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MEMOIRS

INDIAN MUSEUM

Vol. VII, 1918-1922.

EDITED BY
THE DIRECTOR
OF THE

ZOOLOGICAL SURVEY OF INDIA.

QUA

Caleutta :
PUBLISHED BY THE DIRECTOR, ZOOLOGICAL SURVEY OF INDIA.

1918-1922.

ANSE

. 1.—A contribution towards the Revision of the Passalidae of the World. F.H.

CONTENTS.

GRAVELY

. 2.—Observations on the Shells of the Family Doliidae.

. 3.—On a collection of Oligochaeta from the lesser known parts of India and from

(Published 19th December, 1918).

(Published 28th July, 1919).

Eastern Persia. J. STEPHENSON

. 4.—Report on the Parasitic Nematodes in the collection of the Zoological Survey

of India.

(Published 27th April, 1920).

H. A. Bayrıs and R. DAUBNEY
(Published 22nd December, 1922).

E. W. VREDENBURG

Page

191

263

LIST OF PLATES.

Plate I (Passalidae) =
Plates II—VIII (Mollusca)
Plates IX—XI (Oligochaeta)

Follow page.
144

7 a Al

AA
AU

ERRATA.

[ee

P. 3, line 24 from top of page, for homaloguous read homologous.

P. 11, line 6 from top of page, for Mitorhinus read, Mitrorhinus.

P. 13, line 12 from top of page, for Pseudacathinae read Pseudacanthinae.
P. 56, line 15 from top of page, for eucadorensis read ecuadorensis.

P. 66, line 15 from bottom of page, for euacadorensis read ecuadorensis.

P. 70, in the description of text-fig. VIII, 3, for punctipectis read punctipectus, and text-
fig. VIII, 5, for Eshscholtz read Eschscholtz.

P. 71, line 6 from top of page, for Malagassalus read Malagasalus.
P. 86, line 10 from top of page, for Epishenus read Ephisphenus.

P. 88, in the description of text-figs. XI, 2 and XI, 3, for javanus read javensis, and text-
fig. XI, 4, for lamellidens read lamellatus.

P. 89, line 19 from bottom of page, for Acera us read Aceraïus.

P. 92, line 21 from bottom of page and page 93, line 6 from bottom of page, for occulidens
read. oculidens.

P. 93, line 3 from bottom of page, for Trapezockilus read Trapezochilus.

P. 108, line 15 from top of page and page 109, line 8 from top of page, for Brittain read
Britain.

P. 115, line 13 from top of page, for Bonatao read Banahao.

P. 117, line 5 from top of page, for Vissale read Viscaya, line 6, for Banalao read Banahao;
and line 15 for glabriventis read glabriventris.

INDEX.

[.V.B.—An asterisk (*) preceding a line denotes a new variety or subspecies; a dagger (f) indicates a new species; a double
dagger (+) a new genus or subgenus; a double asterisk (**) a new family or subfamily ; synonyms are printed in italics.]

Aceravinae

Aceraius
aequidens
alutaceosternus
assamensis
borneanus
comptonv
germari
grandis
helferi
himalayensis
hirsutus
illegalis
kuwerts
laevicollis
laevimargo
T lamellatus
laniger
manor
moschleri
oculidens
peltostictus
perakensis
pilifer
pumilio
rectidens ..
sodalis
tavoyunus
tricornis .
virginalis
wallacei

Acuaria \
anthuris ..
calcarata

19, MON 1, SO, WN, Wes

Page

76, 121

76, 88
90, 93

90, 92

Si 109
76, 92, 93
89, 93

91, 93

Be 91
91, 93, 123
a6 91
88, 89, 92
91, 93

91, 93

16, 92, 93
102

310293
90, 93

108

102

91, 93

35 87
al ayy
321

321

Page

Acuaria contorta 321
longeornata 321
phoenicopteri 321
spinifera .. 321
squamata 321
uncinata 321
Acuaria (Acuaria) anthuris . 319
Acuaria (Echinuria) jugadornata 39 321
leptoptili .. 319, 320, 321
Acuarlinae 319,321
Aelurus i 270
fulgens .. 270, 336
Allantonema 340
Alococerus 51
Amplicaecum 56 287
+ varani + „. 281, 288
Analaches Ur, Br ll, M2
australiensis 98
bicavis 8
brachymetopus 8
dubius 8
infestus Cc
laevigatus 8
laticauda 8
paraplesius 8
punctithorax ds. 8
Ancylostoma 265, 33D, 357
caninum 50 BR), BOLO
ceylanicum 339, 336, 337
duodenale Bain, Soil
malayanum 336
Ancylostomidae 335
Ancylostominae .. 835, 337
Angiostomoidea . 303
Anguillulidae 343

