two lamine in the typical Mongooses.
I think, however, it is probable that the close folding of the ear
is an adaptation to the known burrowing habits of this group,

* Ann. Mag. Nat. Hist. (8) xvi. p. 354, pl. xv. fig. 4, 1915.

354 MR, R. I, POCOCK ON THE.

since the Civets, Genets, and Palm Civets, with normal ears, are
either scansorial or terrestrial, but not fossorial *. In that case,

Text-figure 3.

A. Head of Cynictis penicillata, from a spirit-specimen, with ear open. X 3.
1B} » Suricata suricatta, 55 es 5 Fy be
C. Ear of Wungos smithii, open.

D. ,, Suricata suricatta, closed.

BR. ,, Cynictis penicillata, closed.

v., upper valvular lamina; s., supratragus (plica principalis), forming lower
valvular lamina; 6., bursa; a.é., antero-internal, a.e., antero-external,
p.e., postero-external ridges.

it seems to me possible that the method of closing the ear by
means of two lamine, as above described, may serve the purpose

* Not ascertained in the case of the Galidictinee.

EXTERNAL CHARACTERS OF MONGOOSES. 355.

of excluding dust and dirt without at the same time excluding
sound-vibrations to the same extent as does the arrangement for
folding seen in Swricata.

Facial Vibrisse.—In number and disposition the tufts of
vibrissee are quite normal and agree with those of the Viverride.
But they vary a good deal in development in different forms.
Broadly speaking, they are longer and more numerous in smaller
than in larger species, as may be seen by comparing J/wngos
gracilis with M. mungo or Ichneumia albicauda with Ariela
fasciata and Crossarchus obscurus. For instance, the upper
genal tuft consists of about three bristles in J, gracilis and
Ariela fasciata, and generally at all events of only one, which is
not always detectable, in J/. mungo and Ichnewmia.

An exception to this generalisation is seen in Atilax palu-
dinosus, one of the largest members of the group. The normal
vibrisse are long, and the anterior mystacials and the submentals
are unusually copious and long, giving to the muzzle a hirsute
appearance not seen in other species where these particular
bristles are comparatively poorly developed (text-figs. 1, 2, & 3).

Rhinarium.—This organ presents no features by which it can
be distinguished from that of all genera of Viverride. Within
the group of the Mongooses it is tolerably constant in form. Its
upper anterior margin, viewed from above, is evenly convex from
side to side and not mesially notched or sulcate; from the front
it is nearly straight or lightly convex, with rounded angles. On
the upper surface the narial slits converge slightly, or somewhat
markedly, as in Atilax, and are bordered externally by a narrow
rim of naked integument ; the posterior border is usually lightly
concave, but in Atilaw the hairs of the muzzle overgr it
to a greater extent, forming an angular excision in the naked
skin. The infranarial portion of the anterior surface is always
well developed, and generally transverse from side to side along
the lower margin; but in Atidlax paludinosus, in which the
whole rhinarium is broad, the infranarial portion is especially
deep and its edges diverge a little upwards and outwards from
the middle line. The median sulcus, with which the anterior
surface is marked in some forms, never apparently passes higher
than the upper edge of the nostrils.

As has been pointed out by Gray, Thomas, and others, the
inferior edge of the rhinarium may or may not be continued
down the middle line of the upper lip as a strip of naked skin.
This strip is present in the genera Wungos, Helogale, Ichneumia,
Atilax, Bdeogale, and Cynictis*, and absent in Rhynchogale +,
Ariela, Crossarchus, and Suricata, in which the skin of the
upper lip is continuously hairy across the middle line. When
this strip is present, it is always grooved, and in a great majority

* Tn the ‘fauna of South Africa, Mamm. i. p. 73, 1900, Mr. W. L. Sclater places
Cynictis, with Suricata and Crossarchus (=Ariela), in the category in which the.
lip is undivided. This is an error.

+ I have never seen a fresh or alcohol-preserved adult example of this genus.

St
lor)

MR. R. I. POCOCK ON THE

Text-figure 4.

WAU;
Oe

A>» Arh UY
NEST PON
Wie, Uf far oe PR
fii Mis S/N

Jiepebtens

Maa) ’
Mayr ris
OT fem

yal SSeS

a je we Ss

Z

Vy, \\

LT PWN

Yo Ti SA
AY 0~ WS

rin Sie
ye sate °
WOGAISSR
MD ANA VX
WY Ys vy
5
4, \ |
yi

A. Rhinarium and upper lip of Atilax paludinosus, from the front.
B.

” 2” 29 ”? ” from above.
C. 3 A Me Tchneumia albicauda, from the front.
D. 0 5 66 5 2 from above.
E. 5 3 8 Mungos smithii, from the front.
=
F. ) » ” $3 PD from above.
aN u .
G. » » 5p oy 7 from the side.
H. a A 55 Suricata suricatta, from the front.
I. % » 5 Ariela fasciata, we
K. a % Crossarchus obscurus, i

(All natural size.)

EXTERNAL CHARACLERS OF MONGOOSES. 357

of cases the groove is continued upwards on to the rhinarium.
In Cynictis, however, the groove does not quite reach the
rhinarium and is confined to the upper lip, where it is nothing
but an impressed line, not a definite gutter, dividing the median
naked area of skin. Inthe other genera of Mongooses possessing
this feature, the median naked area forms a gutter capable of
expansion and contraction. When contracted, it closes up com-
pletely and is represented superficially by a linear groove, the
right and left hairy areas of the upper lip being in contact
in the middle line. In Cynictis, therefore, we have a condition
of the upper lip nearly intermediate between the condition seen
in Mungos, with the guttered upper lip, and Ariela, with the
undivided upper lip.

The depth, or height, of the upper lip beneath the rhinarium
varies. In most genera it is less than the depth of the rhinarium.
But in Jehnewmia albicauda the lip is deeper—as deep, indeed,.as
the rhinarium. ‘The same applies to Crossarchus obscurus ; but
in this animal the rhinarium is relatively much deeper than in
Ichneumia, owing to the unusual depth of the infranarial portion
in front. Hence the lip itself is also relatively deeper. It is
the combined depths of the rhinarium and lip which impart to
Crossarchus the very characteristic somewhat pig-like appearance
about the snout-——an appearance not noticeable in any other species
-of Mongoose except Swricata *

Normally in Mongooses, as in other Carnivores, the two portions
of the upper lip to the right and left of the groove are closely in
contact, the groove itself : appearing as a narrow vertical line. The
function of the groove is to help the separation of the two halves
of the hp when “yaised to clear the teeth. When the groove is
obliterated, the snout is raised, thus drawing the lip upwards away
from the teeth, a phenomenon very noticeable in such forms as
Procyon and Nasua. In all Carnivora it seems that elongation of
the snout does not take place without obliteration of the groove ;
but the condition of the snout in Ariela shows that it is not.
true to say that obliteration of the groove always accompanies
-elongation of the snout.

The Feet.

Setting aside the variations recorded below, the feet of Mon-
gooses have the following characters in common. The claws
ave moderately long or very long, curved to a comparatively
small extent, and incapable of being lifted high off the ground
by the retraction of their phalanx upon the outer surface of
the penultimate phalanx, and the tips of the digits at the base
of the claws are never. provided with lobes of skin or thickly-

crowing hair. The digital pads are small. The plantar pad is

* The name Rhinogale, and its’substitute Rhynchogale, suggest a similar modi-
fication of the snout in that genus. The adult of this rare animal is, however,
known to me only from dried skins, in which the real length of the snout cannot be

«determined.

Proc. Zoou. Soc.—1916, No. XXIV. 24

358 MR. R. I. POCOCK ON THE

well developed, cushion-like and trilobed, and the area between
it and the digital pads, whether webbed or not, is naked. The
hallux and pollex, when present, are shortish or very short and
set above the plantar pad; and the hallucal and pollical lobes of

Text-figure 5.

A. Left fore foot of Mungos smithii. XE
Bb. ,, hind foot 5 on »
C. ,, fore foot of Atilax paludinosus. ,,
D. » hind foot oy) 2? ”

the plantar pad are small or obsolete, and detached from the
plantar pad. A single or double carpal pad is always present,
and separated by a naked tract from the plantar pad.

EXTERNAL CHARACTERS OF MONGOOSES. 359

Judging from the analogy supplied by other Carnivores, the
ancestral foot of the Mongooses was pentadactyle and plantigrade,
and furnished with well-developed interdigital webs and naked

soles.

Text-figure 6.

A. Right fore foot of Bdeogale puisa. Xz
B. ,, hind foot op 3 3
Ce os a Ichneumia albicauda. ,,
DS ee torestoon % 453 BS

The feet of several of the genera conform to this type, and those
of Mungos may be taken as an illustration and as a standard

with which the feet of other genera may be compared.
24*

360 MR. R. I. POCOCK ON THE

In a specimen of Mungos smithii, a Ceylonese species, the fore
foot is entirely naked beneath as far back as the carpus. The
digits, when spread, are seen to be slightly asymmetrically
arranged. The four main digits are united by webbing which
extends proximally up to the inner or admedian portion of the
rather small digital pads. Nevertheless they are capable of

a

Text-figure 7.

A.t Right fore foot of Crossarchus obscurus. X 3.

B. - ., hind foot £6 3 »
C. Left fore foot of Ariela fasciata. -
D. ,, hind foot 2p » »

being more widely divaricated than is the case in the arboreal or
terrestrial genera of Viverrine and Paradoxurine. The claws
are longish. The pollex is quite short, does not project laterally
so far as the second digit, and is inserted higher up the foot
than the adjacent portion of the plantar pad; its claw is well

EXTERNAL CHARACTERS OF MONGOOSES. 361

developed, but smaller than that of the other digits. The plantar
pad is swollen, trilobed and asymmetrical, the external lateral
lobe being a little larger and extending higher up the foot than
the internal lateral lobe. The distal margin of the median lobe

Text-figure 8.

A. Right fore foot of Suricata suricatta. X 3.
B. ,, hind foot 5 A 53
C. ,, fore foot of Cynictis penicillata. ,,
DF an hind&toot FA 3 s
E. Left fore foot of Mungos gracilis. sf
F. ,, hind foot 3 *f 3

is broadly truncated, and extends obliquely backwards and out-
wards to the point where it meets the external lobe. There is a
small pollical lobe, but it is detached from and higher up than
the posterior extremity of the internal lateral lobe of the plantar
pad. ‘The larger carpal pad occupies a similar position with

362 MR. R. I. POCOCK ON THE

regard to the posterior end of the external lateral lobe. It is on
the outer side of the middle line of the foot, and between it and
the edging of hair there is a naked area of considerable size.

The hind foot agrees with the fore foot in all essential respects,
but the digits are more symmetrically disposed and the claws are
shorter. ‘There is a small detached hallucal lobe, and above the
latter and on the corresponding area on the outer side of the foot
there are feeble indications of right and left metatarsal ridges.
The whole of the metatarsus is naked, and the nakedness extends
to the tip of the calcaneum (text-fig. 5, A, B, p. 358).

Sketches of the feet of several examples identified as IMuwngos
mungo, the commonest of the Indian Mongooses, show a close
general resemblance to the feet of MU. smithii described above, -
except that the pollex and hallux are set a little higher and the
edges of the main interdigital webs are a little more emarginate ;
but without further examples of I. smithii, it would be unwise
to attach systematic importance to these differences.

An example of J. auropunctatus from Chittagong also has
feet of this type; but the interdigital webs are more emarginate
than in JZ, muwngo—that is to say, when the digits are separated
the edges of the webs project to a rather lesser extent beyond the
lobes of the plantar pad. In this species, moreover, the hallux
is much smaller than the pollex, a disparity in size not noticeable
in the examples of I/. smithit and MW. mungo examined.

The only representative of the African species of J/ungos
I have examined is a spirit-specimen of JZ. gracilis*. In this
individual the feet are much more delicately formed than in the
Indian species, being narrower, with smaller pads and with the
webs considerably shallower even than in J/. awropunctatus, and
both hallux and pollex are small and set high up the foot ; but
the claws of all the digits are short. Except for the shortness of
the claws and the retention of the hallux, the feet of IW. gracilis
are nearly intermediate in structure between those of J. mungo
and Cynictis (text-fig. 8, E, F, p. 361).

Although the number of species and specimens of this genus
examined is small, a certain amount of variation in the depth of
the interdigital webs is noticeable. This feature will probably be
found useful for distinguishing species when investigated in forms
hitherto unexamined.

One other character is known to be variable, as Thomas has
shown for the African and Blanford for the Indian species,
namely, the extent to which the heel is covered with hair.
This feature may vary within the limits of a single species,
e. g. M. ichnewmon; and in MW. urva the upper part of the
metatarsus as well as the tarsus is hairy.

In Helogale the feet recall those of the Indian species of
Mungos, but, if anything, are more robust, with the webs

* Peters figured the soles of the feet of this species under the name JZ. ornatus
(‘Reise nach Mossambique,’ Siug. pl. xxvi., 1852). The shallowness of the webs
as shown, but details of the plantar pads are not indicated.

Dp

EXTERNAL CHARACTERS OF MONGOOSES. 363

more deeply emarginate. The hair on the carpus reaches the
carpal pad, which has a supplementary lobe at the base on
the inner (pollical) side. In the hind foot the hallux is small,
smaller than the pollex, as in Mungos awropunctatus, and the
heel is hairy, when the hairs are not worn off.

Peters’ figures of the feet of typical H. undulata from
Mozambique differ in some respects from those of the examples
of this genus I have seen (‘ Reise nach Mossambique,’ Siiug.
pl. xxv., a, 6). In the fore foot the hair does not reach the
carpal pad, which is small and single, and the pollex is set higher
up thefoot. In the hind foot a larger extent of the underside is
overgrown with hair. But the complete absence of detail in the
outline of the plantar pads does not attest care in the execution
of these figures *.

The feet of an example of Atilax paludinosus from South
Africa differ from those of JZungovs in one or two particulars,
notably in the complete suppression of the interdigital webs,
the digits being separated right down to the plantar pad. Both
pollex and hallux are long. The plantar pad is elongated and
distally narrowed, the apex of the median lobe being less
truncated than in Wwngos, and the two lateral lobes are set
relatively a little farther back. Small pollical and_hallucal
lobes are retained, but are detached from the posterior angle
of the internal lateral lobe of the plantar pads. In the fore foot,
the carpal pad is elongated and set on the external side of
the middle line of the naked carpal area. The hind foot shows
no distinct traces of metatarsal pads, and in the example
examined the whole of the metatarsus was naked beneath, and
a naked strip of skin extended along the underside of the heel
to its tip, but, as Thomas has shown, the degree of hairiness of
the tarso-metatarsus varies considerably within the species, this
area sometimes being naked as in the specimen described above,
sometimes the heel alone being hairy, and sometimes the hair
extending nearly as low as the plantar pad. Jam not aware
whether geographical races have been studied from the standpoint
of this character, or not.

The absence of the interdigital webs in this Mongoose
constitute, in my opinion, a valid reason for resuscitating the
genus Atilax (text-fig. 5, C, D).

In an example of Ichnewnia albicauda from Dutfile (White
Nile) the feet are slender and longish, with decidedly emarginate
webs, recalling in these respects those of Mungos gracilis rather
than of I. mungo or M. smithii. The hallux and pollex, about
equal in size, are set well above the plantar pad. The carpal
pad is semielliptical, of moderate size and higher than the

* In systematic works, Helogale is*merely distinguished from Mangos by the
suppression of the diastema between the canine and pm.? of the upper jaw,
pm.! being absent, as sometimes occurs in Mungos. As living animals, Helogale
and Mangos are very different in appearance, the former being a squat little
creature with a comparatively short tail and a broad head with short, pointed
muzzle.

364 MR. R. I. POCOCK ON THE

pollex, with only a narrow strip of naked skin above it. In
the hind limb the hairs, as recorded by others, extend all over
the back of the metatarsus practically down to the hallux.
Judging from dried skins, there does not appear to be any
marked variation within the species in the hairiness of the
metatarsus (text-fig. 6, C, D, p. 359).

This Mongoose is more digitigrade and stands higher on its.
legs than any member of the group of which I have seen living
specimens, not excepting even perhaps Atilax paludinosus.

In Ariela fasciata the fore foot closely resembles that of
Mungos except that the digits and the naked area behind the
plantar pad are relatively a little shorter, the claws longer, and
the interdigital webs somewhat shallower. The hind foot is.
rather narrower than in dMZungos, the lateral interdigital webs.
are much more deeply emarginate, whereas the median web
between the third and fourth digits is about as deep as in
Mungos but ties the toes a little closer together. The hallux is
as large as the pollex. The sole of the foot is naked back to the
tip of the caleaneum, and a little behind the plantar pad there
are traces of suppressed metatarsal pads (text-fig. 7, C, D, p. 360).

The fore foot of Crossarchus obscurus does not differ from that
of Ariela fasciata except that the carpal pad is larger and has a
small supplementary lobe on its inner or pollieal side, as in
Helogale, but detached from the main part of the pad. The
hind foot is relatively shorter than in Ariela fasciata, owing to
the shortness of the third and fourth digits. In the specimens
examined, the heel, when unworn, is hairy *, the naked area of
the metatarsus corresponding exactly with that of Helogale.
This area exhibits a pair of low elongated metatarsal pads, of
which the external projects farther forwards than the internal
(text-fig. 7, A, B).

In Cynictis penicillata the hind foot, as has been often stated,
is hairy below down to the plantar pad, and differs from that of
the genera hitherto recorded in the total suppression of the
hallux. The whole foot is comparatively long and narrow, and
the webs are very shallow, those between the second and third
and the fourth and fifth digits extending only slightly beyond
the plantar pad on each side, and although the web between the-
third and fourth digits is a little deeper, it only passes about half-
way up the admedian margin towards the digital pads, which,
like the digits themselves, are narrow. The claws are long. The
three lobes of the plantar pad form a tolerably evenly cordate
mass narrower than in Mungos and Crossarchus. The fore foot.
closely resembles the hind foot, but the claws are longer, the
interdigital webs are a little deeper, and the pollex, carrying
a long claw and set high above the plantar pad, is retained,
although short. The area between the plantar pad and the small
submedian carpal pad is quite naked (text-fig. 8, C, D, p. 361).

* Perhaps a variable character, since Thomas (P. Z. S. 1882, p. 86) mentioned:
the nakedness of the hind soles amongst the generic characters of Crossarchus.

EXTERNAL CHARACTERS OF MONGUOSES. 365

The genus Paracynictis *, recently established for the species
described by de Winton as Cynictis selousi, seems to resemble
Cynictis in the structure of the feet except that the pollex is
suppressed, as well as the hallux. In this respect the feet
resemble those of Swricata and Bdeogale.

The feet of two species of Bdeogale, namely, B. puisa and
B. crassicauda, have been figured by Peterst. Although the
details of the plantar pads are not very clearly shown, the
drawings are tolerably accurate, judging from a spirit-preserved
example of B. puisa, ticketed “ Zanzibar (Sir J. Kirk),” in the
British Museum. In this example the feet are, on the whole,
very symmetrical with respect to the plantar pads and the digits.
The latter are shorter and a little thicker than in Mungos, but
are webbed approximately to the same extent. There is no
trace externally of hallux or pollex. The carpal pad is large and
submedian in position, but with a slight external inclination.
Behind it there is a small triangular area of naked skin, and in
front of it a broad naked area separates it from the plantar pad.
All trace of the pollical lobe has disappeared with the pollex.
In the hind foot there is a semicircular area of naked skin
behind the plantar pad. Apart from this, the entire posterior
surface of the metatarsus is covered with hair; and judging from
dried skins, the hair in some species extends right down to the
plantar pad. In other respects the structure of the feet in this
genus is apparently tolerably uniform (text-fig. 6, A, B).

The last of the tetradactyl Mongooses is Suricata. In length
and narrowness the feet resemble those of Cynictis. The claws
are perhaps a little longer, and the webs are deeper, but they are
not so deep as in Mungos, being developed to approximately the
same extent as in Ariela. On both the fore and the hind foot
the web between the third and fourth digits is deeper than the
others, and on the hind foot the web between the third and fourth
is deeper than that between the fourth and fifth digits. The
digits are less symmetrical than in Oynictis, and markedly
asymmetrical as compared with those of Bdeogale. The carpal
and plantar pads are normal in development, and the external
lobe of the plantar pad of the hind foot is larger, sometimes
much larger than the internal lobe. ‘The area above the plantar
pad on the hind foot is naked to the tip of the heel, and towards.
the heel this area rises into a wide, low, laterally expanded pad-
like eminence which gives a sinuous outline to the naked tarso-
metatarsal area t (text-fig. 8, A, B).

* Ann. Mag. Nat. Hist. (8) xvii. p.177,1916. An examination of dried skins
suggests that the area between the carpal and plantar pads may be overgrown with
hair, thus contradicting the generalisatiofi (p. 358) as to the nakedness of this area
in Mongooses.

+ © Reise nach Mossambique,’ Saug. pls. xxvii. & xxviii, 1852.

{ Accounts of the metatarsal area vary. ‘Thomas and, following him, W. L.
Sclater correctly described this area as naked. But Mivart and, following him,
Flower and Lydekker wrongly described it as covered with hair. Its naked condi-
tion does not seem to be subject to variation.

366 MR, R. I. POCOCK ON THE

From the accounts above given, it is clear that the feet of
Mongooses exhibit a wide range of variation in such characters
as the numbers of the digits, the hairiness of the tarso-metatarsal
area, and the presence and extent of the interdigital webs. These
characters are either invariable or subject to much less variation
in other groups of corresponding rank amongst the Adluroidea.
There is only one group of the suborder, however, which possesses
feet structurally recalling those of the Mongooses, namely, the
Galidictine, the feet of which I have recently described and
figured *. -Between the feet of Galidictis and one of the penta-
dactyle, semiplantigrade Mongooses, like Mangos, there appear
to be only two differences which call for notice. In. Galidictis
(and in Galidia) the pollex and the hallux are set lower on the
foot and project therefrom on a level with the internal lateral
lobe of the plantar pad, and the pollical and hallucal lobes of this
pad are better developed and in contact with the internal lateral
lobe. Hence the plantar pad is quadrilobate, whereas in J/wngos
and all other genera of Mongooses the plantar pad is trilobate.
It may also be added that the metatarsal and carpal pads in
Galidia and Galidictis are better developed than in the Mongooses
and are double.

These differences are interesting because they show that the
feet of the Galadictines are of a more primitive type and, on the
whole, more Viverrine than are those of the Mongooses. Never-
theless it cannot be claimed either that the feet of M/wngos differ
more from those of Galidictis than they differ from the feet of
Bdeogale, Atilax, or Suricata, or that the feet of Galidictis differ
more from those of Mungos than they differ from the feet of the
Paradoxurine genera or of Hupleres.

The Glandular Anal Sac.

The presence of a glandular anal sac in Mongooses has long
been known ; but its invariable occurrence within the group has
been disputed. I have found it without exception in all the
specimens I have examined, even in those belonging to species
in which its existence has been denied. Cuvier, for example,
said that the Marsh-Mongoose, which he named Atilax vansire *,
is without it. It happens, on the contrary, to be rather ex-
ceptionally well developed in that form (text-fig. 9, B, C). It is
also present, though small, in Mungos auropunctatus, despite
Mivart’s statement { that in a living example he examined “ the
anus opened most distinctly on the surface of the body, and not
into a saccular depression.” Since Mivart was probably the
authority for Blanford’s declaration § that “this character is

* Ann. Mag. Nat. Hist. (8) xvi. pp. 351-356, pls. xiv., xv., 1915.

+ St. Hilaire & Cuvier, Hist. Nat. Mamm. ii. pt. 54, pl. 198, 1826.

~£ Proc. Zool. Soc. 1882, p. 178.

§ ‘Fauna of Brit. India’: Mammalia, p.119, 1888. It isa pity Blanford did not
particularise the species, and say whether his information was based upon his own

observations or not. It may here be recalled that Murie and others entirely failed
to find the large anal sac in a living Spotted Hyzna.

EXTERNAL CHARACTERS OF MONGOOSES. 367

ill-marked "or absent in some of the common Indian species”
[of Mungos], it may be explained at once that Mivart evidently

Text-figure 9.

Oe ) yy <\)

“A. Longitudinalfsection{of anal area of Atilaa paludinosus, 8. p., aval pouch ;
gl., cutaneous glands of pouch; o., orifice of anal gland ; a., anus ; 7., rectum ;
sce., scrotum with testis.

B. Anal area of the same. Lettering as in A, with p., penis.
C. Anal pouch of the same, closed.
D. Anal pouch of Ichneumia albicauda, 2. Lettering as in A, with v., vulva.
E. The same of Cynictis penicillata (young g). Lettering as in B.
F. The same of Bdeogale puisa, 9. Lettering as in D.
G. Glans penis of Crossarchus, from below.
HH. The same of Suricata.
I. The same of Cynictis.
K. The same of Mungos mungo.

368 MR. R. I. POCOCK ON THE

did not distinguish between the orifice of the sac and that of
the rectum which lies within it. That this would be a difficult,
task in a living animal is quite credible. Indeed, in such a form
as M. auropunctatus, it is easy at first sight to believe that the
sac itself is merely an enlarged anus, that its thickened rim is
the anal sphincter, and that the small rectal orifice within the
sac arises from constriction of the gut just within the anus;
and this view would be strengthened by the discovery of the
apertures of the anal glands well outside the inner orifice and not
within it, where, by the analogy of other carnivores, they should
be situated if the orifice in question were the anus. There are
reasons which make me judge that conclusion to be wrong.
In the first place, the external orifice of the sac is not shut by
the constriction of a circular sphincter muscle as the anus is ;
but when closed it forms a transverse, sometimes crescentically
upeurled, rima by the juxtaposition of its upper and lower
margins (text-figs. 9,C, and 10,A). In the sécond place, the
walls of the sac, sometimes at all events, show short hairs
projecting from the cutaneous follicles, suggesting its origin from
the involution of hairy circumanal integument. Finally, the
inner orifice itself is provided with a sphincter muscle. The
external position of the apertures of the anal glands, although
unusual, is not without parallel in the Carnivora, as is attested
by the condition seen in Hyena and Proteles.

In the Mongooses the position of these apertures varies.
Usually they are set one on each side of the anus, and tolerably
close to it (ungos mungo) or removed to some distance from it.
(Ariela, Suricata). In other cases they are placed somewhat
(Ichnewmia) or considerably (Cynictis) higher up in the anal
sac (text-fig. 9, D, E).

The secretion of these glands is always liquid and foul-smelling
and often copious. But, in addition, the free wall of the sae is
provided with well-developed cutaneous glands of the ordinary
kind, sometimes fairly uniformly distributed (Aédaa), sometimes
especially active and large in a half circle above the anus
(Mungos), sometimes located in definite paired areas of the sac.

Three of the genera, however, call for more detailed notice,
since the published descriptions of their glands do not agree in
all particulars with my observations.

When claiming for the first time the close and congeneric
affiliation between Crossarchus obscurus and Ariela fasciata,
Thomas added the following paragraph (P. Z. 8. 1882, p. 86,
note) :—“ Since the above was written, Prof. Mivart has pointed
out to me that the researches of Chatin into the structure of the
anal glands of the Carnivora (Ann. Sci. Nat. (5) xix. p. 89, 1874)
fully confirm the opinion here expressed as to the generic re-
lationship of the Striped Mongoose (C. fasciatus) with C. obscurus.”
Chatin does not, however, mention C. obscurus in the paper
referred to; and that Thomas was misled by Mivart is clearly
shown by the remarks of the latter on the subject in question

Qk >

i a

EXTERNAL CHARACTERS OF MONGOOSES,. 369

Text-figure 10.

. Closed anal sac of Helogale undulata, 2. v., vulva.
. Anal sac of the same, partially distended.
. Anal sac of Helogale undulata, g, on a larger scale, fully distended, and

showing glandular pouches or depressions. a., anus; 0., orifice of anal
gland; se., scrotum.

. The same of Ariela fasciata, 2. lettering as in preceding figures.
. The same, dissected and seen trom within, showing the single pair of normal

anal glands (a.g.), that of the left side opened to show oritice (0.) into sac;
7., rectum turned aside.

_ Anal sac of Crossarchus obscurus, 2, cut from below and spread. open.

r., rectum, cut open; «., fine cutaneous ridges extending*from upper margin
of sac to anus; other lettering as above.

_ The same anal sac shown partially distended when the tail is raised.
_ Anal sac of Suricata suricatta, 2, distended to show the glandular depressions,

with the scrotum-like swelling between the vulva and the lower rim of the
sac.

370 MR. R. I. POCOCK ON THE

(P. Z. 8. 1882, p. 183) :—‘*The anus opens into the middle of a
very large and deep fossa, into which several pairs of anal glands.
also open. The structure of these parts is described by M. Chatin
as they exist in both species. The condition found in C. obscurus
is described by himin....C. R. Assoc. frangaise, i. p. 557, 1872.
The parts of C. fasciatus are described and figured by him (under
the name of Herpestes fasciatus) in the Ann. des Se. Nat. vol. xix.
(5th series) 1874, p. 89, figs. 29-33 & 38. No less than five
pairs of glands are arranged about the anus, and pour their
secretion into the capacious and naked anal pouch.”

Reference to these two papers shows that the second is merely
an amplified edition of the first. C. obsewrus is not mentioned
in either. It is C. fasciatus that is described in both, and Chatin
does not even quote the first publication in the second. He was
clearly unacquainted with C. obscwrus, at all events so far as the
organs under discussion are concerned. Hence Mivart’s above
quoted summary of Chatin’s description of the anal sac and
glands in Crossarchus applies solely to C. fasciatus, and Thomas’s
claim of aftinity between the two species derived from Mivart’s
information falls to the ground.

In a maleand a female example of Crossarchus obscurus (text-fig.
10, F, G), I find the anal sac and glands resemble in a general way
those of typical Mongooses of the genera Mungos (= Herpestes),
Bdeogale, and Helogale. The anus is sunk ina central subcircular
depression surrounded by an upstanding thickened rim which is
about equal in thickness to the transverse diameter of the
depression. The skin of the superior or caudal area of this
thickened rim is furnished with a number of fine longitudinal
laminze which pass into the central depression above the anus,
where they break up into wrinkles set for the most part trans-
versely between the anus and the two orifices of the anal glands.
These orifices are quite conspicuous, open in the ordinary position,
and lead into glandular reservoirs of average size.

Thus the anal glands and anal sac in both sexes of Crossarchus
obscurus differ very considerably from those of the male Ariela
fasciata described by Chatin.

A female of Ariela fasciata, or of a closely-allied form brought
by Mr. G. Blaine, F.Z.S., from the Sudan (text-fig. 10, D, E),
has a very large anal sac as compared with that of most species
of Mongooses. Its surface is marked with three pairs of integu-
mental folds or depressions—two pairs above and one pair at the
sides of the anus. The depressions of the upper of the supra-
anal pairs are situated close together immediately beneath the
upper rim of the anal sac, and are sunk in a common fold of the
skin. Those of the lower of the supra-anal pairs are beneath
them, but more widely separated and nearer the anus. Those of
the lateral pair are placed far out towards the lateral rim of the
anal sac. They are much longer than the others, and approxi-
mately follow the curvature of the rim of the sac, but come to an

EXTERNAL CHARACTERS OF MONGOOSES. SHE

end on each side before reaching the middle line beneath the
anus.

Hach of the depressions or folds above described contains a
small glandular pit with several secreting pores, and the glands
beneath these pores appear to be simple enlargements of the
ordinary sebaceous or follicular glands of the skin. It is quite
clear that none of them represents the true anal glands of other
Carnivores. The orifices of these are situated on each side of
the anal sac close to the inner margin of the lateral folds. The
glands themselves are a pair of large muscular sacs filled with
dark-coloured, strongly smelling, oily fluid which escapes to the
exterior through the orifices above described. These glands
are quite different from the smaller glands of the anal sac
in their size, their saccular character, and the nature of their
secretion.

The character of the anal glands in this Sudanese female
example of Ariela fasciata explains much that was puzzling in
Chatin’s description of the glands in a male of the species from
South Africa. As has been already stated, Chatin assigned to
that species five pairs of anal glands opening by as many orifices
in folds upon the surface of the anal scent-pouch. These glands
he named the anterior, the lateral, the intermediate, the lateral
posterior, and the median posterior. It is not easy to homologize
all of these precisely with the glands I have described in the
female. But judging from their position and size, the lateral
glands appear to be the true anal glands, the median posterior
are evidently those that I have called the upper supra-anal,
while the intermediate and anterior probably correspond respect-
ively to the lower supra-anal and the lateral, the lateral portions
being undifferentiated in the female.

The principal difference, however, between the glands of the
two animals consists in this:—In the female the secondarily
specialised glands appear to be ordinary sebaceous or follicular
glands but little modified and quite distinct from the true anal
glands, whereas in the male they have been modified so as to
resemble approximately the true anal glands—that is to say, each
consists of a wall of secreting cells surrounding a sac or hollow for
storing the secretion, which is similar in nature to that of the
anal glands in being brown in colour and feetid in odour. This
secondarily acquired similarity—an exceedingly interesting fact—
seems to have misled Mivart into thinking that the two normal
anal glands present in all Afluroid Carnivores had become broken
up, as it were, in the male Ariela fasciata into the five pairs of
glands described by Chatin (Ann. Sci. Nat. (5) xix. pp. 89-93,
pl. iv. figs. 29- 30, 1874).

The existence of a pair of saccular anal glands in Suricata was,
as Mivart states, recorded by Daubenton. To this Mivart adds :—
“¢ The anus opens into the middle of a very deep fossa, deeper than
that of Bdeogale and like that of Crossarchus. ...I1 strongly

372 MR. R. I. POCOCK ON THE

suspect, from the form of the anal pouch, that there are here, as
in Orossarchus *, several pairs of anal glands.”

This is not the case in the examples of Suricata suricatta
I have examined.

Ina male the anal sac resembles that of Mungos mungo in a
general way, but is relatively larger. It is marked on each side
of the anus by a transversely oblique slit-like depression, and not
with several depressions as in Aviela fasciata. Its surface is
pitted with comparatively large and quite conspicuous hair-
follicles, and from most of these, perhaps all normally, a single
hair or a little tuft of hairs arises. Under pressure liquid
secretion can be squeezed from the pores of these follicles, which
are particularly numerous in the oblique depression above
described. When expanded, the depression is seen to curve
downwards towards the lateral margin of the anal pouch, and
the orifice of the anal gland is situated just below its deepest
portion, but is concealed within it when the depression is un-
expanded. ‘There is a single moderate-sized gland on each side,
as Daubenton said; and I can find no evidence for the multi-
plication of similar glands such as Chatin described in the male
Ariela fasciata. On the contrary, sections of the highly glandular
hair-follicles show them simply imbedded in the thickened skin
of the anal pouch without coalescing to form composite glands
with reservoirs for the storage of secretion. In its glandular
character the anal pouch recalls that of the female Ariela fasciata,
with the exception that the hair-follicles are more irregularly
scattered and not aggregated in so many definite integumental
depressions.

Tn a female (text-fig. 10, H) the gland is similar to that of the
male, and apparently as well developed. The two orifices of
the anal glands perforate the walls of the sac nearly midway
between its lateral border and the anus. The integument of the
sae round about them is pitted with large hair-follicles. These
also extend to the middle line of the sac both above and below
the anus, and there is an aggregation of larger pores lodged in a
depression just above the orifice of the gland on each side. As
in the male, this depression lies in the crease of skin formed
when the superior part of the sac closes over the inferior pait
when the tail is lowered. There appears to be no definite
storage-sac beneath this cluster of follicles, and, as in the female
Ariela fasciata, the only reservoirs for secretion are those of the
pair of anal glands proper.

The only other genus which requires particular mention in
this connection is Helogale. In the male and the female of the
species identified as Helogale undulata (text-fig. 10, A, B, C)
the anal sac is well developed and provided with supplementary
pouches. The anus itself lies in the centre of a slight depression
defined above by a fine but distinct cutaneous ridge. On each

* By Crossarchus Mivart meant the species referred in this paper to Crossarchus
obscurus, which he erroneously believed to have multiple anal glands, and Ariela
fasciata, iv which many anal glands had been described by Chatin.

EXTERNAL CHARACTERS OF MONGOOSES. 373

side of the depression there is a small supplementary pouch a
little higher up than the anus. In the middle line of the sac,
nearly midway between the anus and the root of the tail, there
is a moderately large unpaired supplementary pouch, and on each
side of this, but a little lower down and about half-way between
the anus and the margin of the sac, there is another moderately
large pouch. The orifices of the anal glands open just below
these in a line with the anus, but outside the central depression
in which the anus lies, Thus the anal sac of Helogale approaches
that of Suricata, and differs from the sac of any species of
Mungos 1 have examined.

Function of the Anal Sac.

On the material at my disposal I was unable to substantiate a
higher grade of development of the anal sac and its associated
glands in the male than in the female, or vice versa. On the
evidence, therefore, this composite glandular structure cannot be
included in the category of sexual organs except on the plea that
the secretions may help the sexes to find each other.

Captive Mongooses have the habit, also observed in Genets
and Civets, of rubbing the glandular surface against the walls or
projecting angles in their cage or against the legs of chairs and
tables in a dwelling-room. Hence it may be inferred that one
of the functions of the secretion is to make the animal’s sur-
roundings smell of itself, and the scent so applied serves, I
believe, the purpose of familiarising the Mongoose with every
square yard of its environment, so that, independently of vision,
if need be, it can find its way with precision over any road it has
once travelled *.

Several of the Mongooses, too, like Cynictis, Swricata, Ariela,
and Helogale are gregarious tT; and of one species of Helogale at
least the habit of hunting in packs has been recorded. One of
the larger Indian species of Mongoose (J. vitticollis) has been
seen combining in pairs in pursuit of prey. And since specialised
cutaneous glands very often attain exceptional development in
gregarious animals for the purpose, presumably, of helping indi-
viduals to keep together, it is not improbable that the secretion
of the glandular anal sac has a functional significance in that
respect in some of the Mongooses.

Finally, Hodgson’s record $ that the secretion of the paired
anal glands in Mungos urva is ‘‘ aqueous, horribly fcetid, and
projectile to a great distance by the living animal,” suggests that

* Tt is well known that most mammals have the habit of keeping to definite
beaten tracks. ‘The advantage of this to species like rats and rabbits is very evident.
TI have frequently seen rats escape from dogs by knowing exactly the position of a
pipe or of a hole in a wire-net fence. They make a bee-line for the spot at full
speed, and, apparently without ever seeing the hole, go straight through it, knowing
the precise direction to take by complete familiarity with the track, owing, I believe,
to the scent it holds. ;

+ See W. L. Sclater, ‘ Fauna of 8. Africa’: Mammalia, vol. i. p. 69, 1900.

< Quoted by Blanford, ‘ Fauna of Brit. India’: Mammaha, p. 129.

§ Journ. As. Soc. Bengal, vi. pt. 1. p. 563, 1837; also my paper in Ann. Mag.
Nat. Hist. (8) vill. p. 756, 1911.

Proc. Zoon. Soc.—1916, No. XXV. 25

374 ON THE EXTERNAL CHARACTERS OF MONGOOSES.

this species at least makes use of the secretion in the same way
as the Skunk. I have never seen this species alive, and have
never noticed any Mongoose practise the habit ; but I have seen
the secretion of a dead Marsh-Mongoose (A. paludinosus) issue,
under pressure of the gland, in a narrow jet as if propelled by a
squirt; and since Mr. W. L. Sclater states that this animal is
able to diffuse a strong odour described as ‘“‘ sweet-sickening”
from its anal glands, I suspect it is endowed with the same
power as J. urva. ‘To me the scent of the secretion in 4. palu-
dinosus is very nauseous.

The External Genitalia.

In the male the glans penis is always short and smooth, and
emerges close in front of the scrotum as in the Felide and
Nandinia. he orifice of the urethra is apparently always a
large and elongated slit opening in the middle of the underside
of the glans and not at its tip. The glans is laterally compressed,
usually much longer than wide, attenuated towards the apex, and
carries a bone or “ baculum” which reaches the tip. In Suricata,
however, it is markedly piriform and considerably expanded
proximally, so that its width nearly equals its length (text-fig. 9,
Ces bee Pel .@

The vulva is only a short distance below, or in front of, the
inferior edge of the anal sac. The naked area is sometimes
broadly continuous with that of the sac, as in Bdeogale puisa
(text-fig. 9, F), sometimes connected with it by a narrow naked
tract, as in Ariela fasciata and Crossarchus obscurus (text-fig. 10,
G), or separated therefrom by a narrow tract of hair, as in
Helogale wndulata (text-fig. 10, A).

In the female of Surieata Mivart drew attention to the
presence of a perineal swelling just below the anal sac, and
compared it to a small scrotum (text-fig. 10, H). A somewhat
similar swelling is present in the female of Mungos smithit
T examined. This swelling occupies the position of the perfume-
gland of the Civets and Genets. Since Mivart did not dissect
the swelling in Suwricata, it may be recorded that it appears to
consist of fatty and not of glandular tissue. It is, in my opinion,
exactly comparable to the scrotum-like excrescence so often
noticed in female Hyzenas and to the apparently similar structure
observed by Lénnberg * in a female Cryptoprocta. Since the
Mongooses resemble the Hyzenas and Cryptoprocta in possessing
a large anal sac, and have been compared with them in other
respects, the remarkable differences in the structure of the penis
in the three groups must be borne in mind. In the Hyenas
(and Proteles) this organ is very long, fleshy, pendulous, and the
glans is short and boneless. In Cryptoprocta the glans is ex-
ceedingly long, copiously armed with strong spicules, provided
with a long bone, and emerges a long way in front of the
scrotum,

* Bih. Svensk. Vet.-Akad. Handl. xxviii. Afd. iv. no. 8, 1902.

ON THE SITATUNGA OF THE SESSE ISLANDS. BS

12. Notes on the Sitatunga or Marsh Antelope of the
Sesse Islands, Lake Victoria Nyanza. By Major
R. MEINERTZHAGEN, F.Z.S.

[Received January 14, 1916: Read April 4, 1916.]

(Text-figures 1 & 2.)

INDEX.
SYSTEMATIC: Page
Limnotragus spekei sylvestris, subsp. n. ............ 380
SHITE O10 G vauen: Nae eee R ey ete aT Bots ea sta goauneiees seeent aches, O795. 580

BuUGALLA ISLAND.

The southern end of Bugalla, the main island of the Sesse
‘Group, was visited on the 21st, 22nd, and 23rd October, 1915.
In all seven mature buck were examined in the flesh, whilst
twenty-two mature buck and over fifty other buck, females and
young, were observed, some at very close quarters. The animals
were found to be so plentiful at one spot, that as many as twelve
warrantable buck, nine females, and five young were seen from
one anthill.

Colour, etc.—Old males appear typical of mainland specimens ;
a few white flecks and occasionally an obsolete stripe can be
observed on the flank. Skin black and inclined to be hairless
between the horns. Hair very thin on the under parts, long and
coarse on the back.

Adult females were seen to be both dark brown and red, the
latter colour predominating in the proportion of seven to one.
Red females invariably had red young. Brown females were
never seen with young, and they may be old ones past bearing.

All young seen were red, with white spots and flecks on the
flanks and hind quarters. Solitary young were often observed
lying in the open and away from their mothers. This is pro-
bably accounted for by the complete absence of any natural
enemies on the island.

Skin very greasy. On placing a freshly shot buck in the water,
a film of oil floated to the surface.

Skull and horns.—The amount of white at the tips of the
horns depends on the age of theanimal. It seems that the horns
of very old buck tend to lose altogether the white tips, which
are invariably present in younger individuals. Old buck wear
their horns to a considerable degree by rubbing them on trees
and anthills. This exposes the yellow under surface of the horn
and adds considerably to its beauty.

The shape of the horns (text-fig. 1) is different from the usual

25*

Text-figure 1.

376 MAJOR R. MEINERTZHAGEN ON THE

mainland type, having a wider splay. In specimens from the
mainland the horns do not usually diverge to any great extent,
the tips, however, nearly always showing a tendency to splay
out considerably, whereas in the Bugalla type the tips splay out

pee sin eet rater eee OTe SOE =e Sees cesar sess ala eT

but slightly. Whether this type is constant or not on Bugalla
Island, I cannot say, but I never saw a ‘ mainland” or ‘“‘ Nkose”
type on Bugalla Island, neither did I see a “mainland” or
“ Bugalla ” type on Nkose Island.

Photograph of the skull and horns of (A) Nkose, (B) Entebbe, an exaggerated type, (C) Bugalla Sitatunga.

e

SITATUNGA OF THE SESSE ISLANDS.

377

The skull, horn, and body measurements (in inches) of Bugalla

and Nkose Sitatunga are as follows :—

Horns.

No. |Locality.| Sex. ]

Length | Girth

of curve. at base.
A. | Bugalla.| g¢ | 24/s 7/8
B. | Bugalla.| ¢ | 22/6 7»
C. | Bugalla.| g | 23°’s 73/4
D. | Bugalla.| g | 24 Te
E. | Bugalla.| ¢ | 23
F. | Nkose...| ¢@ | 22%/4 79/5
G. | Nkose...| ¢ | 231/4 7°/8

Tip to

tip.

Skull

1625/)5| 111/,

114/s
14°/s

|173/g

| Op

| 62/4

| 115/e
11"/2
11"/4

|
|

114/16
| 117/s

length. |

Bopy.

nan

Height. | eneth | rail, | Weight.
35> | 5912 | 84/> | 204 Ibs.
3 | @l 9 | 175
35. | 6s | 81s 196

san iiliies 83/, | 200

_ 84l/y | 62 Gia 180
39 65 91/, | 296
a7. | 6 | 9 | 219

male shot at Entebbe, which measured 10°52 inches.

Norr.—The only mainland skull I have been able to measure is that of an adult

Feet.—An indiarubber-like pad stretches for 23 inches from
the heel of the main hoof, with a small patch of short bristly

hairs 1} inches from the heel (text-fig. 2 A.).

The feet splay considerably, and the hoofs can easily be moved
up and down in a freshly killed specimen, so that they form an
50°. The late Captain R. B. Woosnam told me
he suspected that when walking in swamps, they catch hold of
stalks of weeds and papyrus to assist in progression, and this
appears quite possible.

angle of about

The feet measurements (in inches) are as follows :—

| |
Hoofs, | Width
Specimen.| Sex. Locality. |Fore and! a-p*, | 4-8, C-E, |C-D,on|} atE
Hind. | straight.) straight.| straight.) curve. | across
| | base.
fh F. | 61/6 1's 1:41 3 1°75
GAS Os g || Bugalla. { Be ee [carte a hesie cu eee ol eee
ii pity I Oe Aca 1:39 | 8 173
J Beepeanees 3 Bugalla. ?) H. 6 Wy 1 WAG 1:27 3 1:59
4, EL |: 63/4 1 | wens 1:78
Cnt! o | eae jee | Re ale | aes |) ein, | ae
‘ a Oy Pa aE 175
Date: é Bugalla. ; H. 63/5 1/s 1:27 31/3 1:62
‘ Ira F. | 63/g Wig | eee ere ee
FENG Rea 3 Bugalla. ; H. 6 Vg 7 hg 1:28 3 I/g 1°63
ioe (ins Genk 61/4 1 1°68 27/5 2°32
1D eoemate 3 Nkose. | d H. 6 V5 1 1:62 9 lb/6 9:09
ak FB. 61/5 1 171 27/s 2°34
Ga eet: 3 | Nkose. ; E. 61/5 Tale 1:63 27 /g O17
* For explanation of these letters see text-fig. 2 A.