Anisakinae
Anisakis
Anthropoides virgo
Antilope cervicapra
Aponeleides
Aprocta
Aproctonema
entomophagum
Arboricola torqueola
Ardea cinerea
manillensis
Arrox
agassizl ..
Ascaridae
Ascaridia
columbae
compar
cristata
perspicillum
stroma
Ascarinae
Ascaris
ardeae
ardearum
circularis
helicina
hexametra
curva
lumbricoides
reticulata ..
sulla
suum
triquetra .
vitulorum
vulpis
Ascaroidea
Askarinae
Aspidodera
Atractidae
Atractis
cruciata
dactylura
opeatura .
Atractonema
Aulacocyclinae ..
Aulacocylus

Index.

.. 340, 341

bo
Me)
et

D D bw ND bv
© ©
BR bo

ND by
CO =I
D ©

bo
er)
aq
ISS) [8S 18S) TS) TSE) 8) LS)
(en) fer)
O0 fer)

bo
fer)
D
SS OO
en &
a ©

° 340

2,5, 9, 12, 13, 120, 125, 126

5, 13, 14, 17, 120, 123

Aulacocylus aruensis
bicuspis ..*
cavicornis. .
celebensis
deyrollei ..
edentulus
errans
felderi
glabriusculus
mastersl ..
parryl
perlatus ..
platypus ..
pygmaeus
rouxl
teres
tricuspis ..

Aurelius
dohrni ..

Auritulus

Balearica pavonina
regulorum
Barbus tor
Basilianus
andamanensis
indicus
sikkimensis
Belascaris
marginata
masculior
melis
mystax
vulpis
Bradynema
Branchiodrilus
Branchiura
sowerbyi ..
Bucein
cannele
perdrix
Buccinum
dolium
perdrix

Page

18, 20

123

19, 20

if, Is, 19, 20
1819220
18, 20

17, 20

18, 20

se Lit, AD
17, 18, 20
16

ve 8
18, 19, 20
13, 14, 18, 19, 20
77, 103
104

13, 21

293

305, 306, 342
80, 86

. 194, 200

A 164

150, 162, 166
150, 162, 163, 167
16,

Buccinum tessellatum

Bungarus candidus
fasciatus ..
Bunostomum

Caccabis chucar
saxatilis ..
Calidas
Camallanidae
fCamallanides
fprashadi
Camallanus
americanus
bungari
confusus ..
dumerilii ..
Tkachugae

microcephalus

roseus
seurati
tripinosus
undulatus
aureus
pallipes
Capillaria columbae
Capra falconeri ..
Cassidaria
Cassididae
Cassius
Caulifer
Cephalobus
butschli .
Tseistanensis
Ceracupes
arrowl
austeni
fronticornis
Cervus axis
Cetejus
acutangulus
australiensis
infans
schenklingi

sodalicus .

8.11,.91..102,103, 122

Index.

Page

156, 163, 164, 166, 167

328

273, 314, 325, 330, 331

337

293

322

54 325
325, 326, 327

322, 323, 325, 326, 327, 328

.. 323, 324
328

325

ie 325
322, 323, 324, 325
323, 324, 325
325

325

336

. 269, 336
330

ily 337
.. 146, 148
146, 148, 149
51

1B, 117

341

ae 341
.. 341, 342

. 14,21, 120, 125, 126

21
21
21
337

8
77, 101
8
8
8

Chaetogaster
bengalensis
limnaei
punjabensis
spongillae

Chamaeleon calcaratus
vulgaris

Cheilospirura

Chelonia

Chevreuxia

Chilomazus borealis

Chitra indica

Chlamydonema felineum .

Chondrocephalus
fcordiger
granulifrons
purulensis
fquinquecornutus

Ciceronius
antanarıvae

Ciconia nigra

Circus cineraceus

Clemmys leprosa

Cobboldina

Cochlea rugosa
striata s. olearia

Caelopeltis monspessulana

Colinus virginianus

Coluber
helena

Comacupes
basalis
cavicornis
cylindraceus
felderi
foveicollis
mason
minor
stoliczkae

Coniger

Contracaecum
Teugonium
incurvum
microcephalum
rosarium .
Tschizothoraeis

2, 10, 11, 15, 43, 44

8 10, MENU, A

.. 45,46
11, 43, 44, 45.
70, 71

321

303-

af 331
14, 120, 123
15, 16
15,16

14, 16

ah 18
15, 120, 123.
192016

123

15, 16

a 22H
.. 281, 283
.. 284, 285
282, 283, 284
282

At 282:
. 285, 286

AV Index.