378

MAJOR Rh. MEINERTZHAGEN ON THE

Text-figure 2.

A

9

Aieesga

i f
if main Hoe
ES i f

Lediarubber-like Pad i

Short bristly Heel E
hairs
B

A

LL
ge ey

AAT

as
S

A. Diagram of foot of the Bugalla Sitatunga, for use with tables of
measurements on pp. 377, 381.
B. Hoof and false hoof of the Nkose Sitatunga, one-half natural size.

C. Hoof and false hoof of the Bugalla Sitatunga, one-half natural size..

SITATUNGA OF THE SESSE ISLANDS. 379

Habits —These Sitatunga appear to have developed the habits
of Waterbuck, living in the forest in the heat of the day and
coming boldly into open grass-land at other times. At about
5.30 p.m. they come out and walk straight away from the forest,
not hanging about the edge as Bushbuck do, Most animals
return to the forest about 8.30 a.m., though some were seen still
feeding on the open grass at 10.30 a.m. In the forest they lie
up in the densest thickets, but once in the open they appear to
be almost fearless. The firing of a rifle within 200 yards of
grazing animals did not always induce them to seek cover.
Though the human smell was always noted with suspicion, it was
not always treated with alarm.

The majority of the animals seen were on the edge of the
forest which grows along the shores of the island, but several
were observed among the reeds on the lake edge, and among bush
on the very tops of the grass-covered hills.

These Sitatunga are both browsers and grazers. ‘hey are
particularly fond of feeding along the edge of the forest, and on
two occasions buck were seen on their hind legs like goats,
browsing off forest shrubs. The stomachs of three buck examined
contained grass, leaves, and a little bark.

When alarmed the noise is a deep grunt, and I was unable to
distinguish between that made by the two sexes.

A slow stately walk seems to be the usual mode of progression.
I never saw one trot. When they make off they go clumsily,
dragging their hind legs with the slightest suggestion of a
kangaroo’s gait. Their action when running or walking is very
high. They are very averse to facing a hill, either up or down,
and one buck, which I compelled to gallop down a steep hill,
tripped over himself twice and completed the journey most
clumsily, but much to his own alarm, for not content with barking
and grunting at every bump he took on his downward journey,
he continued loudly to advertise his concern for nearly half an
hour later.

It will probably be found that this Bugalla type of Sitatunga
is entitled to subspecific rank, but not having had the oppor-
tunity of examining mainland specimens, no further remarks will
be made on this point.

Nose I[suAND.

Nkose Island, the southernmost of the Sesse Group, was visited
on the afternoon of the 23rd October, 1915. It is about
2000 yards long and 300 broad. Throughout its entire length
it is covered with dense forest, which overhangs the water’s edge.
There is practically no undergrowth, but the tangled mass of
roots, creepers, and fallen trees makes progression difficult and
affords dense cover for the Sitatunga, which appear to lie up in
the thickest parts during the daytime. At the southern ex-
tremity of the island there is about an acre of short grass.

380 MAJOR R. MEINERTZHAGEN ON THE

In all I saw twelve warrantable buck and numberless females
and immature specimens. I killed one buck, and the rest of the
party Killed an adult buck, an immature buck, and captured
a female and young, about a month old. I had exceptional
opportunities for close observation, for sitting by a game-
path I actually stroked several beasts as they walked slowly
past me.

These Sitatunga differ remarkably from the Bugalla type.
Their colour, length of skull, type of horn, greater size and
weight, and different shape of feet appear to entitle them to sub-
specific rank, and I propose the name of Limmnotragus speket
sylvestris for this island form.

Colour, etc.—The adult male is of a uniform dull mouse-colour,
and not a dark brown. ‘The legs have no sign of reddish marks.
There is no light mark on either the throat or under side of the
neck. The hair is thicker than in Bugalla specimens, and the
skin can seldom be seen through it, except on the under parts.
All females seen were red, no brown ones being observed. All
young were bright red, the one captured being well marked with
white flecks and spots on the flanks and hind quarters.

Skull and horns.—The shape of the horns is well exemplified
by the photograph (text-fig. 1). The horns of all I saw were of
this “ bushbuck ” type.

The animal is larger and heavier than the Bugalla type, as
will be seen by referring to the table of measurements on p. 377 ;
the smallest Nkose skull is 3 inch longer than the largest Bugalla
skull.

Feet.—The hoofs are, perhaps, the most distinguishing feature,
and this is well exemplified by text-fig.2. The length of the
hoofs differs but slightly from that of Bugalla specimens, but they
are much stouter and stronger. The difference in shape of the
false hoofs in the two forms is shown in text-fig. 2, B, C.
Neither of the three specimens shot had the small hair-patch on
the pad of indiarubber-like skin behind the hoof, whereas all the
Bugalla specimens had such a small hair-patch. The feet of the
young one captured were not abnormally long, in fact, no longer
than one would expect to find in a young Bushbuck of his
age.
AMeererients of the feet are given in the table on p. 377.

Habits.—They are of necessity entirely browsers and eat a lot
of bark. They live in dense dry forest, seldom seeing the light
of the sun, for the small patch of open grass at the southern
end of the island was particularly devoid of tracks. It is un-
doubtedly these peculiar surroundings which have produced such
a peculiar form. Whether such a form exists or not on other
such small afforested islands of the Sesse Group is not yet known.
On Nkose Island there must be at least 200 individuals.

SITATUNGA OF THE SESSE ISLANDS. 381

Appirionatu Note.—Since writing the above I have been able
to examine three adult males shot on Nkose Island in November
1915. The skulls, horns, feet, and colour bear out what is said
above, the measurements (in inches) being given below :—

| |
| |
f so 3 Horns.
| ae} es | i)
Hoof, | 2 | S| lee
: 3 oS on tet) =| Ss
Speci- : Wore |) SF | 9 || a oS |e ah eels Skull
| Sex./Locality. 2 | 8 | 2 a |®a2/ ao |e 5
men. and e/< Be Bi Se le al S length.
; BS m aay) a esl a
Hind.| * Ellie 5 Bl ae esi es
A A <3) A wae] 25 |-FAQ eae
| Hoe Bas

5) 8 | 229
69) 2155] 221, 7

3 | Nkose. ‘tf He
I...) g | Nkose. if 6/4 Li A Bis | zal) 201/4| 74/4) 61/.| 12

(F 16

3 UH 16

|
e
(=)
ee
=
=
uo
—
=
oo

6 61/4) 11/4) 31/4 2°32
|

. | 61/5) 13/g
. | 63/5) 11/4)

*69| 31/4 | 2°36) |
64 ae a19| 21/4) 71/4 11s 12%s

| Nkose.

* For explanation of these letters see text-fig. 2 A, p. 378.

ON PATTERNS CONSPICUOUS IN NATURE. 383.

13. An Experimental Determination of the Factors which
cause Patterns to appear Conspicuous in Nature. By

J.C. Morrram, M.B. (Lond.)*.
[Received March 4, 1916: Read May 9, 1916.]

(Text-figures 1-20.)

INDEX.

MorpPHoLoey : Pages
Experimental Analysis of Patterns ..................655 383-410
Conspicuous Patterns among Indian Lepidoptera... 410-418

ETHOLOGY :

Revealing power of Patterns... .............cseeeeeeuenes 383-418

INTRODUCTION.

It is an undisputed fact that patterns often render animals
inconspicuous in Nature. These patterns have definite cha-
racters on which their inconspicuousness depends, and in a
previous paper (P. Z.S. 1915, p. 679) some of these characters
were defined. Just as against any single background, or against
any series of backgrounds, patterns can be placed which will
appear inconspicuous, so other patterns can be placed which
will appear relatively conspicuous. Experiments were carried
out to determine the characters which render patterns con-
spicuous, and these are dealt with in Part I. of this paper.
Having defined the factors for conspicuousness, the Indian
Diurnal Lepidoptera were examined to see whether any of
these insects presented patterns which must render them con-
spicuous. Part II. deals with this consideration.

Part I.
‘SCHEME OF DESCRIPTION.

Experiments were carried out with artificial patterns,
against artificial backgrounds. These are described under four
headings ;—

(A) The consideration of plain objects against plain back-

grounds.

(B) The consideration of patterned objects against plain back-

grounds.

(C) The consideration of plain objects against patterned back-

grounds.

(D) The consideration of patterned objects against patterned

backgrounds.

The experimental conditions are shown in text-fig. 1; standard

* Communicated by the SECRETARY.

384 DR. J. C. MOLTRAM ON

candles were used. In order to obtain a series. of backgrounds
ranging from dark to light tone, the object was fixed on a
glass plate and was illumined separately from the background,
as shown in text-fig. 1, B;.by moving the background near to
or away from its illumination, and by using backgrounds of
different tones, it was possible to obtain every grade of tone,
from black to white.

In these experiments tone is alone considered; colour was
kept constant by using only black, white, and neutral greys.

(A) Plain Objects against Plain Backgrounds.

Plain objects may enter into consideration of pattern, because
they are in reality objects covered by a very small pattern.
The visibility of plain objects was found to be affected by the
following factors :—the human eye, the lighting, the atmosphere,
the background, and the object.

The human eye, even when it does not present some gross
defect, nevertheless is found to vary from individual to indi-
vidual; so that the readings made by one person cannot be
directly compared with those of others. Working with the
same eye, several factors affect visibility ; if the eye be allowed
to become fatigued, the greatest distance at which objects can be
seen is much reduced. Experiments showed that from thirty to
forty observations could be made during two hours without
encountering fatigue effects.

Some time must be allowed for the eye to become accommodated
to a sudden change in illumination: for a change from daylight
to almost complete darkness, twenty minutes is necessary ;
working with two standard candles, it was found that fifteen
minutes must be allowed.

The eye was also found to vary somewhat from day to day:
health and general fatigue are probably the cause of these
variations. Owing to this, one cannot directly compare the
reading of one day with that of another. The accuracy with
which measurement can be made is indicated in experiment
moa:

The effect of the opacity of the air on visibility does not enter
into these experiments, as no measurements beyond eighty feet
were made, and observations were not made during fogs.

The Effect of Lighting —Experiments showed that the greater
the illumination the greater the distance at which objects can be
seen, all other factors remaining constant.

The Effect of the Background.—A plain object is visible at a
great or small distance according as to whether the difference in
the amount of light coming from the object and the background
is great or small. A white object is more visible against a black
background than against a grey one, and more visible against
a dark-grey than against a light-grey one. Experiments were
not carried out to define this relation more accurately ; but the

PATTERNS CONSPICUOUS IN NATURE. 385

Text-figure l.

O

1 foot.
a

Ground plan of apparatus used for measuring visibility of objects and blending
distance of patterns. A = when object and background are illuminated by the
same light. B= when illuminated by different light.

c=standard candles, F=screens, E=eye, B=background, o=object,
G=glass plate.

EXPERIMENT No. 1.

A. Candles 1 ft. apart : conditions as in text-fig. 1: candles distant 1 ft. from glass
plate on which object was placed: background of grey paper 2 ft. from glass
plate. Object of black ueedle-paper, 36 sq.mm. The following ten readings
(in feet) of the greatest distance at which the object could be seen, were made at
intervals of five minutes :—72, 72:8, 72°9, 72°8, 74°4, 73°4, 74:1, 73°5, 72°7, 72°9.

B. Candles 1 ft. apart: patterns 2 ft. 1 in. from candles: pattern consists of
alternate black and white squares, 25 sq. mm.: pattern covered, 16 sq. cm.
The following ten readings (in feet) of the distance at which the pattern
blended into an even grey tone, were made at intervals of five minutes :—
31, 33, 33°6, 34°f, 34, 33°9, 34°5, 33°7, 33°8, 343, 34.

following experiment (no. 2) was made as it has a bearing on
the relative visibility of patterns, as will appear later. It shows

386 DR. J. C. MOTTRAM ON

that a constant-in-tone contrast between object and background
does not result in a constant visibility.

Text-figure 2.

N

09

OC OF OF Og

'222f OY

7 gz 3 & 5 fet,

Relative visibility of objects.
Ordinates = distance at which object is visible.
Abscissee = distance of candles from object.

X = white object against black background.
© = black object against white background.

EXPERIMENT No. 2.

Experimental conditions as in text-fig. 1. Candles 1 ft. apart.

Materials:—Backgrounds: white, of white Bristol board 100 sq. em.; black, of
black needle-paper 100 sq. cm. Objects :—White, of white Bristol board
45 sq. cm.; black, of black needle-paper 4°5 sq. em.

Results (mean of three observations).

When distance from white square was black square was
candles to object visible against black visible against white
was background at background at
1 foot 66°5 feet 41°6 feet
2 feet 504, 344 ,,
3 oP) 44-1 ” 32°2 22
Te 40°9 ,, 3
5 ep BA 30°77 ,,

This experiment shows that a white object on a black back-

PATTERNS CONSPICUOUS IN NATURE. 387

ground is more visible than a black object on a white ground.
It can be seen (text-fig. 2) that the higher the illumination the
greater the difference ; at low illuminations the curves of visibility
approach one another (they would meet at complete darkness).
The same was found to be the case when object and background
were of different shades of grey, instead of black and white. The
light-grey object against the dark-grey background is more visible
than the dark-grey object against the light background.

It was thought that the lower visibility of the black object on
the white ground might be due to the dazzling effect of the
large area of white. It was found that reduction in the area of
the white background by means of black diaphragms produced
the opposite effect, and, further, that the nearer the diaphragm
was brought to the object the less visible the object became. It
was further found that when the white square on the black
background was similarly surrounded by white diaphragms, the
same effect resulted (see experiment no. 3).

EXPERIMENT No. 3.

Experimental conditions as in text-fig. 1. Candles 1 ft. 6 ins. apart;
distance of candles from object, 2 ft.

Materials :—Backgrounds of white Bristol board and black needle-paper, 100 sq. cm.
Objects of same materials, 4°5 sq.cm. Black and white square diaphragms,
total size 100 sq. cm., with a central square hole : -

No. 1. Size of central hole was 64 sq. cm.

3” 2. 29 29 39

bP) 3. 39 29 16 39

32 4. ” 9 4, 33
5 1

bP) o, 9 33 33

3) 6. eb} 2 0 16 29

White object on black background was visible at 45°5 feet.
When surmounted by No.1 white diaphragm, at 40°7

29 »” 2 ) ” 37°6 29
oy) 2) 3 oy) ” 348 ”
23 22 4 29 » 29°5 »
) 3 5 2 2 24°0 2:
) oD) 6 2 ” 13°4 9

Black object on white background was visible at 38°4 feet.
When surmounted by No. 1 black diaphragm, at 32°2

bb)

29 22 2 bb) 39 30° 39
29 29 3 39 39 30 33

b) 29 4 22 33 29°6 39
39 23 5 ” 39 21:7 3)
39 ey} 6 39 39 15°5 39

(Above readings are the mean of three observations.)

It would thus appear that light tone on dark is more visible
than dark on light. This has been considered to be due to the
eye recognising the object, in one case by a_ positive image,
in the other by the absence of stimulation. This fact is of
considerable importance in regard to the visibility of animals
in Nature: those exhibiting large areas of light tone must

5

388 DR. J. GC. MOTTRAM ON

be considered to be, other things being equal, much more
conspicuous than those which do not. For instance, a light-
coloured butterfly flying across a meadow, or down a hedgerow,
is visible at a much greater distance than a dark one.

This greater visibility of light-toned objects in Nature can
easily be demonstrated by comparing the visibility of black
and white discs against a great number of natural backgrounds :
only against snow and certain parts of the sky is the white
the less visible; against the vast majority of backgrounds the
white is very much more visible.

Text-figure 3.

OL

OF

4 1GAR. 859 3S G . 15 ub Lh. | OIG
Visibility of objects:‘in proportion to size.

Ordinates = distance (in feet) at which object is visible.
Abscissee = size of object, in square millimetres.

ExprEriment No. 4.
Experimental conditions as in text-fig. 1. Candles 1 ft. apart and 2 ft. from object.

Materials :—Black and white backgrounds of Bristol board and black needle-paper.
Objects :—Black needle-paper of the following sizes: 2°5, 3°7, 4°8, 72, and
8:8 sq. mm.; and white paper of the following sizes: 1, 1°6, 19, 28, and
3°5 sq. mm. :
The above diagram shows the distance at which the black objects were visible
against the white background (X) and at which the white objects were visible
against the black background (©).

An even more convincing way of demonstrating this fact is to
take a series of artificial backgrounds, from white, through grey,

PATTERNS CONSPICUOUS IN NATURE.

389

to black: find the background against which black and white
are equally visible under some natural condition of lighting—
for instance, in a wood; now compare this background ith

Sty Ge WT

Z
b
|

C.

Text-figure 4.

+
/
/ P
Ye /
J i
/ i/
Seam mais /
44 eee /
_X- /
2-9
SQ
<- DS ds RZ.

Diagram showing the correspondence between the visibility of objects of different
shapes (circle, square, isosceles triangle, and rectangles) and the concentration

of their areas.

Candles 1 ft.

EXPERIMENT No. 5.

apart; objects distant from candles 1 ft. 6 ins.

Objects of black needle-paper, area 16 sq. mm.; background of white Bristol board.

Objects. Distance at which Inverse figures.
visible, in feet.
Circle sae iaantsaeeeee 59 170
Square.......... te 58 173
Isosceles gsomella oe 90° . 53 190
Baan, O92 sccoe cc cde. 51 198
Rectangle, 16X11 ...........: 38 263

(Mean of five readings to nearest whole number.)

Proc. Zoou. Soc.—1916, No. XX VI.

26

390 DR. J. C. MOTTRAM ON

Circumference
Area
Circle QV mx _.3'56
a ae
é Ax 4.
NSO AUIS RG) memebpadtiodacoatod besesade " ==
wv Xv
(2427 2)x 4:83
ane ara a)da
Tsosceles triangle of 90°...... een ee,
bx 5
JR@C NENA, ASK oc coosoonons0 2 Paar
8 dx 85
Rectanele; 4 <2 3. cce eee 5 —
a? x

In text-fig. 4. the inverse visibility figures (X) are conventionally plotted with the
numerator of the periphery over area (©). It can be seen that the visibility
curve closely follows the concentration of area curve.

the surrounding natural backgrounds: it will be found that it
is very much lighter in tone than the lightest natural back-
ground which can be found in the wood.

On referring to the diagram (text-fig. 2, p. 886) it can be
seen that the difference in visibility between white and black
is greater at high illumination than at low. It follows that
at night white is, in Nature, not nearly so conspicuous with
regard to black as it is during daylight.

Diurnal animals pr esenting a large area of white or light tone
must therefore be considered to be. much more conspicuous than
nocturnal animals similarly patterned.

The Effect of the Object’s Characters on Visibility—The cha-
racters, size, and shape will affect the visibility of an object
when all other factors are kept constant. When the contrast
in tone between object and background is great (the object
being light in tone and the background dark), the human eye
is able to define an object subtending an angle of approximately
one minute. Distinction must be made between the ability to
define or focus, and that to see: the eye cannot define a star,
although it may be able to see it.

Keeping the shape of the object. constant, experiments show
that visibility is directly proportional to size: the larger the
object, the greater the distance at which it is visible.

WV hether the relation between size and visibility is the same
for all shapes has not been determined; though during the
course of this and other investigations a large number of
shapes have been examined, no exceptions have been noted.

The visibility of objects is dependent upon their shape. Circles,
squares, triangles, and rectangles of the same area are not
equally visible. Experiments show that the more concentrated
the area the greater the visibility.

In the following experiments concentration is measured by
the ratio of circumference over area, and it can be seen that the
distance at which the object is visible is inversely propertiongue
to this ratio (see experiment no. 9).

PATTERNS CONSPICUOUS IN NATURE, 391

As a circle is the most concentrated form that an object can
have, therefore it is the most visible form. The ratio ‘cir-
cumference over area only gives the concentration for simple
figures. Objects can be made of the same area and of the
same circumference but of different concentration. In these
cases concentration must be represented by the moment of area
round the centre of area.

Text-figure 5.

TAS k.

z. 4.

Four figures of the same area and of the same circumference but which are not
equally visible. The concentration of their areas is estimated by the length of
the cord joining the centre of area of the two triangles.

EXPERIMENT No. 6.

Candles 1 ft. apart. Objects distant from candles 2 ft. 1 in.

Objects of black needle-paper, of the same area and having the same circumference,
composed of two triangles as shown in the figure; the longest side measured
1'1 cm., the shortest “5 cm., and the angle opposite the longest side was a right
angle. Background of white paper.

Distance
Object. at which visible.
NOs secede Yo secreence 51 feet.
bb) 2 50 33
gE Nd i» BS rh ee
ye 45°5 ,,

(Mean of six observations.)

The figure shows the lengths of the cords joining the centres of areas of the two
triangles; it can be seen that when the cord is short and the area therefore con-
centrated, then the visibility is great, and vice versa.

392 DR. J. C. MOTTRAM ON

In experiment no. 6 mathematical expressions are avoided
by presenting the concentration as the length of a cord. Areas
of the same size and circumference, but of different concentration,
are dealt with in this experiment; and the same conclusion is
arrived at, namely, that the more concentrated the area of the
object the greater its-visibility.

‘This completes the consideration of plain objects against plain
backgrounds. The following facts have been observed :—

1. The greater the difference in the amount of light coming
from an object and background, the greater the distance
at which the object is visible.

2. A constant contrast in tone between object and back-
ground does not ensure a constant visibility. Dark
objects against light backgrounds are less visible than
light objects against dark backgrounds. ‘This difference
is greater at high illuminations than at low ones.

3. The larger the size of the object, and the greater the
concentration of its area, the greater the distance at
which it is visible.

It follows that a plain object will appear conspicuous against a
plain background when the contrast in tone between object and
background is great, and when the object is lighter rather than
darker in tone than the background, and when its size is great
and area concentrated.

(B) Patterned Objects against a Plan Background.

As long as the pattern of an object against a plain background
is visible, the object must be visible. It follows that visibility
will to some extent depend upon the blending distance of the
pattern.

The following factors were found to affect the blending distance
of patterns: lighting, contrast in tone between the components
of the pattern, size of the components and shape of the com-
ponents, and the relative size of the components.

1. Lighting.—Experiments showed that the better the illu-
mination the greater the distance at which the pattern was
visible. A pattern which by day appears conspicuous, on account
of the long distance at which it can be seen, at night may be
difficult to see (e. g., the Zebra).

2. Contrast in Tone between the Components.—The greater the
contrast, the greater the blending distance of the pattern. A
chequered pattern of black and white is visible at a greater
distance than one composed of two shades of grey.

3. Size of Components.—The larger the components the greater
the blending distance, all other factors remaining constant, as was
shown in my previous paper (Joc. cit.). If there be components
of more than one size, then the smaller will blend first and the
larger at a greater distance.

PATTERNS CONSPICUOUS IN NATURE. 393

4. Relative Size of the Components.—For any given pattern

there is a particular proportion of the components which gives
the greatest blending distance.

Text-figure 6.

6 77.”
il

272M.
47 IN.

A specimen of the patterns used in Experiment No. 7.
ExprertMent No. 7.

Candles 1 ft. 6 ins. apart. Objects 2 ft. 2 ins. from candles.

Background grey. Objects square, 9 sq. em. in size, divided horizontally into five
black and white stripes of 6 mm. (see text-fig. 6).

Pattern blends at

In No. 1 there is 2/8 black and 6 8 white ..... 24 feet 2 inches.
5. : Sei a Ne Tie ae aa ae
Sb eS 5 4/8 5 Ae Oia ak PAB on HI gg
er eee Gees See | ee 26) Tuan”
ase ee COP... | Ge nee Sle ole

39

A striped pattern is dealt with in the above summary; it shows
that where the amount of black to white, or white to black, is
very small, the blending distance is smaller than when there are
about equal amounts of the two components.

Referring also to experiment no. 9 (p. 397), it can be seen that
for the types of patterns here dealt with there is similarly a
particular proportion of black to white which gives the greatest
blending distance under the experimental conditions.

5. Shape of the Components.——It has been shown that the
visibility of plain objects depends upon the concentrations of
their areas. Experiments show that, similarly, the blending
distance of patterns is proportional to the concentration of the
components of the patterns: the more concentrated the com-
ponents, the greater is the blending distance, as is seen in the
following experiment.

EXPERIMENT No. 8.

The blending distance of black and white patterns, of which the components are of
the same size but of different shape. Experimental conditions: Candles 11 ins.
apart, and 2 ft. from glass plate on which patterns were fixed. ‘

Background behind glass plate. of grey paper, at such a distance that it is of th

394

same tone as the patterns after they have blended.

patterns used.

fa

gure (.

Text-fi

Pattern.
No.

> OE o9 bo

DR. J. C. MOTTRAM ON

Length of cireumference

of component.
28°3 mm.
30)
3255) 1,;
SERB) op
A275 ,
AO on

(Mean of six observations.)

; Text-fig. 7 shows the
The following table gives the length of the circumference of
the component and the blending distance :—

Blending distance.

These were uscd in Experiment No. 8.

Patterns whose components are of the sane size but of different shape.

The experiment shows that the smaller the cireumference of the component, and
therefore the greater its concentration, the greater the blending distance of the

pattern,

PATTERNS CONSPICUOUS IN NATURE. 395

With plain objects the circle is the most visible shape which

they can have; so with patterns, the circle gives rise to the
greatest blending distance.
Patterns composed of only two components require special
consideration, as they have an important bearing on the subject
of conspicuous patterns, as will appear later. It has been seen
that the larger the pattern the greater the blending distance; 1t
follows that, in order to give an object a pattern which will
blend at the greatest distance, only two components must be
used, thus making it as large as possible. To further increase
the blending distance the relative tones of the two components
must be as far removed as possible, and at least one of the
components must have as concentrated an area as possible.

Text-figure 8.

1-6. Patterns in which one component has the most concentrated shape, namely a
circle, and is surrounded by the other component. 7-9. Patterns in which
neither component has the most concentrated shape.

If the object be a triangle, a square, or a circle, then these
conditions would be fulfilled in figs. 7-6 of text-fig. 8. These
patterns would blend at a greater distance than would those
shown in figs. 7-9.

It has been seen that a third factor affects the blending
distance, namely, the relative proportion of the components.

Tf a series of circular objects be made, as in text-fig. 9, and be

396 DR. J. C. MOTTRAM ON

examined against a number of differently-toned backgrounds, it
will be found that the blending distances are not the same for
different backgrounds. If, for instance, they be examined
against a white background, then in the case of nos. 7-7 no
pattern-blending occurs: the objects appear as black spots. If,
instead, the background be light grey, then the white blends

Text-figure 9.

A series of eye-spot patterns used in the experiment*described on p. 401, and set out
in Table I. The uppermost disc is white, the lowest black. The others contain
from above down 4, & 8, 4, 2, 2, and 3th of white, either concentrated in the
centre, or in a ring round the periphery.

with the background and leaves the central black area visible
after the white has ceased to be distinguishable from the back-
ground, This difficulty can be overcome by joining up a number of
two-component patterns, and then finding the blending distance;
this has been done in the following experiment.

PATTERNS CONSPICUOUS IN NATURE, 397

Text-figure 10.

Portion of a series of spotted patterns used in Experiment No. 9.

EXPERIMENT No. 9.

The blending distance of black and white spotted patterns, as shown in above
figure. The percentage of white to black varied from 10 to 60 per cent.
Experimental conditions: Candles 1 ft. 6 ins. apart and 2 ft. 6 ins. distant
from the pattern, which was placed on a glass plate with a grey background
behind, as in experiment no, 8.

398 DR. J. C. MOTTRAM ON

The following diagram (text-fig. 11) shows the blending distance in feet, plotted
against the percentage of the spots to the whole pattern ; the X shows the white-
spot pattern, and the O the black centre. It can be seen that the white spot shows
the greatest blending distance, which occurs when the white is approximately
20 per cent. of the whole.

Text-figure 11

ry

go

20

*yaf uy gounzsip Suipusige

“
°

Percentage of central area.

lo zy) 30 Lo 59 60

Ordinates=blending distance of the patterns in fect. Abscisse=percentage of the
central components of the pattern. @-=the readings obtained when a black
centre pattern was used (text-fig. 10, upper series). X=when a white centre
pattern was used (text-fig. 10, lower series).

It can be seen that the greatest blending distance is produced
by a white-centre pattern ‘containing 20 per cent. of white, and
it is shown also that the white- centre pattern blends at a
greater distance than the black centre. It appears, therefore,
that the blending distance of patterns is affected in the following
ways :—

(1) By contrast in tone between the components: the greater
the contrast, the greater the blending distance.

(2) By size of components : the larger the size, the greater the
blending distance.

(3) By shape of components: the more concentrated the area
of the components, the greater the blending distance.

In order, therefore, to cover an object with a pattern which
will blend at the greatest distance: (1) make the tone of the
components a great contrast, black and white; (2) make the
size of the components as large as possible, by reducing their
number to two; (3) make the shape of the components as con-
centrated as possible, by making one of them a circle.

PATTERNS CONSPICUOUS IN NATURE. 399

The consideration of the visibility of patterned objects against
plain backgrounds can now be resumed. As before mentioned,
a patterned object is visible as long as its pattern is visible; it
follows that objects whose patterns blend at a great distance are
more conspicuous than taose whose patterns blend at a short
distance. :

There remains to be considered patterned objects which are
visible against plain backgrounds after the pattern has blended
at distance. pike

In a previous paper it was shown that, as regards visibility (as
measured by the greatest distance at which the object is visible),
patterned objects against plain backgrounds are neither more nor
less visible than plain ones. It was also shown that, where the
pattern interrupts the margin, the outline of the object appears
blurred and difficult to define after the pattern has blended at

°

. Text-figure 12.

ot

AAR

1-3. Patterns which do not interrupt the margins of the object. 4-6. Patterns
which interrupt two sides, three sides, and one side of the object.

distance. It follows that a further condition must be fulfilled in
order that a patterned object may be as conspicuous as possible
against a plain background: the pattern must present an un-
interrupted margin, for example, as shown in text-fig. 12, 1-6.
If these patterns be viewed from beyond their blending distance,
they will appear more defined than nos. 4, 5, and 6, in which the
pattern interrupts the margin. Apart from this, the visibility of
patterned objects beyond the blending distance and against plain
backgrounds is similar to that of plain objects against plain
backgrounds. A patterned object can be made more conspicuous
_ against a series of plain backgrounds than can a plain object,
because, though a plain object can be made very visible against a
single plain background by a strong contrast in tone with the

400 DR. J. C. MOTTRAM ON

background, nevertheless, when a series of backgrounds are used,
then the object will appear inconspicuous against those similar, in
tone. On the other hand, with a patterned object, when the
background is similar in tone to one of the components, then the
other will make a strong contrast and cause the object to be
easily visible.

It has been seen that the greatest blending distance which a
pattern can have is one in which the white or lighter component
is concentrated in the form of a circle, and that an object is most
conspicuous against a wide series of backgrounds when it presents
a pattern of only two components. Experiments were therefore
made to discover which of two component patterns, the light-
centre one or the dark, is the more conspicuous against a wide
series of backgrounds. A series of discs were made, as shown in
text-fig. 9, and examined against a series of backgrounds in the
following manner :—-

Text-fig. 1, B (p. 385) gives a ground-plan of the experimental
conditions ; it can be seen that the backgrounds are illuminated
separately from the discs, which are fixed to a glass plate. By
moving the backgrounds towards or away from the light, a
continuous and wide range of tone in the background can be
obtained. First, the background of white paper was moved so
as to exactly match in tone that of the white in the discs.
Under these conditions the all-white dise was invisible; of the
rest, the dise with a white centre, 7/8 of the whole, was found to
be the least visible, and next the dise with 6/8 white centre.
The most visible dise was the all black.

The discs were examined in a similar manner over a wide
series, and in each case the three least visible discs and the most
visible were noted ; the following table gives the results.

It can be seen that, except against backgrounds lighter than
the white in the dises, the black-centre eye-spots are less visible
than the white, and the appearances of the discs as seen from a
distance show that the white-centre discs are the more visible.

The greater visibility of the white-centre eye-spots is especially
marked when the backgrounds approach the dark end of the
series; and as, as already shown, the backgrounds in Nature are,
for the most part, of dark tone, it follows that white-centre eye-
spot patterns of two components must be more conspicuous than
black-centre patterns. Similarly, the white-centre pattern must
be more visible than any other combination of black and white,
because in this pattern the white is most concentrated, and there-
fore has the greatest visibility and the longest blending distance.
Tt follows that against a series of plain backgrounds, and especially
a series of relatively dark tone, the white-centre eye-spot pattern
is the most visible one that an object can have.

401

PATTERNS CONSPICUOUS IN NATURE.

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402 ' DR, J. GC. MOTTRAM ON.

The most conspicuous pattern that an object can have against
a series of plain backgrounds may now be defined :—

(1) The pattern must consist of wo components.

(2) The components must differ widely in tone (black and
white).

(3) The lighter component (white) must be concentrated at
the centre in the form of a circle.

(4) The darker component must surround the white so that
nowhere is there an interrupted margin.

(5) If the series of backgrounds be of low tone (as they are in
Nature) then there must ue more white than black in the
pattern.

(C) Plain Objects against Patterned Backgrounds.

(1) If the object is visible after the pattern has blended at
distance, then the factors which control its visibility are the
same as those of plain objects against plain backgrounds. The
outline of the object will, however, appear blurred just as when
a patterned object, whose pattern is interrupted at the margin,
appears blurred when viewed against a plain background. The
conclusions are similar, the only difference being, that in one
case the object is plain and the background patterned, whereas
in the other the object is patterned and the background plain.

(2) If the object becomes invisible before the pattern of the
background blends at distance, then the object may be seen
against one component of the background; in which case the
factors controlling visibility will be similar as are those of plain
objects against plain backgrounds, except that the near presence
of an area of different tone will affect the visibility of the object.
A black square on the white component of a checkered back-
ground will be less visible than on a plain white background
(see experiments, nos. 2 & 3). With this exception, the factors
controlling visibility are similar to those of a plain object against
a plain background. The object may be visible against two or
more components of the background. If the object is of the
same tone as one of the background components, then the object
will appear as a projection from the margin of one component.

Experiments were carried out to discover whether the factors
controlling visibility were different from those of a black object
against a white background, and it was found that, except for
the decrease in visibility due. to the presence of an area of black
(in this case touching the object), the visibility was similar to
that of a plain object on a plain background, except that the
effect of shape of the object was rather different from its effect
when dealing with plain objects against plain backgrounds, as
seen in the following experiment *.

* This consideration has an important bearing on concealment b, indented or
scalloped margins; a series of experiments has been carried out from this point of
view which, however, are only of present interest in so far as they show that an even
margin is a "factor for conspicuousness.

PATTERNS CONSPICUOUS IN NATURE. 403

Text-figure 15

Objects of the same size but of different shape lying at the junction of a black and
white background as used in Experiment No. 10.
ExpreriImMent No. 10.

The visibility of black objects projecting from the margin of a large black mass.
The objects are all of the same size but of different shape, as shown in the text-

figure.
Experimental conditions: Candles 1 ft. 3 ins. apart and 3 ft. 6 ins. from objects.

Object no. 1 visible at 57 feet.

be) oP) 2 99 54 rh}
2 7° 8 2” 49° ,,
A 3 32s

39 bb)

-Compare with Experiment No. 5.

The object may be visible against two or more components of
a patterned background, and may be different in tone from

either component.
Text-figure 14.

/ Ld J

1. A grey disc placed over the junction of a black and white background. If the
dise approaches in tone more nearly the white of the background than the black
then from a distance it will be seen as in no. 2; if more nearly the black, then

as 1n no. 3.

If a grey disc be placed over a black and white junction, as in
text-fig. 14, and be viewed from gradually increasing distances,

404 DR. J. C. MOTTRAM ON

a point will be reached at which one half of the disc is blended
with one component of the background, whilst the other half is
seen as projecting into the other component. If the grey disc
more nearly approaches the black in tone than the white, then
at a distance it will appear as in text-fig. 14, 3; if it more
nearly approach the white, then as no. 2.

The following experiment illustrates this appearance :—

ExPERIMEnT No. 11.

Candles 1 ft. apart and 2 ft. distant from background.
Background half black and half white (7 X 5 ins.).

Objects circular, 20°4 sq. mm., eight in number and ranging in tone from black to
white. Objects placed exactly over the junction of the black and white of the

background.
Visibility distance.
: Blackman. a feet. isuerien Visible as a dark
asp we arg : projection into the white of
a ERY oon one eye Be the background.
o 5B) Ey OG Sa0 bad
5 5 shige 18), 5. 5, Visibl cI
eas a light
3 Light grey oy ” 3 2 projection into the black of
8. White 2 SRO ics the background.
ADODBOOaS oy) oy)

An object was prepared of such a grey tone that it was neither seen as a white
nor as a black projection into the background’s components. This object was the
least visible disc and visible at 16 ft. 4 ins.

Some similar experiments were carried out with backgrounds
composed of different tones of grey instead of black and white,
and it was found that the objects likewise appeared as projectors
into one or other of the components, according as to whether
the object more nearly approached in tone one or other of the
components.

If, for instance, the background was made of two dark grey
tones, then only the darkest objects appeared as black projectors
against the lighter of the two components.

As in Nature backgrounds are:dark in tone rather than light,
it follows that light grey or white discs will be more visible
under these conditions, 7.¢., when seen against two or more
components of a patterned background, than dark discs.

As mentioned ‘in the last experiment, there is one tone of grey
against which black and white are equally visible, and when the
object is of this particular tone it never appears as a projective
from one component on to the other. Against backgrounds
composed of tones other than black and white, there is similarly
one grey tone which the object may have which will cause it to
give a similar appearance.

An experiment was carried out with discs of this tone to
discover whether the effect of size of the object was similar or
not to that found when dealing with plain objects against plain
backgrounds. As seen in the following experiment, the effect is
similar :—

PATTERNS CONSPICUOUS IN NATURE. 405

EXPERIMENT No. 12.

Candles 1 ft. apart and distant 2 ft. from background.
Background as in Experiment No. 11.

Objects of various sizes were made of grey discs of such a tone that they were seen
neither as black nor as white projections into the components of the background.
These objects were placed over the junction of the black and white components
of the background. The following table gives the visibility of the discs :—

Size of Distance at which
grey disc. visible.
419 sq. mm. 27 feet 5 inches.
302 =, Bi iis Obes
20°4 ” 19) 33. 5 2s)
6-1 2 9 22 1 ”

The effect of shape was found to be different, as in the case
of objects of the same tone as one of the components of the
background, (See experiment no. 10.)

Text-figure 15.

3 by

Objects placed on more than two components.

A few observations were made with objects placed in front of
more than two components, as in text-fig. 15, 7 & 4, but no special
difference was noted from those when only two components were
covered by the object.

Another arrangement of an object against a patterned back-
ground remains to be mentioned. The object may more or legs
resemble, both in tone, in shape, and in position, one of the
components of the background; in this case, though it may
appear to be invisible on account of this similarity and thus
to come under a separate category, nevertheless this is not so,

Proc. Zoou. Soc.—1916, No. XX VII. aT

406 DR. J. C. MOTTRAM ON

as it will fall naturally under one of the conditions already
considered, For instance, a very light grey object may resemble
the light square of the checkered background, both in shape, size,
and position, as in text-fig. 15, 2; but it can also be considered
under the heading ‘“ Objects falling on one component of the
background.” If it falls as in text-fig. 15, 3, then on two com-
ponents of background ; if as in no. 4, then as an object visible
after the pattern has blended at distance.

It appears, therefore, that a direct resemblance does not
necessitate a separate consideration.

Conclusions.—The factors which make for the conspicuousness
of plain objects against patterned backgrounds appear to be
similar to those when plain backgrounds are used. If the
pattern of the background interrupts the object’s margin, then
outline blurring occurs. The near presence, or contact with
the object, of an area of tone similar to the object makes it less
visible.

(D) Patterned Objects against Patterned Backgrounds.

If a patterned object be viewed against a patterned background
from gradually increasing distances, several different appearances
may be seen.

(1) The pattern of the object may blend before the pattern of
the background, in which case it will appear as a plain object
against a patterned background.

(2) The pattern of the background may blend before the
pattern of the object. The object will then appear as a
patterned one against a plain background.

(3) The patterns of object and background may both blend
and yet the object may still be visible against the background,
in which case the object will appear plain against a plain
background.

These three conditions have already been dealt with.

(4) A fourth appearance may occur. The object may not be
visible, although neither its pattern nor the pattern of the
background have blended.

Before dealing with this appearance the first three must be
briefly considered. It has already been shown in my previous
paper (loc. cit.) that though a patterned object is not less visible
than a plain one, nevertheless, if the pattern interrupts the
margin, then its outline after pattern-blending appears blurred
and indistinct, as compared with a plain object of the same tone
as the patterned one after blending.

It has also been mentioned (p. 402) that outline blurring
gimilarly occurs when the object is plain and the background
patterned. It might therefore be concluded that against
patterned backgrounds the outline of a plain object would appear
just as blurred as that of a patterned one beyond the blending
distance of the background’s pattern; but when both the object’s

PATTERNS CONSPICUOUS IN NATURE, 407

pattern and the background’s pattern are blended, and provided
that both interrupt the junction of object and background, then
the blurring effect of these two interruptions are added, and
‘eause the junction of object and background to appear much
more indistinct than when only one pattern interrupts.

In text-fig. 16, if in each case the insects remain visible after
‘the patterns are blended, then the outline of the patterned one
will be the more indistinct when the tone of the patterned insect
after pattern-blending is the same as that of the plain insect.

Text-figure 16.

Visibility of insects on different backgrounds.

It follows that animals with a pattern which interrupts the
margin will be less visible than plain animals against a patterned
background as well as against a plain one; and further, that
against a patterned background an uninterrupted margin will be
as necessary for conspicuousness as against plain backgrounds ;
an fact, experiments appeared to show that it was more necessary,
because the blurring effect of the background pattern required to
‘be counteracted.

In Nature, a pattern which interrupts the margin must be a
great aid to concealment against patterned backgrounds, because
the backgrounds are irregular and the animal must often be seen
with one or more of its margins against a single component of
the background, as shown in text-fig. 16.

The conclusions as regards conspicuousness which have al-
ready been made must thus apply to patterned Queer against

27

408 DR. J. C. MOTTRAM ON

patterned backgrounds. There remains to be examined only
the fourth condition, in which the object cannot be seen though
both the object’s and the background’s patterns remain visible..
This invisibility is due to a great similarity between the patterns.
of object and background in size, shape, and relative tone of the
components. It is obvious that the most conspicuous pattern
will be inconspicuous against a background made of a similar,
or closely similar, pattern. For this reason, under these special
conditions, it is not possible to define a pattern which will be
especially conspicuous, unless the pattern of the background is
also defined.

It may be concluded, therefore, that the pattern which has
been called the, white centre eye-spot pattern is a most con-
Spicuous one against every background with the exception of
backgrounds themselves composed of eye-spot patterns. It follows.
that in Nature the white eye-spot pattern must be very con-
spicuous, provided that this type of pattern is not continuously
found as a background.

White centre eye-spot patterns in Nature.—Natural backgrounds.
were examined in order to discover whether this type of ‘pattern
was to be found and to what extent, and it was at once noticed
that they were very uncommon; a morning’s foray amongst
woods, fields, hedgerows, and broken country resulted in only
a few examples. It is not difficult to make such patterns.
artificially out of doors; for instance, by laying round white
stones on circular patches of dark moss, by placing shining leaves.
over dark rough ones, by viewing pierced leaves against the light,
and in many other ways. Natural eye-spot patterns may be
conveniently described under the following headings :—

1. On bare ground.—(a) Due to irregularities of the surface.
Working with plasticene the pattern can be produced by
shallow conical pit with a flat bottom, by a truncated cone lying
on its base, or by a cylinder standing in the middle of a cylindrical
depression ; in each case top lighting 1s necessary. Viewed from
above, a light centre dark-margin circular pattern is seen; the
pattern. does not perfectly reproduce the one desired, because
the centre instead of being lighter in tone than the background
is either of the same tone or somewhat darker. It is evident
that by artificial methods this pattern can only be reproduced
with ditfticulty, and thus its occurrence in Nature must be very
rare. By prolonged search isolated examples are to be seen.

(b) Due to the surface being of broken tones; a light stone
or one reflecting the light from the sky when lying on a circular
dark patch will give rise to the pattern. Examples of this
nature are not difficult to find, but they are never numerous
avd always isolated.

On grass and other short vegetation.—Except for flowers,
rina are considered elsewhere, the eye-spot pattern is very
rarely seen ; occasionally light reflected from a shiny leaf supplies
an example.

PATTERNS CONSPICUOUS. IN NATURE. 409

3. Rank vegetation—Light from a shiny leaf, or a leaf in
strong light against shadow, occasionally forms the pattern.

4. Scrub, hedgerows, and wood margins.—Circular leaves in
very strong light against dark shadows often give rise to a
pattern which is somewhat like a white centre eye-spot one,
but distinction must be made between a pattern consisting of
light spots on a dark background and the pattern under con-
sideration. The first is common in Nature ; the second requires
a light centre, a dull margin, and a background of a different
tone. Occasionally this arrangement is to be seen among
vegetation, but only isolated examples are to be found.

5. Light woods. — Here are sometimes to be seen patterns
similar to those described under the previous heading. Where
sunlight penetrates through foliage and falls on dark ground
or foliage beneath, white spots of light result, and when these
happen to fall on dark objects they produce the white centre eye-
spot pattern. According to the frequency with which they happen
to fall on dark objects is the prevalence of the pattern. When
the ground beneath the trees is much broken in tone, several
may be seen from a single station. Several conditions are,
however, necessary for their production—an uncovered more or
less vertical sun, a not completely dense canopy of foliage, and a
broken ground beneath.

6. Heavy woods.—lf the fohage be not too dense the pattern
may be produced as described in no. 5.

7. Sky.—On looking up at the sky through foliage, white
spots are to be seen in the intervals between the leaves and
where there are holes in them. If a white spot happens to be
surrounded by a dark shadow or a deep-toned leaf, then a white-
centre dark-margin eye-spot results. A small number of these
are always to be-seen. It may be pointed out that though man
is not accustomed to view foliage in this way, many animals of
low stature and whose eyes are set looking upwards as well as
forwards must frequently take this view.

8. Water—Very small pools of water when they reflect the
sky and when, as is often the case, they are surrounded by a
ring of moist and therefore dark-toned ground, have the appear-
ance of the eye-spot pattern. Foliage overhanging water or
floating upon it also rarely gives rise to the same pattern, the
sky reflected from the water forming the white centre and the
foliage the dark ving. Drops of water and dew’ under some
conditions of lighting give rise to an abundance of the pattern
of a transitory nature.

9. Flowers.—By far the most common examples in Nature
of the eye-spot pattern are to be found in flowers. A dark
centre eye-spot is as common as a light centre. There can
hardly be a doubt that flowers are purposely conspicuous ; it is
therefore noteworthy that their patterns conform to the rules
which experiments have decided must be followed in order that
a pattern may be conspicuous in Nature.

410 DR. J. GC. MOTTRAM ON

Flowers are, as arule, circular ; their patterns consist of seldom
more than two components, one being concentrated in the middle-
in the form of a cirele, and there is usually a strong contrast in
tone (and colour) between the two components.

It might be thought that these arrangements of pattern in
flowers were due to convenience of growth; but the eccentric
shapes and patterns assumed where special animals are sought
for the purpose of fertilisation indicate that flowers are not
forced by growth to assume the circular shape and eye-spot.
pattern.