Page Page
"Contracaecum spiculigerum a: 281 Dioctophymoidea > Fe 331
tricuspe .. Sn Au 284 Diplotrema Te 6 oe 226
Corvus corone .. oe be 315 Diplotriaena in At SUS, BIG, SLU
Cosmocephalus .. Ae oe 321 trieuspis .. ws oo Bl, uly
Coturnix communis ue 28 293 Diplotriaeninae .. + A 317
dactylisonans : aA sf 293 Dispharynx ats 27 = 321
Crocodilus acutus Ae a 289 Doliidae +. 0 a, IAG, Wir, Was
americanus ae ot 289 Doliopsis BR ue 147, 179, 180
niloticus .. oe Ae 289 Dolium 145-151, 154, 155, 161, 162, 166, 167,
porosus .. Les ne 288 MAL, 174, 175, 176, 178, 180, 184, 187
‘Crocopus phoenicopterus .. = 292 ampullaceum 20 .. 156, 168
Crossocephalus .. vs a 303 antiquum 23 -. 181, 183
Cruzia uy ag Ail 309 arabicum an ag re, US)
Cruzidae 309 bairdii elie ceo 90 147
Cucullanus vwiparus = 5 325 EASE ENON DL .. 181
Cylindrocaulus .. .. 14,21, 120, 125, 126 chinense 2. 173, 16 he felon ling 182185
bucerus Le ER IB Mit cinguliferum ee .. 1498 180
patalis .. DA a 9] costatum .. 149, 150, 156, 158, 163, 167,
‘Cynaelurus jubatus aie a 270 us, 169, 12, 18
Cyon dukhunensis es “= 336 a BAER u oy, 1g
Cyphoproculus 49 costatum martinr Be .. 180, 182
Roue té ne crosseanum 146—149, 171, 180, 184, 185
4 cumingii .. ue iy 179
denticulatum AR Se 181
D deshayesi I: vs 179
Deletrocephalinae 56 931332 dolium .. ae a 169
Diaphanocephalus en 331, 332, 334 dunkeri .. os Ey 182
+minutus 331, 332, 333, 334, 335 Jasciatum 145, 146, 148, 149, 156, 160,
willeyi .. a 73311332 161, 171, 180
Dichogaster SE Kr 937. 240. 257 Jimbriatum 149, 150, 156, 162, 163, 165,
“to 167, 168, 169
SANS NE de 2% 258 :
el os galea bo 1165, WG, Ge, lal, Az, Ay;
N ee a N a 4 176, 178, 182, 184, 185
“bolaui malabaricus .. ae 257
hela palmicola a galea luteostomum .. Se LB, 1176)
: ie, u hochstetteri I 55 178, EO
crawl 7 ee N 258 - .
Meath vibe japonicum 58 174, 176, 178
Didimoides = 5c) OOS .
ee : lischkeanum ” 156, 160, 168
Didimus i =: N 72 :
: losariense se 5 Jee), 1162
duplicatus Je de 74.
nn - luteostoma las, 174, 175% Koss
ee a fs # i maculatum 149, 150, 161-163, 165-173,
idymus congoensis SL ee 8 £79) 182,2 18552186
a 8 magnificum 173, te) Ue, 110
curvilineatus a 8 martini .. oh ah 172
laevisternus 4e af 8 melanostoma 171, 174, 175, 176, 178, 182, -
latipunctus 8 185
ruwenzoricus ose es 8 minjac .. 156, 162, 167, 168, 169
Dioctophymidae , is 331 modjokasriense la, 170, 192, 126

Index.

Page
Dolium muticum ; .. 180, 181
olearium . 165, ICE MT Thee, lalaı
orbiculatum 181
ormarense a 172, 173, 180
perdix . 148, 149, 179, 182, 184, 185, 186
pomum 181
procellarum 182
pro-orbiculatum 181
ringens 181
stephaniophorum 181
subfasciatum 181
tenue 168
tessellatum 146, 148, as 150, 155, 156,
161-163, 165, 167-173, 179, 180, 184
testardi 179
townsendi 173, 180, 183
varicosum . 145, 186
varvegatum 156, 162, IAL, 173- 179, 182,
184, 185
verrillii 181185
zonatum .. 146, 147, 148, 171, 180
Dolium (Eudolium) arabicum 183
| einguliferum — 183
crosseanum es a 183
fasciatum a. 145,183, 185, 186
hochstetteri 183
muticum .. 183
ormarense ae a 182
stephaniophorum 183
subfasciatum A EN 183
tessellatum 149, 156, 182, 185, 186
verrillii a 183
zonatum . 183, 185, 187
Dolium (Malea) camurum 183
orbiculatum a: 183
pomum so MGB}, IL, CS, 187
pro-orbiculatum _ 183
ringens 183, 185, 187
Drawida 194, 200, 202
barwelli 202
*barwelli impertusus 200
bournei 202
pellucidus 902
Dujardinia 288
dujardini 288
helicina 288