In conclusion it may be said that, except in the case of flowers,
white-centre dark-margin eye-spot patter ns are rarely to be seen
in Nature and are almost always isolated. Sunlight penetrating
through foliage on to broken ground and sky views through
foliage are the two most common causes. As regards flowers,.
eye-spot patterns are very common, but the centre is as often
darker than the margin as vice versa.

It follows that animals presenting this type of pattern must.
be considered to be conspicuous in Nature.

Part II.

Having by experimental methods defined the types of pattern
which render an object conspicuous, attention was turned to the:
animal kingdom to discover whether examples of these types.
could be found and, if present, what was their distribution.
Search was made among the Lepidoptera because their wings offer-
a plain, flat patterned surface, and thus the complicating factor |
of solidity is avoided. Rather than search through a large
amount of material, it was decided to deal thoroughly with a
definite amount, viz., the Indian Lepidoptera. Moore’s ‘ Lepi-
doptera Indica’ was the work chosen, because of its good coloured
illustrations of each species. On glancing through these plates.
several types of pattern were found which previous consideration
showed would render these insects conspicuous. The first type:
to be dealt with is shown in text-fig. 17. It can be seen that
the pattern consists of a central white, or light yellow, area
surrounded by a black margin, so that the four wings combined
present an irregular, white-centred, black-mar eined pattern.
The margin of the Wings is, except in two cases (nos. f and 4),
not scalloped. The black marginal band is sometimes broken
by onal spots or bands of light tone, but only in the case of
no. 2 is the margin interrupted by patter n.

This type of pattern presents, therefore, those characters which
previous consideration has shown must render the insect con-
spicuous in Nature: the table on p. 412 gives its distribution
among genera of the Indian Lepidoptera

Salatura (text-fig. 17, 7) and Acidalia (no. 2) do not conform
to the type in several respects. In Salatwra the centre white

PATTERNS CONSPICUOUS IN NATURE. 411

area is broken up by dark bands, and in Acidalia by black spots,
besides which there is a half-tone area at the centre of the wings.
They are introduced for several reasons, as will appear later.
Certain butterflies are presumed to be protected from the attack
of enemies by ill-flavour ; further, it has been noted that these
insects are conspicuous in Nature (and it has been suggested that
they are conspicuous in order to warn enemies) ; and lastly, it has
been noted that the pattern and coloration of these insects are

Text-figure 17.

Types of all the genera illustrated in ‘ Lepidoptera Indica’ which present patterns
of the first type under consideration.

1. Cethosia. 2. Acidalia. 3. Catopsilia. 4. Elymnias. 5. Apatura. 6. Appias

& Huphina. 7. Salatura. 8. Pareha. 9. Catophaga. 10. Limnas. 11.

Eurymus. 12. Kibreeta, Nirmula, & Terias. 13. Ivias. 14, Hyposcritia.

15. Anapheis. 16. Telchinia. 17. Chrysophanus. 18. Stiboges. 19. Daimio.
20. Callosune.

mimicked by insects which are not thus protected by ill-flavour,
in order that they may gain protection by means of a false
cloak. The pros and cons of this contention cannot be discussed
here, but it is remarkable that many of the insects presenting
the type of pattern under consideration belong to what are con-
sidered to be protected genera, or to what are considered to he

412

DR. J. C. MOTTRAM ON

TaBLeE IJ, .
| ls Meg
ob
| |B a
| a o-5
Family. Subfamily. Genus. | 2 || ai : | ae
| [S|] 2) e\s8
oO x =! me ost Ibs Be
| c= fi |S iios | 2 be 8)
s 3) iS) = Rsv PUES
Ba B/A/e |e 4 |
Nymphalide. | Eupleine. Limnas ........... «S| Xx |
Salatura .........| ood x || X
Elymniine. Elymnias .........| Dai pene lies 5X
Nymphaline. Apatura .......-.. x x
— Sy ——s | —— ||.
Argynnine. Cethosia ......... ees
Acidalia x Socal | X
Acreinee. TEHRAWE proton bno..oe | x | x
Telchinia ........ | xX || X
Riodinide. Nemeobiine. Stiboges ......... | x | acelPos
Pieride. Pierinz. | Anapheis seg aueee | x || x
ZY DIGS sosroccomooe | X | | | X
Huphina ......... | | X || | | x
Hyposcritia ...... | Se os | meee bee Sor
Catophaga ...... x || | x
Coliine. Kibreeta ......... Pe S< iil xx
INO EPOMG ~ ono noncne esses line | Xx
Terias Aigte| Mell cee | | x
Catopsilia ...... | x | Sool | xX
ELE UGS enone hese x | gon || cca |] 2S
Callosume ......... a SG ill cco ison |] 2S
| Hurymus ......... x | | xX |
Lycenide. Lyczenopsine. | Castalius ......... | x | x |
—— \ sn lkie ae
Chrysophanine. | Chrysophanus .... X | | x |
Hesperiide. Celeenorrhinee. IDO seooedeeaooal| one | xX | x |
5 iB 24 18) | se) |. Ne a6 |

unprotected insects mimicking protected.

or mimicking.

Tt can be seen that
out of 24 genera forming Table IT., 5 are described by Moore as
protected and 3 as exhibiting mimicry ; whereas out of all the
600 genera described only 41 are mentioned as either protected

No reason can be given why the other 16 genera

(for the most part belonging to the Pieride) presenta conspicuous

pattern *.

It is, however, noteworthy that the sexes are alike,

with the exception of Hurymus and Chrysophanus, where the

pattern is confined to the male.

Several of the genera are

amongst the commonest of butterflies, and at certain times collect

* The Pierine and Coliinz are considered by some observers to be “ protected ”

insects.

PATTERNS CONSPICUOUS IN NATURE. 413

together and migrate in immense swarms. Salatura was intro-
duced into this table because it shows a considerable resemblance
to the next type to be considered. ‘ithe bands of dark tone
crowning the central light area have been drawn too boldly and

of too dark a tone, which makes the resemblance closer than it
really is.

Text-figure 18.

"Types of all the genera illustrated in ‘ Lepidoptera Indica’ which present patterns
of the second type under consideration.
1. Parhestina. 2. Parantica. 3. Orinoma. 4. Caduga. 5. Penthema. 6. Par-

anticopsis. 7. Delias. 8. Calinaga. 9. Cadugoides. 10. Caduga. 11.

Neurosigma. 12. Metaporia. 13. Hestina. 14. Radena. 15. Prioneris.
16. Bahora.

Examples of the second type of pattern which must render
the insect conspicuous in Nature are shown in text-fig. 18. It
can be seen that the insects present an uninterrupted margin,
the pattern nowhere reaching the margin, and that at the
margins there is an area of dark tone, whilst the centre of the
wings is much lighter in tone. As before, there is no scalloping
or irregularity of the margin. It follows that this pattern

414. DR. J. C. MOLLTRAM ON

conforms to the factors which have been considered to make for
conspicuousness. ‘Table III. shows the distribution of this second
type of pattern. At a short distance the central patterned area
will become blended and give rise to a light grey tone, and the
insect then has an appearance similar to the type first considered.
As before, it can be seen that out of 18 genera, 7 are protected
and 7 mimic: in this case, therefore, the conspicuous pattern
is accounted for in the case of 14 out of the 18 genera; 4 remain
unaccounted for. It is noteworthy that in all cases the sexes are
alike.

Tasxe IIT.
| | } |
rd
® .
use
| | | | Ec
~ 6 5 | | | te, 5 =
Family. Subfamily. Genus. fe hu ney ae ems
| | Be reas) ese ay t=
| litres oS a | Sos Ish
| Penis) 2 ae sel
| [=e A RB ae
Nymphalidz. | EKupleine. TEOMEDG 00 cha 008 800 ests ee eXeotl |
Tirwmala ......... Wecoliane eso
| IETORE teonan acopoe OX xX
Parantica......... XG 2X |
Caduga ... 1.021... oe 2S |) on |
Satyrine. | Orinoma ......... | xX] | xX
Elymniine. | Melynias ......... | x x
Nymphaline. | Hestina............ “98: jf ono] 2S" x
Parhestina ...... Bis lattes os | xX |
Neurosigma ...... | xX || | 38
Penthema ......... | | Xx jf x
Calinagine. | Calinaga ......... | &< |
Papilionide. Papilionine. | Cadugoides Are | X || ex |
Paranticopsis ... x | x
Pieridee. Pierine. | Metaporia ...... sos gta eR [Poa |
DY ATER eiseaearate Penton ll ace Ito als
Prioneris ......... mies |X FER OOM
Eroniine. Pareronia ...... | | x Gilt same ils ons
SS || oe eos | fe |
3 | 8 | 18 Farolan tape coat) retiree lla
| \ |

Conspicuous pattern combined with an absence of secondary
sexual dimorphism is so frequently associated with a protected
species, that-attention must be drawn to the fact that some of
the unaccounted-for genera in both tables may be protected
genera, although not mentioned as such in the work consulted.

Finally, all the insects which are mentioned by Moore as being
either protected by ill-flavour or mimicked by other species, are
briefly considered in order to see whether or not they present

PATTERNS CONSPICUOUS IN NATURE. 415:

patterns which experimcnts have shown must be conspicuous in
Nature *,

Text-fig. 19 shows their patterns. No. 3 is like Salatura, a
stage between the first and second types. There is, however, an
absence of a defined dark margin to the hind wings, the margins.
are not scalloped, and the pattern does not interrupt the margin,
though it approaches near to it; it thus presents some of the
characters making for conspicuousness.

Text-figure 19. _

“se

Types of all the genera mentioned in ‘ Lepidoptera Indica’ as being “ protected”

beyond those already given in text-figs. 17 & 18.

1. Hestia. 2. Menama. 8. Piecarda. 4. Bimbisara. 5. Calliplea. 6. Con-
dochates & Neptis. 7. Euplea & Pademna. 8. Stictoploea. 9. Penoa &
Crastia. 10. Cynitia. 11. Danisepa (the dark tone of this imsect should
be darker). 12. Isamia. 18. Stabrobates. 14. Libythea (? protected). 165.
Hrgolis.

Nos. 2, 5, 7, 8, 9, and 12 are conspicuous in so far as they
present a large, dark, unpatterned area; their margins are not
scalloped or interrupted by pattern ; the marginal spots, when

* Mimicry within the Papilionine is only referred to once in an indefinite
manner.

416 DR. J. C. MOLTRAM ON

present, would, however, tend to mask the outline. As to whether
or not these insects are conspicuous in Nature must depend upon
the tone of the backgrounds against which they are commonly to
be seen; if the backgrounds be light in tone they would be
conspicuous insects, but if the insects lived in dark forests, for
instance, they would not be especially conspicuous.

No. 17 is similar to the last except that both wings present a
large white patch which must make the insect more conspicuous.
The patch on the fore wing interrupts the anterior margin and
must therefore have the opposite effect.

No. 10 presents a black-centre white-margin pattern which, as
has been seen, is almost as conspicuous as the white-centre black-
margin pattern.

Nos. 4, 6, 13, & 14 present patterns which do not interrupt
the margin but, instead, follow it; there are three central bands
or rows of spots which are srmomadal by black, and the margins
of the wings are not scalloped, thus several factors making for
conspicuousness are present.

In no. 7 the pattern everywhere interrupts the margin, and the
margin of the wing is not scalloped. The pattern is not therefore
a conspicuous one, * the general tone of the insect is light and the
wing-expanse large ; thus, i in spite of an inconspicuous pattern,
the insect might be conspicuous if its natural environment were
of dark tone—if, for instance, 1t were a forest insect. It may be
noted that another species of the same genus (see text-fig. 20, 4)
presents a typical conspicuous pattern.

No. 15 presents no character making for conspicuousness; the
margin is somewhat scalloped, the pattern interrupts the margins,
the insect, as drawn, is coloured a middle brown with a darker
line pattern, and is mimicked by Rohana parisatis. With the
exception of this genus and Hestia, the patterns of these pro-
tected or mimicked insects all show one or more characters
which make for conspicuousness, and present patterns much less
perfectly conspicuous than the two types first dealt with. The
first type conforms very closely to the pattern which experimental
consideration indicates must be the most conspicuous. Even the
larger proportion of black to white tone in the pattern conforms ;
as the backgrounds in Nature are for the most part dark rather
than light in tone, so there should be a greater proportion of
white to black in the pattern. It is not possible to show why
less perfect types are to be found; perhaps they present a stage
in the evolution of the conspicuous patterns, or that for some
reason a more perfect pattern is not required by these insects.
‘On referring to text-figs. 17 and 18 it can be seen that the
mimicking species present patterns which are not so perfect as
the models. Acidalia, for instance, could with justice be removed
from the first series.

As a contrast to these patterns four inconspicuous patterns are
shown in text-fig. 20, q, 2, 3,5; it can be seen that in three the
margin is scalloped; in nos. 7 and 3 the pattern interrupts the

PATTERNS CONSPICUOUS IN NATURE. 417

anterior and lateral margins; in no. 9 it interrupts the anterior
margin. In no. 2 the pattern of eye-spots and irregular bands
is confined to the outer margins of the wings, whereas the rest
of the wings is of an even dull tone (brown); in nos. 7, 3, and 5
the pattern is likewise more or less confined to the margins,
leaving the centre of the wings plain; in no. / the outer margin
is fringed by outstanding scales which cause the margin to.
appear indistinct. Unprotected butterflies show, as a rule, one
or other of these and other characters which cause their outlines
to blend into their surroundings, the pattern is confined to the
wing margins and it interrupts the margin, and the margin is.

Text-figure 20.

SS

Insects with inconspicuous patterns.

1. Pontia daplidice Q. 2. Anadebis himachala 6. 3. Lethe neelgheriensis 2.
4. Hestia hadenii 2. 5. Pazala sikkima 2. 6. Hestia malabarica 2.

scalloped. On the other hand, those insects which have been
considered to present conspicuous patterns show none of these
characters: their outlines are not scalloped, their patterns are
not especially confined to the margin and do not interrupt the
margin.

Finally, it may be said that whilst the inconspicuous pattern
of insects conceals their outline, the silhouette of an insect
against its surroundings (the patterns may or may not mimic the
backgrounds), the conspicuous pattern accentuates the margin.

418 DR. J. C. MOTTRAM ON

CoNcLUDING REMARKS.

Many experiments and observations have shown that the
patterns and coloration of animals are related to their environ-
ment; such terms as Protective Resemblance, Obliterative
Shading, etc., indicate the lines of research along which know-
ledge has been acquired, and which is conveniently condensed in
the following table by Prof. Poulton. The basis of this classifi-
cation is a resemblance, or otherwise, of the animal’s coloration
to its natural background. Further differentiation is achieved

TABLE LV.

A. Apatetic colours = colours resembling some part of environment.
(1) Cryptic (a) procryptic = protective resemblance.
(6) anticryptic = aggressive resemblance.
(2) Pseudo-sematic = false signalling.
(a) pseudo-sematic = protective mimicry.
(6) pseudo-episematic = aggressive mimicry or alluring.
B. Sematic colours = signalling colours.
(1) Aposematic = warning.
(2) Episematic = recognition marks.

by division according to the utility or function which this re-
semblance, or the reverse, has. These functions have to do with
the escape from enemies, the procuring of food, and recognition
by members of the same and other species. It follows that the
patterns of animals must be closely related to the visual percep-
tion of their enemies, their prey, and their friends. A classifica-
tion from this point of view would seem, therefore, to be the
most natural, and the following table was therefore prepared.

TABLE V.
pees ie for the absorption of Light Rays:
| ai ene f for the absorption of Heat Rays:
eBence Duro nS, Excretory products ete.
| other animals :
| to the eyes of _ * 5 NO.
| enemies = protective coloration.
Inconspicuous | PRs ae Teer ae Io tee ae
| to other 0 nhereyes of = aggressive coloration.
| c : prey —
| animals : A
| totheeyesof _ 5
Colour | laa a friends fae
and 1€ ee (1 ann =i eit ae
| perception o : cain 3 3
Pattern. Bikes aaa to the eyes of _ attr acting and repelling.
enemies and warning coloration.
| Conspicuous | tothe eyes of _ to allure prey as in
| boreune: rey - Mantid
| animals : prey : S
to the eves of _ social signals,
| friends ~ sexual signals.
|

PATTERNS CONSPICUOUS IN NATURE. 419

If the consideration of pattern from this aspect be of value, then
an experimental analysis carried out with artificial patterns and
the human eye must be a sound foundation for the study of the
subject, at any rate, as regards the visual perception of mammals,
provided the human eye is not widely different from that of
mammals as a whole. The results of this line of investigation
show that patterns of animals will bear such an intense study,
and indicate that many details of pattern may be of value
although they have, up to the present, and on negative evidence,
been considered to be unrelated to the visual perception of their
own and other species. ,

In view of the fact that sight is a most valuable organ of
perception, and therefore a most powerful weapon in the struggle
for existence, it follows that a study of pattern from this point
of view is likely to throw light on some of the important problems
of Nature.

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MISS DOROTHEA BATE ON FOSSIL VERTEBRATES. 42]

14. Ona small Collection of Vertebrate Remains from the
Har Dalam Cavern, Malta; with Note on a new
species of the genus Cygnus. By Dorotuma M. A.
Batt, Hon.M.B.0.U.*

[Received April 14,1916: Read May 9, 1916. ]
(Text-figures 1 & 2.)

INDEX.
Systematic : Page
OMG DES CHOWBUTD, 30s Ts sein ococedoahoos seneda oodsannoobboonsssccossooee LA

The researches of Spratt and Leith Adams, and later those of
Dr. Cooke and Mr. Tagliaferro, on the extinct fauna of the
Pleistocene cave-deposits and fissures of Malta have already
yielded a rich harvest. The excavations of the two former
extended over a long period. Dr. Leith Adams, for instancef
spent six years in the island, a great part of this time being
devoted to investigating its cave and fissure depositst. That
there is still scope for yet further research is shown by a col-
lection lately sent for examination to the British Museum
(Nat. Hist.) by the Curator of the Malta Museum. This task
was very kindly entrusted to the present writer by Dr. A.
Smith Woodward, F.R.S.

The literature dealing with the subject is very scattered, and
the records extend over a great number of years. Therefore,
before making a few observations suggested by a study of this
collection, it has been thought useful to workers on the paleeon-
tology of the Mediterranean Region to give as complete a list as
possible of the vertebrates of which remains have been obtained
from the Pleistocene of Malta. Leith Adams published a similar
list in 18774, but this contains a record of only twenty-one
species, although Hlephas falconert and Myoxus cartei are in-
eluded. It is gratifying to find that this number has since been
nearly doubled, although no extensive systematic excavations
have been carried on.

List oF SPECIES.

MAMMALIA.

1. Ursus arctos (¢) Linn.

2. Vulpes sp.

3. Canis sp. (size of C. lupus).

4. Leithia melitensis Leith Adams sp.

* Communicated by Dr. A. SmitH WoopwarpD, F.R.S., V.P.Z.S.

+ Nat. Hist. & Archzol. of Nile Valley and Maltese Islands, Edinburgh, 1870.

+ Quart. Journ. Geol. Soc. vol. xxxiii. 1877, pp. ee —“On gigantic land-
tortoises’. _ trom the Ossiferous Caverns of Malta . . together with a list of their
fossil fauna.”

Proc. Zan, Soc.—1916, No. XXVIII, 28

422, MISS DOROTHEA BATE ON

5. Hliomys sp.

6. Arvicola amphibius Linn.

he » pratensis Baillon.

8. Hqwus sp.

9. Cervus dama (¢) Linn.
10 ., elaphus var. barbarus Bennet.
11. Hippopotamus pentlandi Meyer.

12. % melitensis Forsyth Major.
13. Hlephas mnaidriensis Leith Adams.
14, » melitensis Falconer.

AVES.

15. Strix melitensis Lydekker.*

16. Hutolmaétus fasciatus Vieiil. sp.
17. Gyps melitensis Lydekker.t

18. Anser sp.

19. Branta leucopsis Bechst. sp.

? 20. », bernicla Linn. sp.
21. Cygnus faleonert Parker.
22. >» musicus Bechst.
23. » equitwm, sp. n.
24, t, MRSD:
25. Anas sp.
9

26. Marmaronetta angustirostris Ménétr. sp.
27. Columba melitensis Lydekker.*

28. Grus melitensis Lydekker. f

29. Otis tarda Linn. sp.

30. Tetrax sp.

REPTILIA AND BATRACHTIA.

31. Testudo robusta Leith Adams.
a2. » spratti Leith Adams.

Sion » robustissima 'Tagliaferro.
34. Lutremys europea Gray.

35. Lacerta sp. .

36. Batrachia undetermined.

Besides the above, remains of several domesticated species have
been recorded by Dr. Smith Woodward from the Har Dalam
Cavern, from which rude pottery has also been obtained in soine
quantity =.

I have omitted from my list both Hlephas falconert of Busk
and Myoxus carter of Leith Adams. The specimens described
under the former name seem hardly sufficiently distinct to be
separated from H. melitensis, to which species they were re-
ferred by Lydekker. The same author also pointed out that

* Cat. Foss. Birds in Brit. Mus. 1891, pp. 13, 124.
+ Proc. Zool. Soc. 1890, pp. 404, 408.
{ See Ashby, Zammit & Despott in ‘Man,’ Jan. & Feb. 1916, vol. xvi. Nos. 1 & 2.

FOSSIL VERTEBRATES. 423

MM. cartei was evidently a synonym of Leithia melitensis; he
further showed that the mandibular ramus figured by Leith
Adams as that of a young specimen of Leithia is undoubtedly
that of an Hliomys.

The occurrence of Arvicola amphibius and A. pratensis is given
‘on the authority of Leith Adams, and I do not know if any
specimens have been preserved. It ought to be mentioned that
Dr. Caruana reported having found a portion of a lower jaw of a
Hyeena in the island of Gozo.

So far as I am aware no thorough investigation has been made
of the present-day mammalian fauna of the island, but it would
seem to be but poorly represented in species, for Sir John Murray
enumerates the indigenous mammalia as follows :—‘ The rabbit,
weasel, hedgehog, Norway rat, species of mice, and bats” *.

T should like to take this opportunity of recording my grateful
thanks to Dr. A. Smith Woodward, F.R.S., to Dr. C. W. Andrews,
F.R.S., and to Mr. W. P. Pycraft, for kind help and for giving
me every facility for studying the remains of fossil and recent
birds in the National Collection.

The Remains from the Har Dalam Cavern.

The small collection under notice was obtained from the
Har Dalam Cavern, and the adherent matrix shows that the
‘Specimens were embedded in a layer of red eave-earth. They
are rather fragmentary, but a comparatively large number of
Species are represented and range from a small Hlephas to the
extinct rodent Leithia. Most numerous of all are the avian
remains, which include those of a hitherto undescribed swan and
several other species not previously recorded as occurring in a
fossil condition an the island. It will be remembered that
Dr. Cooke had already carried out some investigations in this
cave, the chief results of which have been described by him and
Dr. Smith Woodward 7.

MAMMALIA.

Of mammals, there are examples of four species only, two of
which call for no special notice here, for Hlephas melitensis is
represented by a scaphoid onty, and Cervus elephus barbarus by a
metatarsus and a phalanx. The remaining two are Leithia
melitensis and a small species of Hqguwus, the specimens of which
each show some points of interest.

Leithia.—A small number of rather fragmentary remains
of Leithia are included in the collection. A few of these
agree in size with the larger corresponding specimens in the
British Museum Collection, but the others are so very much
larger that they almost suggest the existence of a second species,
though it behoves one to be careful with regard to size alone

* Scottish Geog. Mag. vol. vi. 1890, p. 453.

+ Proc. Roy. Soc. vol. liv. 1893, pp. 274-288.
28*

A494 MISS DOROLHEA BATE ON.

as a character, more especially when dealing with island forms,
if island form this be.

One specimen shows very distinctly the distal joining of the
tibia and fibula which, according to Weber *, is an important.
character distinguishing the Myomorpha from the Sciuro-
morpha in which these two bones are only joined proximally.
Lydekker 7, on the other hand, attached little weight to this
point, although admitting that a distal union is unknown among
living Sciuromorphs. Unfortunately, there is not yet sufficient.
material available to settle definitely the question of the syste-
matic position of the genus, though the latter author was
probably correct in suggesting that Leithia constituted a separate
family, Leithiidee. This view is strengthened by the fact that.
two further species of Zetthia, not yet described, have been
discovered by the writer in the cave-deposits of the Balearic
Islands. This greatly extends the known range of the genus,
which is, no doubt, another representative of the ‘‘ Tyrrhenian ”
fauna preserved in the Pleistocene deposits of the islands of the
western Mediterranean region.

Through the courtesy of Mr. J. Wilfrid Jackson I have been
able to examine an imperfect left mandibular ramus of a small
species of fox from the Pleistocene of Malta, belonging to the
collection of the Manchester Museum. So far as I am aware, no
fox is found in the island at the present day, and still further
interest is given to this specimen in that this occurrence of a
small carnivore for which Lezthia would appear to be a suitable
prey, suggests that the abundance and tendency towards an
increase in size in the rodent can only be explained by the theory
that it was at any rate more or less arboreal in habit. That it
was not highly specialised for a fossorial mode of life is shown
by the shape of the skull and the curvature of the incisors f.

Equus.—Finds of Equus-remains in the Pleistocene cave and
fissure deposits of Malta have been very few up to the present,
and, so far as I am aware, none has been recorded from the other
islands of the Mediterranean, though their occurrence in the
Genista Cave, Gibraltar, has been noted by Dr. Hugh Falconer
in his list of species from that locality §. Further work in this
region will probably yield other finds of a similar kind.

The present collection includes a left upper pm. 2, which is
believed to be that of a small horse, for its crown pattern shows
the small enamel-fold described as fold 5 (“ pli caballin)” by Prof.
H. F. Osborn, who considers its presence a means of distinguishing
molars of Z. caballus from those of #. asinus ||. It imdicates an
animal of about the size of a New Forest pony. The greatest.

* “Die Saugethiere,’ Jena, 1904, p. 489.

+ Proc. Zool. Soc. 1895, p. 862, footnote.

+ I am indebted to Mr. M. A. C. Hinton for information kindly given me on
this point. :

§ Pal. Mem. vol. ii. p. 555, London 1868.

|| Osborn, H. F., “The continuous origin of certain unit characters as observed
by a Palwontologist,” Harvey Lectures, Ser. 1911-12, pp. 200-1, fig. 8.

FOSSIL VERTEBRATES. 425

height of the specimen is 63 mm., the antero-posterior width
30 mm., and the thickness 21 mm. Perhaps its chief interest
les in its association with remains of one of the small elephants
and those of a large chelonian, thus showing definitely for
the first time the contemporaneity of the Hqwuws-remains
with the rest of the extinct Pleistocene fauna of the island.
For Leith Adams wrote that he had no evidence of such a
contemporaneity ; and, further, an imperfect metacarpus, pre-
viously obtained from the Har Dalam Cavern, was found in a
superficial layer, and included with remains of man, domestic
animals, and rude pottery*. This last example, now in the
British Museum Collection, consists of the proximal half only of
the metacarpus, the greatest diameter of the articular surface
being 38 mm.; it is a somewhat stouter bone than one in the
collection of the Manchester Museum. Mr. J. Wilfrid Jackson
has made some interesting notes on this last specimen which he
very kindly placed at my disposal, and which I feel I cannot do
better than quote in full :—

“The Manchester Museum possesses a short and extremely
slender adult metacarpal bone of an Equine which was found
many years ago in one of the Malta caves associated with
remains of Cervus barbarus, fox and tortoise. In length the
bone measures 160 mm., whilst the width at the middle of
the shaft is only 20°7 mm. (index 7°72). The width of the
distal end = 29 mm.

“The slenderness of the bone is very remarkable. According
to Prof. J. C. Ewart (Proc. Roy. Soc. Edin. xxx. pt. 4, 1910,
p. 291), the cannon bones in fossil and recent Asiatic wild asses
are long and slender, the length of the metacarpal being at least

eight times the width at the middle of the shaft. The index of
the Malta bone is 7°72, and therefore suggestive of ass rather
than horse, as in the latter the index is never more than 7°5.

‘“‘ However, assuming it to belong to horse, it would indicate
an animal of about 10 hands in height (according to Ewart,
op. cit. p. 297, footnote), z.e. slightly higher than a typical
Shetland pony. In a specimen of the latter with a height of
36:5 ins. the metacarpals measured 143 x 25 mm. (jide Ewart).

““T believe the British Museum possesses a metacarpal from
Auvergne (? Pleistocene) which measures 173 x 24 mm. (index
7°20), which would indicate a slender-limbed animal under
11 hands at the withers.

“ Prof. Ewart writes me that he has also a record of a
156 x 25 mm. metacarpal from Seine Inférieure, which means
‘a horse about 9:2 hands.”

AYES.

Owing to its geographical position Malta receives many visitors
on migration, which accounts for the large number of recent

* Proc. Royal Soc. vol. liv. 1893, p. 281.

426 MISS DOROTHEA BATE ON

species recorded from the island, which nowadays has only a very
small indigenous avifauna. Numerous lists of these have been
published ; the latest and most complete is one which appeared
only last year (1915), and was compiled for the Malta University
Museuin of Natural History by M. Giuseppe Despott, Curator
of that Institution. This brings the record down to December
1914, and contains about 50 species not included ia previous lists,
while the total comes to over 300.

As might be expected, avian remains from Malta are far less:
plentiful than those of associated mammals, and an exact deter-
mination is often further hampered by the fragmentary condition.
of many of the specimens, and occasionally by lack of recent
material for comparison. It seems most probable that the birds.
whose remains occur in the cavern deposits were; at least par-.
tially, resident in the island. It is not surprising to find species
represented that nowadays only occur accidentally or on migra-
tion, for the whole character of the extinct Pleistocene fauna of
the island shows that the climate, vegetation, and probably the
extent of the land surface, were very different from those obtaining
at the present day. The fact that anserine birds, including
several extinct species, are so largely represented leads one to-
suppose that they flourished when there were considerable tracts
of low-lying and marsh lands, probably before the final submer-
gence of the land (part of which is now known as the Medina
Bank) which connected Malta with Sicily and formed a northern
extension of the present Tripolitan coast-line.

The present collection includes the distal half of a humerts
believed to be that of the Brent Goose (Granta bernicla), for it
only differs from recent specimens with which it has ‘been com-
pared in being very slightly larger. Other limb-bones appear to-
be those of the Barnacle Goose (B. lewcopsis). The former species
has already been somewhat doubtfully recorded from Malta,
while the writer has obtained remains of the latter from a
Pleistocene fissure in Menorca; at the present day these geese
occur very sparingly in the Mediterranean, and probably then
only on migration.

Remains of several species of Swans have already been obtained
from the Maltese cave-deposits, including the very large extinct
form, Cygnus falconeri, described by Parker *. Of this bird he
wrote (p. 123) that it “was rather generalized in character,
being somewhat of a goose, possessing as he did longer legs and
shorter toes than the typical swans. It would appear, however,
that ...... this bird had its wings of the full relative size: the
immense ulna shows this.” Later, he suggests that ‘“ perhaps he
was altogether more terrestrial,” but I think this was meant as
opposed to swimming habits and did not refer to any loss of
power of flight. The same author (loc. cit.) also described and
figured some specimens believed to represent C. musicus, at the

* Trans. Zool. Soe. vol. vi. pp. 119-124, pl. xxx.

FOSSIL VERTEBRATES. 427

same time suggesting the possibility of there having been remains
of more than these two species of swan in the Zebbug Cave.

A few specimens in the British Museum Collection are said
by Lydekker* to ‘“‘indicate a swan of considerably smaller size
than C. musicus.” None of the remains in the present collection
agrees with these last, and only one phalanx is somewhat doubt-
fully referred to C. falconeri. A left femur wanting the inner
condyle agrees very closely in size and form with the corre-
sponding bone of C. musicus (Brit. Mus. 449 d), and there seems
little doubt that it ought to be referred to this species, which has
already been recorded from Malta both in a fossil state and as an
accidental visitor during severe winters.

The present collection from the Har Dalam Cavern includes a
few remains of an anserine bird, believed to be a small swan,
which it has been impossible to identify with the corresponding
bones of any of the species with which I have been able to com-
pare them, either from Malta or among the recent skeletons in
the osteological collection of the British Museum. With the
small amount of material available it cannot be said definitely
that these specimens all represent a single species, but it is
believed that this is so at any rate in the case of a proximal
portion of a left humerus, a right coracoid, and a right
metacarpus. Besides these, the proximal portions of two ulne
and perhaps a radius might also be included. It is suggested
that this species be known as

CYGNUS EQUITUM, sp. Nn.

Right metacarpus (text-fig. 1).—It is proposed to take this
specimen as the type. It is in a good state of preservation,
but has the distal extremity abraded and the central portion of
the third metacarpal is absent. It is peculiarly interesting on
account of its being relatively very much shorter and stouter
than the corresponding bone of any recent species of swan or
goose with which I have been able to compare it. This character
seems to indicate without much doubt that it belonged to a bird
in which the power of flight was already considerably reduced.

The following measurements, given in millimetres, will show
the comparative size of this bone in the Maltese bird, in two
recent species of swan, and in Tachyeres.

| C. equitum. | C. musicus. C. olor. | Tachyeres.
| |

Greatest length of meta- | |

CATPUS Hence sala Sea | 90 139 137 61
Greatest diameter of shaft of | | ,

second metacarpal ........ ... 8 10% | 10 6
Greatest thickness of proximal :

EVIO MENTO aoa acadce anaccy vos 11 13 Hil 8

* Cat. Foss. Birds in Brit. Mus. 1891, p. 110.

428 MISS DOROTHEA BATE ON

It will be seen from the above that the relative proportions
of this bone in C. eqguitwm and Tachyeres are not very different,
which suggests that, as in the latter, C. eguitwm might have

Text-figure 1.

A. Right metacarpus of Cygnus equitum.
B. Right metacarpus of C. musicus.

Both natural size.

FOSSIL VERTEBRATES. 429

lost its power of flight only when the bird attained its full size
and weight. In Zuchyeres the young are said to be able to fly *

In the large extinct Cnremiornis calcitrans it is not only this
bone which had been enormously reduced but likewise the other
bones of the wing, while the keel of the sternum had almost
completely disappeared ; whereas in Tachyeres, although there is
already some reduction in the size of the ulna and radius, the
sternum appears to be normal.

Text-figure 2

A. Proximal portion of left humerus of Cygnus equitum.
B. Right coracoid of C. equitum.

Both natural size.

Compared with that of C. olor, the metacarpus from Malta is
relatively a very much shorter and stouter bone; also the third
metacarpal is separated from the second for a comparatively
much shorter distance, causing the articular ends to be more
massive. ‘The first metacarpal is in keeping with the rest of
the bone, being large and stout. The proximal articular surface
is much flatter than in either C. olor or C. musicus owing to the
pre-axial border being less raised. ‘lhe comparative proportions
of this bone perhaps approach, on the whole, more nearly to
those of C. musicus, which, judging from the skeletons which

* Owen, “On Cnemiornis,’ Trans. Zool. Soc. vol. ix. 1875, p. 266.

430 "MISS DOROTHEA BATE ON FOSSIL VERTEBRATES.

I have examined, appears to be a stouter-limbed bird than
C. olor.

Humerus (text-fig. 2.).—The proximal portion of a left
humerus believed to belong to the same species as the above
metacarpus shows a similar characteristic stoutness of build, and
is unlike any specimen with which it has been compared. It is.
actually very much smaller, but in comparative proportions agrees
fairly closely with the corresponding povtion of the humerus of
C. musicus, except that the general outline is squarer and the
head and trochanter are stouter, while the subtrochanteric fossa
is more definitely defined and much deeper, and the groove
separating the head and the trochanter is more deeply excavated.

Coracoid (text-fig. 2B).—A right coracoid is also believed to.
be that of C. equitwm, being of corresponding size and showing
the same general characteristics as the two bones described above.
This specimen is in a good state of preservation, only wanting
the outer portion of its sternal border and the point of the sub-
clavicular process. In comparative proportions it is not unlike
the corresponding bone of C. mwsicus, although its ventral aspect
is rather different owing to the wider base from which the sub-
clavicular process springs and the greater thickness of the ridge
between the head and the main body of the bone. The surfaces
of contact with the sternum are wide and shallow.

Ulna.—The collection includes the proximal portions of a
right and left ulna, which I have been unable to identify with
any recent specimens to which I have had access. They appear
to agree in size and robustness with the limb-bones described
above and are provisionally ascribed to the same species. Their
dorsal aspects show no roughened surfaces for the attachment of
the flight-feathers.

Radius.—The distal portion of a radius with about two-thirds
of the shaft is more doubtfully assigned to this species, as it is
perhaps comparatively rather larger than the two ulne.

Two species of Bustards are represented in the collection by a
few fragmentary remains. Both these species occasionally occur:
as stragglers to the island at the present day, but neither has
been previously recorded in a fossil state.

The distal portion of a right tibio-tarsus and the proximal
portion of a left scapula are referred to Tetrax campestris, while
the distal extremities of two tarso-metatarsi are ascribed to.
Otis tarda, one being that of a male and the other that of a
female bird.

TEMP.-MAJOR H. M. EVANS ON THE STING-RAY. 431i

15. The Poison-Organ of the Sting-Ray (Trygon pastinaca).
By Tempy.-Major H. Murr Evans, M.D. (Lond.),
Ihidelghil(O-(U0e).”

[ Received March 3, 1916: Read April 18, 1916.]

(Text-figures ]-7.)

Part I. HistortcaAL SUMMARY.

The question of the presence of true poison-organs in fish is:
one which has exercised the minds of observers ‘for centuries.
From Aristotle down to the end of the nineteenth century the
presence or absence of a poison-gland in the Sting-Ray has
remained an unrevealed secret, although many observers have
felt convinced that something besides the laceration by its
serrated spines was necessary to cause the pain and inflammation
that resulted from injuries produced by it. Even Bottard, to:
whom I am indebted for most of the early historical literature,
denied the presence of a poison apparatus; and the ‘ Cambridge
Natural History’ merely states (p. 177) that ‘“‘among Hlasmo-
branchs the Hagle-Rays (4 é¢obatis) and Sting-Rays (Zrygon) have
barbed or serrated spines on the tail, which inflict wounds far
more severe than those caused by mere mechanical laceration ;
but, except the mucus secreted by the gland cells of the skin,
which may possess venomous properties, no special poison-forming
glands in connection with the spines are at present known.”

Dr. Antonio Porta contributed a paper on venomous fish to:
the * Anatomischer Anzeiger’ of March 1,1905. “It can be
seen,” he says (p. 235), ‘by what I have quoted above, that until
now it was not known that a poison apparatus existed in the
Trygonide and Mylhiobatide. The barb of the Trygon is almost.
sinilar to that of the Myliobatides, but it is longer and narrower.
The said sting shows the lateral margins deceitfully serrated
with the points turned from the back to the front. According
to Moreau, in a fish of a medium size, the dart is very nearly

-. one quarter of the length of the back, but there is nothing exact.

in its proportion. In nine specimens (7. violacea and [. pasti-
naca) of medium size that I examined, I found that the length
of the dart varied from 8-7 to 12°6 cm. It is renewed every
year of the life of the fish, and since sometimes the new one
sprouts before the old one falls off, we find individual fish armed
with two or, more rarely, three or four stings. If we isolate a
sting and examine it, we observe on the ventral aspect two
grooves on either side of a ridge which become shallow in width
and depth towards the base. In these two grooves the poison-
organs are situated, which penetrate to the deepest part of the
groove and there continue laterally and above into two small

* Communicated by the SECRETARY.

432 TEMP.-MAJOR H. M. EVANS ON

tubes which converge at the base, providing a passage for the
blood capillaries, which supply the connective tissue surrounding
the gland.”

‘“‘On making transverse sections of the sting and putting them
under the microscope, we see that the ventral furrows are
occupied by a glandular mass of a more or less triangular shape
with the corners rounded off. It is composed of a great many -
cells of various sizes and shapes (°2—"4 mm. x 510), often joined
to one another to form true glandular follicles, which measure
“7-15 mm. x 510; the connecting and surrounding tissues are
rich in blood-vessels and communicate with the sheath of the
sting. Towards the apex the gland gets smaller, the cells
become less numerous and smaller and are surrounded by much
connective tissue, with which they gradually merge.”

“This gland is similar to that which is observed in the genus
Scorpena and in the greater number of other poisonous fish. It
should be considered as a cutaneous gland. The dart is merely
an arm of defence. It is united to the tail by strong ligaments
and muscles, which only, however, permit a small lateral move-
ment. The emission of the poison takes place in a very simple
manner. The sting introducing itself into the wound, the sheath
is drawn back towards the base and presses on the gland which
thus emits a poisonous liquid, which flowing towards the narrow
apical groove thus inoculates the wound.”

Part IJ. OBSERVATIONS ON SERIAL SECTIONS.

I am in agreement with Dr. Porta as to the position and
general triangular outline of the gland. I have not personally
examined ie! polson-organ of Sear, pena, but the elongated com-
pressed cells pictured by Bottard in the grooves of the spine of
Scorpena are of the same type as the gland-cells of both
Trachinus draco and vipera, and according to that authority the
poison-gland of Scorpena is a less developed type of the gland
found in the Weevers. Having myself made many sections of
the glands of both the Great and Lesser Weevers, I can state
with assurance that the glandular structure found in Trygon is
of a totally different type: in fact, there are many points in its °
structure of a unique character, and the arrangement of the cells
requires careful examination. The gland consists, for the most
part, of a fine mesh, within the interstices of which are groups of
small cells with a vacuolated protoplasm. ‘These cells are grouped
together in regular follicles.

These follicles in other parts are entirely filled with secretion,
so that you have a cystic appearance, a distended cavity, lined by
a layer of flattened cells. The external margin of this glandular
mass shows a well-marked layer of pigment-cells, and external to
this are several layers of rounded epithelial cells, which, however,
are frequently detached in the sections, as shown in text- fig. 2.

If we could understand the origin and ‘relations to other parts

THE STING-RAY. 433.

of the tissues which occupy the lateral grooves, which for con-
venience and brevity I propose in future to speak of as the
glandular triangle (Porta speaks of it having a triangular shape
in section with the angles rounded off), we must study sections
at the root of the spine or dart, before it separates from the whip-
like tail. With the naked eye one can see a special dark pig-
mented patch on the tail where it opposes the spine, on the
surface of which appears some soft whitish epidermis. More-
over, if we view the spine at this point in profile, we notice that.

Text-figure 1.

xy

0"

\Y

.
SS
sos
Sx

A

.
SS
ws

\S
SS
SS

\
OS

Ss
SS

Ce)
SSS SN AG
SS
SE <>
: SRS SSS
See S S>
I4 Se

SOS

.
ESOS SES Se SS SS
0.p.S.

Trygon pastinaca.
Part of gland in groove.

c.c.g. Central canal of groove. e¢.¢. connective tissue. e.g. epithelium of groove.
fg. follicles of gland. o.p.s. osseous part of spine. p./. pigment-layer.
v.a.g. ventral aspect of groove.

the dorsum of the spine becomes free of the epidermis which
has been covering it sooner than the ventral aspect, so that the
dorsum of the spine in this respect is somewhat similar to the
nail on a man’s finger. The epidermis ends rather abruptly on
the dorsum, while, on the other hand, the dentate margin and
the lateral grooves between it and the ventral ridge separate:
gradually from the tail, the last part to become free being the
ventral ridge. On either side there is a gradual invagination
of ectoderm between the tail and the glandular triangle, this.

434 TEMP.-MAJOR H. M. EVANS ON

invagination becoming deeper as one traces it posteriorly.
Microscopically, one can observe that this invagination is carried
out by a specialised portion of epithelium. Where the spine
begins to separate, the flattened epidermal layer and mucous
-cells covering the tail are replaced by a layer of columnar cells

Text-figure 2.

Trygon pastinaca.

Portion of gland of groove, showing follicles full of secreting cells,
and central canal empty.
e.c. Central canal or duct (empty). c¢.¢. connective tissue. d.g. ducts of gland.

Fg. follicles of gland. m.f. muscle-fibres. 2.c.c. nipple of central canal.
0.p.s. Osseous portion of spine.

with superimposed layers of rounded cells, resting on a basement-
membrane in which is a layer of large deeply pigmented cells.
This peculiar layer of epithelial tissue gradually grows inwards,
i.e., towards the middle line on either side, and inserts itself
-between the tissues occupying the grooves and the tail.

THE. STING-RAY. 435

Text-figure a

Trygon pastinaca.
Lateral view of spine, with diagrammatic sections at A, B.

e. Canals. d. tooth. g. glandular tissue. 7.s.e. invagination of specialised
epithelium. m.v.7. median ventral ridge... pigment-layer.

Text-figure 4,

Trygon pastinaca.
Half-section of spine separating from tail, showing invagination of specialised
epithelium and pigment-layer.
ec. Cartilage. d.m.s. dentate margin of spine. e.¢. ordinary epithelium of tail,

m.f.t. muscle-fibres of tail. 0.p.s. osseous portion of spine. — s.e.z, special
epithelium of invagination. ¢.g. tissue of groove. #.¢. tendon of tail.

436 TEMP.-MAJOR H. M. EVANS ON

The tissue of the triangle at this stage consists of the fibrous
tissue which precedes the formation of bone ; the pigmented layer
and the round cells in this way extend over the tissue of the
groove, and the actual separation of the spine from the tail is
accomplished by a division of the round-celled epithelium, so that
when the separation is complete there is a pigment-layer covering
the glandular triangle covered with several layers of epithelium,
and a similar pigment-layer with epithelial covering facing it on
the dorsal surface of the tail.

Text-figure 5.

Trygon pastinaca.

A. Transverse section of spine near base.
B. Lateral canal and nipple. Canal full.
Cees bs s Canal emptying.

e.c. Central canal. d.m. dentate margin. fig. follicles of gland. g.¢. glandular tissue.
1.6. lacune of bone. l.c. lateral canal. m.r. median ridge. 1.l.c. nipple of
lateral canal. s. secretion.

The pigment-layer may also be observed to dip into the trian-
gular area, and carries with it these rounded cells.

If we now examine more carefully the secreting tissue of the
glandular triangle, we notice throughout the length of the
groove, but more markedly near its base, two definite rounded

THE STING-RAY. 437

cavities, occupied by a mass of homogeneous yellow material
staining yellow by Van Giesen’s method. These cavities tend
throughout the whole series of sections to arrange themselves
into two ducts or canals, one lying near the centre of the gland or
towards the ridge, and the other lying towards the lateral
margin in the direction of the teeth. The follicles seem to
empty themselves either into one canal or the other. As each
duct or canal becomes filled with secretion it bulges the pigment-
layer so as to make it protrude externally. On the peripheral
margin of the lateral canal this bulge is surmounted by a curious
nipple-shaped projection which appears throughout the whole
series of sections, at times distended, at times empty and flattened.
Some of the sections show the nipple discharging a fluid from its

Text- figure 6.

Trygon pastinaca.
A-C. Series of sections progressively diminishing towards tip.

em. Elongated nipple. e.n.2. the same enlarged (note bulbous tip containing

secretion). J.c. lateral canal. m.c. canal of median ridge (r.). _ s.f. secreting
filaments. s.f.l. the same enlarged.

tip, and one feels convinced that these sections actually show
the secretion from the lateral canal being discharged externally.
Towards the tip of the spine, where the groove is smaller, the
nipple is prolonged into a filamentous tube.

By counting the sections in series and observing the presence
or absence of a nipple-like projection from the lateral canal, one
is able to estimate approximately the size of these projections.