Echinocephalus spinossisimus

Echinuria
jugadornata
Emys orbicularis
Epeus
Ephydatia nan
Epilaches
puberilis .
Epipertinax
Epiphanus
Epiphoroneus
Epipleurothrix
Episphenoïdes
pectiniger
quaestionis
Episphenus
comptoni
Hache
indicus
moorei
neelgherriensis
pearson ..
Eriocnemis
burmeisteri
dorsalis
gigas
ptox
quadricornis
Erionomus
alterego
palini
planiceps .
platypleura
studti

ftrichostigmoides

Ervopterus
Ervosternus
alterego
Erythraeodrilus ..
Eudichogaster
ashworthi
bengalensis
ffalcifer ..
indica
Tkinneari

9, 100

78, 85, 121, 122

85, 86

ace 106

. 103, 105
96, 97

4, 10, 69, 74
75, 76

74, 75

70, 75, 76

os 69
10, 70, 75, 76
22, 31

ae 75
.. 226, 227
194, 236, 246
. 246, 255
248

252

255

Index.

vi
Page
Fudichogaster parvus 254
fprashadi A 250, 255, 257
fpusillus .. 253
ftrichochaetus 249
Eudolium POS IGS OT, Melis 278
181, 184
Eumelosomus 12
affinis 8
aloysii sabaudiae 8
Eumelus 51
nasutus 7

Eustrongylides ignotus

. 226, 240

Eutyphoeus
bengalensis Me 2 DEB. Dalat
chittagongianus or .. 241, 245
incommodus 240
kempi 242
mohammedi 241
fturaensis .. EN 244
waltoni .. ee 243, 244, 245
F
Falcaustra a ait 303, 309, 310
Towel Gé .. 305, 306, 307, 310
faleatas: AR .. 304, 310
lambdiensis so og) QUE HOD, BIO)
fleptocephala AA .. 306, 310
slamensis ae .. 304, 310
Tstewarti .. 307, 308, 309, 310
Ttestudinis ae .. 304, 310
Falcaustra (Oxysoma) kachugae 304, 309, 310
Falco tinnunculus 315
Felis SE a 270
bengalensis 60 ACD; AO, GRA, SAO
chaus . 269, 322
concolor .. 322
leo . as se co AOU, BRO
pardus .. u. 209; 279; 322, 336
tigris ate .. 269, 270, 335, 336
uncia 3e ae 270
viverrina 269, 270, 322, 330, 335, 336
Ficula > ws ati 21874188
gracilis i 189
Filaria os ot so Ble, QUE
abbreviata aa .. 314, 315

Filaria attenuata
haje
macrophallos
fvarani

Filariidae

Filariinae

Filarioidea

Flaminius

Flavius

Florencioia

Francolinus gularis

Galeopithecus
Galloperdix spadicea
Gallus domesticus
gallinaceus
Galoncus permiciosus
Garrulus glandarius
Gastropoda
Gavialis gangeticus
Geoemydà trijuga
Glossoscolecidae
Glossoscolecinae
Glyphidrilus
papillatus
Gnaphalocneminae
Gnaphalocnemis ..
burmeisteri
monticulosus
tridens

Gnathostoma spinigerum

Gnathostomidae

Gnathostominae

Gonatas
albertisi .
fcarolinensis
cetioides ..
differens ..
germari
major
minimus ..
fminor
naviculator
novaebrattanine

Page

315

.. 314, 315
315, BUG, BI
ee 316
po ales, ally)
314

314

309

290

big 303

.. 290, 300

293

336

315

50 SA 146
Sc Fe 317
.. 304, 310

258

Ae 258

.. 194, 258

258

76, 121

12, 76, 93

330

328

40 ae 329
12, 78, 80, 108, 122, 124
AD 109

108, 110, 111

8, 108, 110

oe 109

109, 111, 122

ae 109

108, 110, 122

108, 110, 111

108, 110, 111, 122
109

"Gonatas pumilio

Index.