Proc. Zoou. Soc.—1916, No. XXIX. 29

438 TEMP.-MAJOR H. M. EVANS ON

A nipple will extend for °3—4 mm., and there is then a gap of
about °2 mm. until the next nipple appears on the lateral canal.
In this way the dentate margin is moistened with secretion, and
as the teeth lacerate the tissue of the victim the poison becomes
inoculated in the wound. Towards the tip of the spine there is
little glandular tissue: the central canal divides into two or more
channels, and the secretion is discharged by means of a canal, of
which there are two on either side, a laterai canal near the base
of the teeth, and a canal resting on the margin of the median
ridge where it looks towards the lateral groove.

Text-figure 7.

[Looe
7 (2S SS
é ee Ss 4 p
KC OR S_
= —————— — UBD Paawo > aS:

—$—$<$__—_—_—__— =
ZL

2 lb EO

SFL. Da

Zhe ==SRES

Trygon pastinaca.

Diagram of portion of spine to show scheme of canals and relations of
nipples and filaments.
e.c. Central canal. 7. canal of ridge. f.g. follicles of gland. J.c. lateral canal.
mr. median ridge. ./.c. nipples of lateral canal. s.f. secreting filaments.
é. tooth pointing forwards.

It is here that these lateral canals give off the hollow filamentous
tubes, which project towards each other and are of such length
that the two make a bridge across the lateral groove. They

THE STING-RAY. 439

terminate by an orifice which is curiously pigmented, and some
sections show the secretion issuing from the tip.

The number of these secreting filaments varies, and apparently
they are provided in order to carry the poison to the tip of the
dart, where the glandular tissue no longer is present. The
width of a filament is from 2 to 5 mm.

It should be mentioned that in one spine of the three of
which I have sections, there is a well-marked nipple projecting
from the central canal near the base (see text-fig. 2), and the
canal can be seen surrounded by a layer of muscular fibres.

A diagram of this arrangement of glandular tissue, central and
lateral Gals, nipples ancl secreting filaments, is given in text-
fig. 7; it is purely diagrammatic, and does not give Sconul any pre-
tence of accuracy the actual relative size of the nipples compared
with the teeth of the dentate margin.

In concluding this account of the microscopic anatomy of the
spine, I must mention with gratitude the help I have received
from Dy. Stuart McDonald, of the University of Durham, whose
laboratory assistant, Mr. Perey Landreth, took the micro-
photographs from which several of the illustrations have been
prepared, and who gave me much technical help.

I must also mention the kindness of Mr. C. Tate Regan, of the
British Museum (Natural History), in looking at my sections, in
giving me references to the literature on the subject, and making g
suggestions as to the form this paper should take.

IT must also thank those members of the Board of Agriculture
and Fisheries, including Mr. Borley, for the help they have given
me in obtaining specimens.

In conclusion, I would add that my investigations began in
1911, and were undertaken in ignorance of the work of Dr. Porta,
who, so far as I can ascertain, was the first to describe the gland
of the groove in the spine of 7rygon.

Summary of the evidence that the gland is really a poison-
organ :—

i. That the nature of the wounds produced are not such as
would happen after a simple laceration ;

ii. that the symptoms of acute pain and inflammation are
similar to the symptoms produced by the stings of the
other venomous fish, particularly the weever ;

il. that the staining reactions of the secretion are similar to
the staining reactions of the poison of Trachinus draco ;

iv. the observations of Dr. Lo Bianco quoted by Dr. Porta.

The observations of Dr. Lo Bianco are very interesting. He
himself saw a young man become extremely pale and fall down
almost senseless for a few minutes, from having received only a
very small puncture while he was in the act of passing a Zrygon
weighing 3 kg. from one person to another. Besides which he

also relates the following most interesting fact. In the month of
29*

440) TEMP.-MAJOR H. M. EVANS ON THE STING-RAY.

September there were in the great tank of the Aquarium of the
Zoological Station of Naples four Zrygon violacea and three
Thalassochelys caretta. One of the Trygons died, and on examin-
ing it he found that the sting was broken and entirely gone.
After a few days one of the 7alassochelys would not eat any
more, unlike the others who ate with great appetite, and re-
mained in a corner of the tank: it lived thus for four days and
died on the fifth. On examining it he found the sting of the
Trygon buried quite 6 cm. under its right fin, piercing only the
skin and muscles; in the part where the sting was buried the
tissue was of a violet colour. The wound was about 3 to 4 em.
in length and breadth, and contained a putrid liquid with a most
offensive smell.

The results of this investigation show :—
1. The origin of the gland from a special layer of epithelium
starting at the root of the spine.
. That the secreting tissue consists of regular follicles with
ducts and central and lateral canals.
i. That the secretion is discharged by means of nipples or
filaments projecting from the canals.
iv. That there is a layer of muscular tissue surrounding the
central canal.

Works consulted.

1. Borrarn’s “‘ Les Poissons venimeux,” 1889.

2. Kopert’s ‘ Giftfische und Fischgifte,” 1902.

3. Porra’s paper in the “ Anatomischer Anzeiger,” xxvi.
1905.

4. Catmertr’s ‘‘ Les Venins,” 1907.

5. Cambridge Natural History, Fishes, 1904.

There are full bibliographies in Bottard’s work and in Porta’s
paper.

MR. E. T. NEWLON ON A BLACK HARE, 44]

HXHIBITIONS AND NOTICES.
March 2ist, 1916.

Dr. 5S. F. Harmer, M.A., F.R.S., Vice-President,
in the Chair.

The Secrerary read the following report on the Additions
to the Society's Menagerie during the month of February,
1916 :—

The number of registered additions to the Society’s Menagerie
during the month of February was 36. Of these 30 were
acquired by presentation, 5 were received on deposit, and 1
by purchase.

The number of departures during the same period, by death
and removals, was 145,

Amongst the additions special attention may be directed
WO) ==

1 Preuss’s Cercopitheque (Cercopithecus preussi), from the
Cameroons, presented by Major Sir George Noble, Bart., F.Z.8.,
on February 21st.

1 Korin Gazelle (Gazella rufifrons), from the Soudan, presented
by Capt. William Dyer, on February 24th.

Mr. EK. T. Newton, F.R.S., F.Z.S., exhibited the pelt and bones
of a Black Hare, for which he was under obligation to Mr. G. F.
Brooke of Leadenhall Market, who had received it with a large
consignment of Brown Hares from Siberia; but, unfortunately,
the locality was not known. This hare is of small size and with
short rabbit-hke ears.- The head and back are black excepting
only a small white spot on the forehead ; and towards the sides
there are numerous long hairs with white tips. Lower down
upon the sides the fur becomes tawny and passes into white
underneath. All the feet, but especially the hinder ones, have
light brown hair up the upper parts.

The skull and limb bones show characters agreeing with those
of the hare; but in size the animal was intermediate between
our common hare and the rabbit.

Mr. D. M.S. Watson, F.Z.8., gave an account of some obser-
vations he had made on the habits and life-history of Platypus
and Hehidna.

449 MR. R. I. POCOCK ON THE

The Alisphenoid Canal in Civets and Hycnas.
Mr. R. I. Pocock, F.R.S., F.L.S., F.Z.S., Curator of Mammals,

gave an exhibition, illustrated by lantern-slides, to show some
points connected with the alisphenoid canal in the Viverride *
and Hyznide, and remarked :—

“Asis well known, the alisphenoid canal is always absent in
the Felide (text-fig. 1, A). In the Viverride, on the contrary,
it is nearly always present, although it is never found in the
Masearene genera Galidia, Galidictis, and Salanoia (text-fig. 1, B),
forming the subfamily Galidictine, and may be present or absent
in Cynogale (Cynogaline) and Hupleres (Kuplerine), two aberrant
genera of Viverride. By Mivart, Flower, and authors inspired
by them, it is also stated to be variable in its occurrence in
Viverri aie a genus closely related to Viverra and Genetta, in
which it is always present.

“ Hxamination of the skulls of Cynogale and Hupleres shows
conclusively that the absence of this canal, when it is absent, is
due to suppression, complete or partial, of its external bony wall.
Nevertheless, when this wall is unossified in these forms, the
channel marking the course of the external carotid artery is very
apparent. This bony wall is also so short in some Mongooses,
e. g. Crossarchus, that a comparatively slight defect in ossification
would convert the canal into an open channel, such as is seen some-
times in Cynogale and Hupleres. In Crossarchus (text-fig. 1, C, D),
Cynogale, and Hupleres, moreover, the foramen rotundum opens
alongside the anterior orifice of the alisphenoid canal into the
posterior part of the temporal fossa close to the sphenoidal fissure
(foramen lacerum anticwm). In the Galidictine the foramen
rotundwum oecupies precisely the same position with regard to
the sphenoidal fissure. This part of the skull in Galidictis, for
example, bears a close resemblance to that of Crossarchus, except
that there is no trace whatever of the alisphenoid canal. Never-
theless, the general likeness alluded to suggests that the absence
of the canal in Galidictis may be due to the complete suppression
of its outer wall (text-fig. 1, B).

‘This interpretation seems to be the one that is currently
accepted ; and since no alternative has, so far as I am aware,
been sugcested, it may be assumed that the same explanation
has been “tacitly extended to those specimens of Viverricula in
which the canal has been described as absent (text-fig.1, KE). The
statement, however, that the canal is absent in that genus is not
true. It is in reality present, its apparent absence being due to
the closure of its posterior orifice and not to the imperfection of
its outer wall.

“ Justification for this view rests upon the following facts :—

“The orifice, lying alongside the sphenoidal fase in .Viver-
ricula, which Flower would doubtless have called the foramen
rotundum, is 1m reality the anterior end of the alisphenoid canal,
which, when complete, opens posteriorly by a small aperture just

™* The term Viverride is here used, without prejudice, in the sense in which
Flower and Miyart and their successors employed it.

ALISPHENOID CANAL IN CIVETS AND: HYZNAS. 443

Text-figure 1.

_ Left cranial foramina of Felis with zygoma cut away. 1. optic foramen ;

2. sphenoidal fissure (foramen lacerum anticum) ; 3. foramen rotundum

4. foramen ovale; A.J. auditory meatus.

. The same of Galidictis.
“Phe same of Crossarchus. al.’ anterior, and al.2 posterior orifice of alisphe-

noid canal with bristle passed through it.

_ The same with outer wall of alisphenoid canal (a/s.) cut away.
_ The same of Viverricula with posterior orifice of canal closed.
- The same with outer wall of canal cut away to show foramen yotundum (8)

opening into posterior end of canal.

_ The same of Genefta, showing the complete alisphenoid canal concealing the

foramen yotundum.

The same with the outer wall of the canal cut away, exposing the foramen

rotundum opening into it.

444 MR. R. I. POCOCK ON THE

in front of the foramen ovale. When this aperture is absent, the
presence of the canal may be demonstrated by cutting away its
outer wall backwards from its anterior orifice. The true foramen
rotundwm will then be revealed perforating the cranial floor at
the posterior end of the canal a little in advance of the foramen
ovale on the admedian side (text-fig. 1, F). Hence in Viverricula
the OP amen rotundun opens into the alisphenoid canal, or, if the
term ‘canal’ be inadmissible for a tube closed at one end, into
the alisphenoid tube representing the canal.

“Tt may be added that this region of the skull in Viverricula
is alike in all specimens, apart from the presence or absence of
the posterior orifice of the canal, which may be represented by a
hole only large enough to insert a needle. Whereas if the
alleged absence of the canal were due to the suppression of its
outer wall, the canal would be represented by a groove, as in
Cynogale and Eupleres, which is not the case, and the foramen
rotundum in Viverricula would have to be described ag a long
tube, to which no parallel can be found in the Aluroidea.

“That the mterpretation above given is correct may be further
shown by comparing Viverricula with Genetta (text-fig. 1, G, H),
Cwwettictis, and other genera where the for amen ora piercing
the skull, may be seen within the alisphenoid canal by looking
through its posterior orifice, the aperture in the skull close to the
sphenoidal fissure being the anterior orifice of the canal and not
the foramen rotundum of the Felide.

* The alisphenoid canal is also stated in current literature to be
absent in the Hyzenide ; and this opinion seems to date from
Turner’s rejection in 1848 (P.Z.5. 1848, p. 81) of Cuvier’s
statement in 1837 that it is present in these animals. Cuvier’s
words are:—‘ Dans Vhyene .... le trou optique, le sphéno-
orbitaire, le rond, le vidien [alisphenoid canal] .et Vovale différent
peu du chien. Jai un individu ou il ya ue canal vidien d’un
cété et pas de l'autre’ (Anat. Comp. ed. 2, ii. p. 471). This
assertion, suggesting that the canal is generally present and
exceptionally absent, is not altogether ‘correct ; nevertheless,
Turner, Flower, and Mivart were wrong in citing the absence of
the canal as characteristic of the family Hyenide. It is usually
absent but sometimes present, at all events in Crocuta (text-fig. 2).
Tt is much shorter than in Viverricula. Nevertheless, its apparent
absence is due to the same process as in that genus, namely the
obliteration of its posterior orifice. There is sometimes no trace
of this orifice ; but quite commonly it is represented by a small
aperture a little in front of the foramen ovale. This aperture
may lead into a very short blindly ending tube, whence a small
hole, also to be seen at the posterior end of the canal an Canis,
penetrates the sphenoid bone (text-fig. 2, B). In other and rarer
cases where this aperture is larger, a bristle passed into it
emerges at a tolerably large foramen. lying beneath the hinder
end of the sphenoidal fissure in the temporal fossa. This foramen
is the anterior end of the canal and not the foramen rotundum
which perforates the base of the skull within the canal behind

ALISPHENOID CANAL IN CIVETS AND HYENAS. 445

that orifice as in Viverricula, as may be shown by cutting away
the wall of the canal.

“The arrangement above described is, so far as my observations
go, much more obvious in the Spotted Hyzena (Crocuta) than in
the Striped Hyzna (Hyena). In both genera there may be no
trace of the posterior orifice, but in Hyena this orifice, when
present, is apparently always quite small and never completes the
eanal. The foramen rotundwm, moreover, is set more forwards,
so that to all intents and purposes it opens direct into the
temporal fossa as in the Felide.”

Text-figure 2.

oy A.M.
STW yy

by ps tton -
“ae:
= We =

A. Left cranial foramina of Orocuta (Spotted Hyena) with zygoma cut away.
1. optic foramen ; 2. sphenoidal fissure (foramen lacerwm anticum) ; 4. fora-
men ovale; A.M. auditory meatus ; a7.’ anterior orifice of alisphenoid canal ;
al, partially obliterated posterior orifice of the canal.

B. The same with the outer wall of the alisphenoid canal (a/s.) cut away, exposing
the foramen rotundum (3) opening into it and showing the posterior orifice
of the canal a/.? with its minute foramen, shut off by bone trom the main
portion of the canal, When this bone is absent, the canal is complete from
end to end.

446 MR. D..SETH-SMITH ON ‘6 INTENSIVE”? POULTRY-HOUSES.

April 4th, 1916.

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

Mr. J. T. Cunntnenam, M.A., F.Z.S., exhibited a number of
skins of fowls produced in the course of six generations descended
from a cross between a male Gallus bankiva and a female’ Silky
fowl. 'The cross was made at the Society’s Gardens in 1910, and
Mr. Cunningham’s specimens were bred from a pair of the F 1’s
given to him in 1911.

The chief points ilustrated were :—

(1) The production of a recessive pile, instead of pure white
recessives; in the pile the female had reddish brown on the
breast and abdomen, the male had no colour on the abdomen,
but yellow on the back and loins. In the first mature plumage
both sexes had reddish brown on the breast.

(2) The production of two types in the coloured dominants, as
well as individual variations. One type was dark, the other
hght: in the former there was an excess of the black colour,
especially about the head, in the latter the head was yellow.
The difference was more conspicuous in the females than in the
males.

Individual differences were shown in comparing a hen with
vinous-red colour over a considerable part of the body, especially
the breast and wings, and another in which there was no vinous
colour, but a neutral drab. These facts seem to indicate that
segregation occurs between colour and white in Mendelian
fashion, but that the segregation 1s not complete, that the colour
is not a permanent unit, but undergoes subdivision.

Mr. D. Sera-Suirn, F.Z.S., Curator of Birds, exhibited lantern-
slide photographs of “intensive” poultry-houses, and remarked
that the Council had decided to hold an exhibition of laying
hens, kept on the intensive system, with a view to educating the
public to the possibility and importance of keeping poultry for
ege-production, even though their accommodation was limited
to a suburban garden or even a back-yard. The system was
explained, and stress laid upon the importance of correct feeding
and. sufhcient exercise, the latter being provided by the birds
beng compelled to scratch for their grain, which must be buried
under deep litter.

The Exhibitor stated that the houses were of three sizes, to
accommodate from six to thirty birds, and had been lent. to the
Society by Mr. Randolph Meech, who was the pioneer of the
system in this country. The exhibition would be open to
the public on April 8th, and some two hundred birds would be
on view.

THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 447

Prof. J. P. Hin, D.Se., F.R.S., F.Z.S., exhibited living speci-
mens of the Crecilian, Siphonops annulatus, collected by the Percy
Sladen Expedition at Theresopolis, Serra dos Orgaos, Brazil, in
October 1913. He also exhibited a series of photographs of
embryos of the same, obtained from eggs laid at University

College.

Mr. G. A. Boutencsr, F.R.S., F.Z.S., read a paper “ On.the
Lizards allied to Lacerta muralis, with an Account of Lacerta
agilis and L. parva.”

This paper wili be published in the ‘ Transactions.’

April 18th, 1916.
Dr. S. F. Harmer, M.A., F.R.S., Vice-President,

in the Chair.

The Secrerary read the following Report on the Additions
to the Society’s Menagerie during the mouth of Mar chiy 1Saice

The number of registered additions to the Society's Menagerie
during the month ‘of March. was 176. Of these 116. were
acquired by presentation, 5 were received on deposit, 50 by
purchase, and 5 were born in the Gardens.

The number of departures during the same period, by death
and removals, was 158.

Aimonegst the additions special attention may be directed
WO Ba

2 Drills (Papio lewcopheus), from W. Africa, purchased March
16th.

1 Bay Duiker (Cephalophus dorsalis), from 'Togoland, presented
by O. H. Bohner, on March 23rd.
_ 2 Thar (Hemitragus jemlaicus), from Chamba, presented by
the Government of the Punjaub, on March 20th.

2 Axis Deer (Axis avis), from India, and 3 Bennett's Wallabies
(Macropus bennett), from Tasmania, presented by Sir Edmund

G. Loder, Bart., V.P/Z.8., on March 21st.

The Secrerary read a letter he had received from Lt.-Col.
R. T. Leiper, D.Sc., E.Z.8., R.A.M.C., on the subject of his
recent investigations in reference to Bilharziosis, the life-history
of the parasite and prophylactic measures.

Mr. C. Tare Reean, M.A., F.Z.8., gave an exhibition of
lantern-slides iusimeiine how certain Genes protect their eges
by carrying them about, either in the mouth (drius, some
Cichlidee), on the occiput (Kurtws), on the abdomen (Aspredo),
or in a special brood-pouch (Syngnathide).

448 MR. R. H. BURNE ON TELEOSTEAN FISHES.

May 9th, 1916.

Dr. S. F. Harmer, M.A., F.R.S., Vice-President,
in the Chair,

The Srcretary read the following report on the Additions to
the Society's Menagerie during the month of April, 1916 :—

The number of registered additions to the Society’s Menagerie
during the month of April was 90. Of these 36 were acquired
by presentation, 23 were received on deposit, 30 by purchase,
and 1 was born in the Gardens. .

The number of departures during the same period, by death
and removals, was 117.

Amongst the additions special attention may be directed to :—

2 Capybaras (Hydrocherus hydrocherus), from South America,
presented by Sir Edmund G. Loder, Bart., V.P.Z.S., on April 18th.

1 Long-haired Armadillo (Huphractus vellerosus pannosus),
from Cordova, presented by Wilfred A. Smithers, C.M.Z.S., on
April 10th.

2 Australian Barn-Owls (Sérix delicatula), from Kalgoorlie,
Western Australia, presented by Dr. J. Vere Arkle, on April 3rd.

3 South-American Cecilians (Siphonops annulatus), from
Brazil, presented by Prof. J. P. Hill, F.R.S., F.Z.S., on April 4th.

Mr. R. H. Burne, M.A., F.Z.8., exhibited preparations from
the Royal College of Surgeons Museum of various Teleostean
Fishes *, showing connections of different kinds between the
swim-bladder and the ear. The fishes belonged to several
distinct families. In some (Berycide, Gadide, Hyodontide,
Notopteride) the connection was shown to be by direct contact
between a process of the swim-bladder and a fenestra in the
periotic capsule, or even (Clupeide) between the swim-bladder
and part of the internal ear; while in others (Ostariophysi) it is
indirect and the swim-bladder is connected with the perilymph
spaces that surround the ear by a chain of ossicles (Weberian
ossicles).

It was suggested that the above connections are probably an
aid in the perception of sound, and, in furtherance of this view,
specimens were shown of the ‘elastie spring” mechanism in
several Siluroids, by which the walls and contained gases of the
swim-bladder can be made to give rise to sonorous vibrations.

* Presented to the College by Col. C. E. Shepherd.

CORRECTION.

On p. 109 of Prof. Poulton’s paper on Moths from Somaliland
“Genus Pachycoa” should read: ‘ Genus Pachycoa, nov.”

NiO: 1555

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

April 4th, 1916.

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

The Minutes of the last Scientific Meeting were confirmed.

Mr. J. T. CunntnenAm, M.A., F.Z.S., exhibited a number of
skins of fowls produced in the course of six generations descended
from a cross between a male Gallus bankiva and a female Silky
fowl. The cross was made at the Society’s Gardens in 1910, and
Mr. Cunningham’s specimens were bred from a pair of the F 1’s
given to him in 1911.

The chief points illustrated were :—

(1) The production of a recessive pile, instead of pure white
recessives; in the pile the female had reddish brown on the
breast and abdomen, the male had no colour on the abdomen,
but yellow on the back and loins. In the first mature plumage
both sexes had reddish brown on the breast.

(2) The production of two types in the coloured dominants, as
well as individual variations. One type was dark, the other
light: in the former there was an excess of the black colour,
especially about the head, in the latter the head was yellow.
The difference was more conspicuous in the females than in the
males.

Individual differences were shown in comparing a hen with

* This Abstract is published by the Society at its offices, Zoological Gardens,
Regent's Park, N.W., on the Tuesday following the date of Meeting to which
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,
to all Fellows who subscribe to the Publications ; but it may be obtained on the
day of publication at the price of Sixpence, or, if desired, sent post-free for
the sum of Stix Shidlings per annum, payable in advance.

18

vinous-red colour over a considerable part of the body, especially
the breast and wings, and another in which there was no vinous
colour, but a neutral drab. These facts seem to indicate that
segregation occurs between colour and white in Mendelian
fashion, but that the segregation is not complete, that the colour
is not a permanent unit, but undergoes subdivision.

Mr. D. Sera-Suirn, F.Z.8., Curator of Birds, exhibited lantern-
slide photographs of “intensive” poultry-houses, and remarked
that the Council had decided to hold an exhibition of laying
hens, kept on the intensive system, with a view to educating the
public to the possibility and importance of keeping poultry for
ege-production, even though their accommodation was limited
to a suburban garden or even a back yard. The system was
explained, and stress laid upon the importance of correct feeding
and sufficient exercise, the latter being provided by the birds
being compelled to scratch for their grain, which must be buried
under deep litter.

The Exhibitor stated that the houses were of three sizes, to
accommodate from six to thirty birds, and had been lent to the
Society by Mr. Randolph Meech, who was the pioneer of the
system in this country. The exhibition would be open to
the public on April 8th, and some two hundred birds would be
on view.

Prof. J. P. Hitn, DSc., F.R.S., F.Z.8., exhibited living speci-
mens of the Cecilian, Siphonops annulatus, collected by the Percy
Sladen Expedition at Theresopolis, Serra dos Orgaos, Brazil, in
October 1913. He also exhibited a series of photographs of
embryos of the same, obtained from eggs laid at University
College.

Mr. G. A. Boutencer, F.R.S., F.Z.8., read a paper “On the
Lizards allied to Lacerta muralis, with an Account of Lacerta
agilis and L. parva.”

This paper is the third and last instalment of a revision of the
Wall-Lizards, of which the first two parts were published in the
‘ Transactions’ in 1905 and 1913.

The author has endeavoured to depart from the empirical
method usually followed in the arrangement of species, by tracing
back the various forms of this difficult group to a hypothetical
ancestor of which Lacerta agilis appears to be the nearest living
representative. The characters of lepidosis and coloration on
which his views are based are discussed, and detailed descriptions
are given of £, agilis and its ally Z. parva, the latter being
regarded as the connecting-link between the first and fourth of
the six sections into which it is proposed to divide the genus
Lacerta. All the species of the fourth section, of which the
type, L. muralis, has been dealt with in the previous contri-

1g

butions, are described with comments on their mutual relation-
ships. ‘Che author’s views on the evolution of markings agree
with Eimer’s well-known theory, but the original pattern is
carried back to a type more primitive than any postulated by
Kimer, in which a light vertebral streak is present. The lines
of evolution are held to be the reverse of those advocated more
recently by Prof. von Méhely.
This paper will be published in the ‘ Transactions.’

Mr. BouLencer also read a short paper containing an account
of some specimens of the Perciform Fish, Tilapia nilotica, with
increased number of anal spines.

Mr. Rosert Gurney, M.A., F.Z.8., communicated a paper on
a collection of Freshwater Entomostraca made by Mr. G. W.
Smith in Ceylon in 1907. The collection contained examples of
39 species, and one species of Copepoda and two of Ostracoda
were described as new, one of the latter belonging to the typically
African genus Oncocypris.

A paper was received from Major R. Mernerrznacen, F.Z.8.,
on the Sitatungas (Limnotragus) of the Sesse Islands. The
author found that the Bugalla Island antelopes of this genus
seem to be of the same race as the mainland form, Lamnotragus
spekei, but that the Nkose Island form, which he proposed as a
new subspecies, diflered in the shortness of its hoofs and other
characters.

The next Meeting of the Society for Scientific Business will be
held on Tuesday, April 18th, 1916, at half-past Five o’clock P.M.,
when the following communications will be made :—

EXHIBITIONS AND NOTICES.
Sips raat ue eee Ede

R. I. Pocock, F.R.S., F.L.S., F.Z.8.
On the External Characters of the Mongooses (Mungotide),

Major H. Muir Evans, M.D., R.A.M.C.
The Poison-Organ of the Sting-Ray (Zrygon pastinaca).

20
The following Paper has been received :—

Dr. J. C. Morrram.

An Experimental Determination of the Factors which cause
Patterns to appear conspicuous in Nature.

The Publication Committee desire to call the attention of
those who propose to offer Papers to the Society, to the great
increase in the cost of paper and printing. This will render it
necessary for the present that papers should be condensed, and
be limited so far as possible to the description of new results.

Communications intended for the Scientific Meetings should
be addressed to

P. CHALMERS MITCHELL,
Secretary.
ZOOLOGICAL Society oF Lonpon,
Recent’s Park, Lonpon, N.W.
April 11th, 1916.

No. 156.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*
April 18th, 1916.

Dr. S. F. Harmer, M.A., F.R.S., Vice-President,
in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

The Secretary read a Report on the Additions to the Society’s
Menagerie during the month of March 1916.

The Secrerary read a letter he had received from Lt.-Col.
R. T. Leiper, D.Se., F.Z.S., R-A.M.C., on the subject of his
recent investigations in reference to Bilharziosis, the life-history
of the parasite and prophylactic measures.

Mr. C. Tare Reean, M.A., F.Z.S., gave an exhibition of
lantern-slides illustrating how certain fishes protect their eggs
by carrying them about, either in the mouth (Arius, some
Cichlid), on the occiput (Kurtus), on the abdomen (Aspredo),
or in a special brood-pouch (Syngnathide).

Major H. M. Evans, M.D., R.A.M.C., read a paper “On the
Poison Organ of the Sting-Ray (Z'rygon pastinaca).”

It has been observed for centuries that the wounds produced
by the serrated spine growing from the base of the whip-like tail
of the Sting-Ray produced very severe injuries and pain and

* This Abstract is published by the Society at its offices, Zoological Gardens,
Regent’s Park, N.W., on the Tuesday following the date of Meeting to which
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,
to all Fellows who subscribe to the Publications ; but it may be obtained on the
day of publication at the price of Sixpence, or, if desired, sent post-free for
the sum of Six Shillings per annum, payable in adyance,

22

inflammation, which could not be accounted for by the laceration
of the wounds alone.

Dr. Antonio Porta in 1905 described a gland in the groove
lying medially to the rows of teeth on either side, which he stated
is similar to the gland found in Scorpena.

Major Evans’s researches do not confirm Porta’s description in
all particulars. The examination of a series of sections shows a
gland of a different type from that found in the Weeyvers,
Scorpena, etc. The points emphasized are :—

(i.) The origin of the gland from a special epithelial structure
at the base of the spine.

(ii.) The arrangement of follicles discharging their secretion
by ducts or canals, communicating with the exterior by means of
nipples or filaments.

(iii.) The arrangement of these nipples at the base of the
teeth.

(iv.) The presence of muscular fibres surrounding the main
canals which are instrumental in discharging the venom.

Mr. R. I. Pococr, F.R.S8., F.Z.S8., Curator of Mammals, read
a paper, illustrated by lantern-slides, “ On the External Charac-
ters of the Mongooses (Mungotide),” dealing principally with
the ears, feet, and anal sac. Reasons were given for restoring
the generic names Ariela for Crossarchus fasciatus and Atilax
for Mungos paludinosus. It was also shown that the Mongooses
differ from other Viverride in the structure of the ears, and that
the type of ear in Suricata is different from that of all other
genera of the family.

The next Meeting of the Society for Scientific Business will be
held ‘on Tuesday, May 9th, 1916, at half-past Five o’clock P.M.,
when the following communications will be made :—

EXHIBITIONS AND NOTICES.
aE EE

Miss Dorornea M. A. Bats, Hon.M.B.0.U.

On a Small Collection of Vertebrate Remains from the
Har Dalam Cavern, Malta, with Note on a new Species of
the Genus Cygnus.

Dr. J. C. Mortram.

An Experimental Determination of the Factors which cause
Patterns to appear conspicuous in Nature.

The following Papers have been received :—
Miss Ontve C. Lopes.

Some Enquiries into the Question of Baits and Poisons for
Flies; being the Report on Experimental Work carried out
during 1915 for the Zoological Society of London.

Miss Wintrrep H. SAUNDERS.
1. Report on Investigations into Stable Manure to check
the Breeding of House-Flies, made during the Year 1915,

for the Zoological Society of London.

2. Report on Trials for catching, repelling, and extermi-
nating Flies in Houses, made during the Year 1915 for the
Zoological Society of London.

3. Report on some Observations on the Life-History of
the Blow-Fly and of the House-Fly, made from August to
September 1915, for the Zoological Society of London,

The Publication Committee desire to call the attention of
those who propose to offer Papers to the Society, to the great
increase in the cost of paper and printing. This will render it
necessary for the present that papers should be condensed, aud
be limited so far as possible to the description of new results.

Communications intended for the Scientific Meetings should
be addressed to

P. CHALMERS MITCHELL,
Secretary.
ZOOLOGICAL Society oF Lonpon,

-Recent’s Park, Lonpon, N.W.
April 25th, 1916.

No. 157.

ABSTRACT OF THE PROCEEDINGS

ZOOLOGICAL SOCIETY OF LONDON.*
May 9th, 1916.

Dr. 8. F. Harmer, M.A., F.R.S., Vice-President,
in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

The SEcrEerary read a Report on the Additions to the Society’s
Menagerie during the month of April 1916.

Mr. R. H. Burne, M.A., F.Z.8., exhibited preparations from
the Royal College of Surgeons Museum of various Teleostean
Fishes ?, showing connections of different kinds between the
swim-bladder and the ear. The fishes belonged to several
distinct families. In some (Berycide, Gadide, Hyodontide,
Notopteridze) the connection was shown to be by direct contact
between a process of the swim-bladder and a fenestra in the
periotic capsule, or even (Clupeidee) between the swim-bladder
and part of the internal ear; while in others (Ostariophysi) it is
indirect and the swim-bladder is connected with the perilymph
spaces that surround the ear by a chain of ossicles (Weberian
ossicles).

It was suggested that the above connections are probably an
aid in the perception of sound, and, in furtherance of this view,
specimens were shown of the “elastic spring” mechanism in
several Siluroids, by which the walls and contained gases of the
swim-bladder can be made to give rise to sonorous vibrations.

* This Abstract is published by the Society at its offices, Zovlogical Gardens,
Regent's Park, N.W., on the Tuesday following the date of Meeting to which
it refers. It will be issued, along with the ‘Proceedings,’ free of extra charge,
to all Fellows who subseribe to the Publications; but it may be obtained on the
day of publication at the price of Stxpence, or, if desired, sent post-free for

- the sum of Six Shillings per annum, payable in advance.

t Presented to the College by Col. C. H. Shepherd.

26

Miss Dorornea M. A. Bars contributed a paper dealing with
a collection of vertebrate remains from the Har Dalam Cavern,
Malta. Birds are most numerously represented therein, and
include some bones of an Anserine bird showing a reduction in
its powers of flight. It is believed to be a hitherto-undescribed
species, and is referred to the genus Cygnus. A list is given of
all the species of vertebrates recorded from the Pleistocene cave
and fissure deposits of the island.

Dr. J. C. Morrram read a paper entitled “ An Experimental
Determination of the Factors which cause Patterns to appear
conspicuous in Nature.”

A series of experiments was carried out with artificial patterns
and backgrounds under controlled conditions of lighting, and a
large number of determining factors were discovered, both as
regards plain and patterned objects and backgrounds. Finally,
the experiments showed that the most conspicuous shape and
pattern which an object can have, when viewed against a series
of plain and patterned backgrounds, was presented by a circular
dise of black, with a central circular area of white. Having
arrived at this conclusion, the Indian diurnal Lepidoptera were
completely examined, in order to discover whether any species
presented patterns approaching this ideal conspicuous pattern.
It was found that a considerable number presented patterns
hardly removed from this ideal, and that a large proportion of
these insects are considered to be “ protécted ” species presenting
“‘ warning coloration.” :

The next Meeting of the Society for Scientific Business will be
held on Tuesday, May 23rd, 1916, at half-past Five o'clock p.m.,
when the following communications will be made :—

K. G. Boutenesr, F.Z.8.
Exhibition of living specimens of the African Lungfish
(Protopterus annectens) and of their Cocoons.
Lieut. R. Broom, M.D.,'C.M.Z.S., R.A.M.C.
On the Structure of the Skull in Chrysochloris.

Pal

Dr. C. W. AnpreEws, F.R.S., F.Z.S.
Note on the Sternum of a Bird from the Eocene of
Nigeria.
Dr. A. Smira Woopwarp, F.R.S., V.P.Z.S8.

On a Mammalian Mandible from the Cretaceous of Alberta,
Canada.

V. LutsHnix.

1. List of Carabide (Coleoptera) collected in . Chopersk
District, South Russia.

2. A new Species of the Genus Platysma (Coleoptera) from
China.

3. Notes on Species of the Genus Platysma from Australia.

The Publication Committee desire to call the attention of
those who propose to offer Papers to the Society, to the great
increase in the cost of paper and printing. This will render it
necessary for the present that papers should be condensed, and
be limited so far as possible to the description of new results.

Communications intended for the Scientific Meetings should
be addressed to

P. CHALMERS MITCHELL,

Secretary.
ZOOLOGICAL Society oF LoNpDon,

ReEcEnt’s Park, Lonpon, N.W.
May 16th, 1916.

No. 158.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.”

May 23rd, 1916.

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

The Minutes of the last Scientific Meeting were confirmed.

Mr. C. Tate Reaan, M.A., F.Z.S., exhibited a specimen of the
rare fish, Centrolophus britannicus Giiuth., the fourth known
example of this species.

Mr. Regan also exhibited a Silver Ling (Molva elongata), nearly
600 mm. long, taken from the stomach of a very large Sun-fish
(Mola mola) that had been caught in a trawl, landed at Milford,
and sent to Mr. W. Howlett of Billingsgate Market, who pre-
sented it to the Natural History Museum. The Sun-fish appears
generally to swim near the surface and to eat small invertebrates,
larval fishes, ete. It is interesting to note that it may descend
to considerable depths (Jf. elongata is usually found at 100 to
300 fathoms) and that it may capture fairly large and active fish.

Mr. E. G. Bovutencer, F.Z.S., Curator of Reptiles, exhibited
living specimens of the African Lungfish (Protupterus annectens).

The Rev. H. N. Hutcuinson, M.A., F.Z.S., exhibited the
plaster cast of a model, four feet long, which he had constructed,
of the Dinosaur, Diplodocus carnegiet.

Lieut. R. Broom, M.D., C.M.Z.8., R.A.M.C., read a paper on
the structure of the skull in Chrysochloris.
Two stages in the development of the skull have been studied.

* This Abstract is published by the Society at its offices, Zoological Gardens,
Regent's Park, N.W., on the Tuesday following the date of Meeting to which
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,
to all Fellows who subscribe to the Publications; but it may be obtained on the
day of publication at the price of Siawpence, or, if desired, sent post-free four
the sum of Six Shillings per annum, payable in advance.

30

The earlier is that of a newly born Chrysochloris hottentota, whose
skull has been cut into microscopic sections and reconstructed,
and a somewhat later stage of Chrysochloris asiatica, whose skull
has been prepared for the study of the membrane-bones. The
following are the most interesting features discovered :-—
' External to the exoccipitals on each side is a large membrane-
bone which partly covers the petrosal or periotic. This is believed
to be the homologue of the bone which occurs in Therapsid and
most primitive reptiles, and usually referred to as the tabular.
The sections prove that it is no part of the auditory capsule.

Along the inner side of the prearticular or “ goniale ”—the
little membrane-bone which supports the underside of the upper
end of Meckel’s cartilage—is a second membrane-bone, which, it
is believed, has not been previously recognised in the mammal
skull. This may be the homologue of the reptilian surangular.

Under the back part of the nasal capsule, and situated between
the capsule above and the alisphenoid and pterygoid below, is a
large membrane-bone of doubtful significance. It is probably
the homologue of the “ postero-lateral vomer” of Parker.

The skull is held to be in some respects highly specialised and
in others degenerate, although also retaining a number of very
primitive characters.

Dr. C. W. Anprews, F.B.S., F.Z.S8., described an incomplete
sternum of a gigantic carinate bird from the (?) Hocene of
Nigeria. Comparison with the sterna of several groups of birds
leads to the conclusion that this specimen, though differing
considerably from the sternum of any living member of the
group, belonged to a very large representative of the Tubinares.
It has about twice the linear dimensions of the sternum of
an Albatross, of which the spread of wing (in the flesh) was
10 ft. 8in. Itis proposed to refer this species to a new genus
Gigantornis, the specific name being G. eaglesomei after its
discoverer.

Dr. A. SmirH Woopwarb, F.R.S., V.P.Z.S., read a paper on a
mammalian mandibular ramus from an Upper Cretaceous for-
mation in Alberta, Canada. The specimen represented an
opossum-like marsupial, and he referred it to a new species of
Cimolestes named C. cuélert in honour of its discoverer, Mr.
William E. Cutler. The close dental series behind the canine
measured 30.mm. in length, and the molars differed from those
of the two known species of the genus in their relatively less
elevated trigonid. The fourth premolar was a large, tumid,
laterally compressed cone, with one well-separated posterior cusp.

Mr. V. LursHnrK communicated the following three short
Coleoptera papers :—(1) A List of Carabidee collected in Chopersk
District, South Russia, (2) On a new Species of the Genus Platysma
from China, and (3) Notes on Species of Platysma from Australia.

31

Mr. E. G. Bounencer, F.Z.8., Curator of Reptiles, described
a new Lizard of the genus Phrynosoma, recently received

among a small collection of reptiles presented to the Society by
Dr. H. G. F. Spurrell, F.Z.8.

Dr. R. W. Suuretpt, C.M.Z.8., communicated some notes on
cases of albinism seen in American animals.

The next Meeting of the Society for Scientific Business, closing
the Session 1915-1916, will be held on Tuesday, June 6th, 1916,
at half-past Five o'clock P.m., when a Discussicn will take place
on the Results published in the ‘ Biologia-Centrali-Americana,’
with special reference to the zoo-geographical relations between
America and Africa.

The Discussion will be opened by Dr. F. DuCane Godman,
F.R.S., F.Z.8., and, amongst others, the following will take
part :—Dr. A. Smith Woodward, F.R.S., Dr. H. Gadow, F.R.8.,
Mr, C. Tate Regan, Mr. R. I. Pocock, F.R.S., and Dr. C. W.
Andrews, F.R.S,

The following Papers have been received :—

James F, Gemuint, M.A., M.D, D.Sc, F.ZS.

Notes on the Development of the Starfishes Asterias glacialis
O. F. M., Cribrella oculata (Linck) Forbes, Solaster endeca
(Retzius) Forbes, Stichaster roseus (O. F. M.) Sars.

S. Maurin, B.A. (Cantab.), FHS.

OnCryptostome Beetles in the Cambridge University Museum
of Zoology.

Sir Joun A.S . Bucky ILL, I M. A., E 5 , FF. ZL. 8.

Notes on the Lepidoptera of Cypr us.

32

The Publication Committee desire to call the attention of
those who propose to offer Papers to the Society, to the great
increase in the cost of paper and printing. This will render it
necessary for the present that papers should be condensed, and
be limited so far as possible to the description of new results.

Communications intended for the Scientific Meetings should
be addressed to

P. CHALMERS MITCHELL,
Secretary.
ZooLoGicaL Socrery or Lonpoy,

Recent’s Park, Lonpon, N.W.
May 30th, 1916.

No. 159.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*
June 6th, 1916.

Prof. E, W. MacBrinz, D.Sc., F.R.S., Vice-President,
in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

An informal discussion on the results published in the
‘ Biologia-Centrali-Americana, with special reference to the zoo-
geographical relations between America and Africa, was opened
by Dr. F. DuCanz Gopman, F.RB.S8., followed by Dr. H. Gapow,
F.R.S, Dr. A. Surira Woopwarp, F.R.S., Mr. C. Tare Reean,
Mr. R. I. Pococn, F.R.S., Dr. O. W. Anprews, F.R.S., Lord
Roruscuitp, D.Sc., E.RS., Prof. J, PS Hin, Se, ERS,
Mr. W. L. Scrarer, Dr. R. BROOM, and the CHAIRMAN.

This Meeting closes the Session 1915-1916, The next Meeting
of the Society for Scientific Business will be held on Tuesday,
October 24th, 1916, at half-past Five o’clock P.M.

* This Abstract is published by the Society at its offices, Zoological Gardens,
Regent’s Park, N.W., on the Tuesday following the date of Meeting to which
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,
to all Fellows who subscribe to the Publications ; but it may be obtained on
the day of publication at the price of Stxpence, or, if desired, sent post-free
for the sum of Six Shillings per annum, payable in advange,

34

The following Papers have been received :—

JAMES F’. Gemuiut, M.A., M.D., D.Sc., F.Z.8.

Notes on the Development of the Starfishes A sterias glacialis
O. F. M., Oribrella oculata (Linck) Forbes, Solaster endeca
(Retzius) Forbes, Stichaster roseus (O. F. M.) Sars.

S. Mavuir, B.A. (Cantab.), F.E.S.

On Cryptostome Beetles in the Cambridge University Museum
of Zoology.

H. G. Newt, A.R.C.S., F.Z.S.

The Karly Development of Cuewmarta: Preliminary Ac-
count.

R. BE. Turner, F.Z.8., F.E.S.

Notes on the Wasps of the Genus Pison and some allied
Genera. :

F. E. Bepparp, M.A., D.Sc., FLR.S., F.Z.8.

On Two new Species of Cestodes belonging respectively to
the Genera Linstowia and Cotugiia.

The Publication Committee desire to call the attention of
those who propose to offer Papers to the Society, to the great
increase in the cost of paper and printing. ‘This will render it
necessary for the present that papers should be condensed, and
be limited so far as possible to the description of new results.

Communications intended for the Scientific Meetings should
be addressed to

P. CHALMERS MITCHELL,
Secretary.

ZOOLOGICAL Society oF Lonpon,
Recent’s Park, Lonpon, N.W.
June 13th, 1916.

PAPERS.

Page
8. Observations on the Cytology of Flagellates and Amcebe obtained from old Stored
Soil. By T. Goopnny, D.Sc., Protozoologist, Research Laboratory in Agricultural
Zoology, University of Birmingham. (Plates I.-IV., and Text-figure 1.).......... 309

9. On some Fresh-water Entomostraca from Ceylon. By Rozserr Gurney, M.A., F.Z.S. °

UBinites el he andeTextatienrant.\iiss saber lens cers ses ae AGRO COOGAN CONG DOL » 000
10. On Specimens of the Perciform Fish Tilapia nilotica with inereased number of anal
APICES a YACHT AG bOULENGHR Hui. He Zc Sica cy se weie siecle ania sales mol ataheer therefore. GA

11. On the External Characters of the Mongooses(Mungotidz). By R.I. Pococs, F.RB.S.,
E.L.S., F.Z.8., Curator of Mammals. (Text-figures 1-10.).........002....20+--. 849

12. Notes on the Sitatunga or Marsh Antelope of the Sesse Islands, Lake Victoria Nyanza.
By Major R. Meinerrauacen, F.Z.8. (Text-figures 1 & 2.) ....2..0....00ss0022 O10

13. An Ixperimental Determination of the Factors which cause Patterns to appear
Conspicuous in Nature. By J.C. Morrram, M.B.(Lond.), (Text-figures 1-20,) .. 883

14. On a small Collection of Vertebrate Remains from the Har Dalam Cavern, Malta;
with Note on a new species of the genus Cygnus. By Dorornna M. A. Barz,

Hon.M.B.0.U. (Text-figures 1 & 2.)..... Rexel ere slo ears Pre UII ON afar iosern genes wee AQE

15. The Poison-Organ of the Sting-Ray (Trygon pastinaca). By Tempy.-Major H. Murr

Eyans, M.D.(Lond.), R.A.M.C.(0.). (Text-figures 17) ae ies 451
Vitlepage ......-.. Sees oR I eo aH Sate aonb Bee SEO OROriGwanOn C poveis i
Inston Council andsOmcersis a «cas «ects eit os ae oor SSO ERE ODL Cote e etdpecsyetees il
Sto ta@ OMperts ere ch raster nyaoherctete hale ints ROL SE a AD OOo Gh Spe Eee OOO FO Mee meas iii
Alphabetical List of Contributors ........ FORA Ar ac bike BED G0 Ucn EN AOC Minis mana lh!

Index Pere ee ee tee eer sneseentese vt eese ee eese se xii

St OE ee TAT aS,

1916, Part IT. (pp. 809-448).

Page
Goopnys yPls.1.—1 Vi, ‘SoilkProtozoa eu can eee sete ee eee ee 309
Gurnzy; Pls. I1.-III. Entomostraca from Ceylon..............+ Binoy. ee)

NOTICE.

The ‘ Proceedings’ for the year are issued in four parts, paged consecutively,
so that the complete reference is now P. Z. 8.1916, p.... The Distribution

is usually as follows :—
Part I. issued in March.

aid] & lence June.
Arp ULE - September.
5p aL Vise December.

‘ Proceedings,’ 1916, Part I. (pp. 1-807), were published on
April 20th, 1916.

The Abstracts of the ‘Proceedings,’ Nos, 155-159, are
contained in this Part,

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