Page
108, 110, 122

304 |

schellongi TE), Ill, 12%
ftenimbrensis 108, 109, 111
tridentatus Ag 109
’Grus antigone . 293, 294
australasiana 278
communis Dts Boe
paradisea 294
“Gyraulus convexiusculus 341
H
Haemonchus 337
+cervinus det 337
contortus 33338
Hamanma 7 321
Hardella = 308
thurgi .. 307, 308
Heliscus 22, 26
Helodrilus 260
caliginosus trapezoides 260
parvus Ei 260
Heterakidae so KR), 20
Heterakinae ee 289
Heterakis 265, 289, 295, 297, 299, 300
bosia 55 PEO, POL
hamulus .. oe 300
isolonche 289, 290, 291
longecaudata 265, 290, 291
maculosa . 292
papillosa .. 289, 290, 291, 300
vesicularis 289
Heterochilus 76, 86, 87
crinitus 87
occulitesselatus 87
wallacer 76, 87
Hieremys annandalei
Histiocephalus 321
Histiophorus 284
gladius _, 2B, a
Hoplochaetella CYL DBR), 220 9297
Tanomala 223
Humannia 321
Hydra 246
Hydrilla 197, 198, 199
_Hydrobioides nassa 146

Hylobates
Hyperplesthenus
glaber

Ithagenes cruentus

Kachuga lineata
smithii
Kalicephalus willeyi

Kathlania
Kathlanidae
Kathleena arcuata
Kaupioloides
Kaupiolus

trigonophorus
Kiluluma

Labiduris
Labienus
compergus
dohrni
gigas
glaber
gracilis
impar
finaequalis
moluccanus
ptox
Tptoxoides
trigonophorus
Lampito mauritii
Lasioperix
Lemur brunneus
Leptaulacides
Leptaulacinae
Leptaulax
angustifrons
anıbarbis
anipunctus
anna
barbicauda

. 289, 292

.. 304, 308

.. 807, 322

331

= 309
303, 309, 310
282

8, 10, 70103
77,103

332

Ko: i 303
77, 78, 79, 103, 122, 124
104, 105, 107

.. 104, 106

106, 107, 125

106

8, 107

=. 106

104, 105, 106, 107

106, 107, 125

.. 105, 107

.. 104, 106

992

302

Be i 112
12, 13, 111, 123, 124, 126
4,12, 111, 112, 123, 124
uh 116

113, 116, 119

la, 1G

112, 114, 119

Index.

Vili

Page
Leptaulax beccarit i io)
bicolor .. 112, 113, 114, 119, 123
cyclotaenius IS IG TLS), 28}
dentatus .. IDR TNS), TIS), 106
glaber bo IOI I)
glabrwentris HUB. TG, Wiz
himalayae ts > las, 1110
humerosus a oo IU. IN)
klugi ae is se 72
MACASSATIENSIS an se 12, LLG
novaeguineae SY ER 112
obtusidens ee 28 112
planus .. 0% 113, 118, 123
punctipectis De dE 73
roepstorfi .. ade 119
fsambawae .. 114, 118
timoriensis sion lute NS)
ursulus .. se a 112
vicinus .. Fe so 119, IA
Leptoptilus crumenifer Sn 35 319
dubius .. ime be 319
Limnaea ae ae = 195
acumınata Er 19541006
chlamys .. Sc as 195
gedrosiana rectilabrum a 195
Lophocephalus .. mS i 51
Lophophorus impeyanus .. .. 289, 290
Lophopodella jo USO ler
Loris gracilis... os 36 300
lydekkerianus ei oe 300
Lucanus interruntus u 22515163
Lucilius ne 44 bee 51
Lumbricidae .. As me 260

M
Macracis ar AR ie 303
Macrolininae MENT A IAI, We, 12,
125
Macrolinus 5 AE al TO, TE CO) 120212
123, 126
andamanensis By se 6805 83
batesi pu el ae 81, 83
crenatipennis se so Sle SB
Tdepressus 80, 81, 83
duivenbodei 821831123

latipennis Br BDA

Page:
Macrolinus nicobaricus 83
Tobesus 80, 82, 83, 125
rotundifrons 80, 83.
sikkimensis 80, 83
sulciperfectus 82,183, 123.
urus oe Jo Cr on 113, 11995)
waterhousei .. 83.
weberi 83
Macrolobus er 51
fMalagasalus 0 ds}, GS
Telypeatus 69, 70:
studti be sie 69
Malea 148, 149, 178, 181, 184
pomum 181
ringens 181
Manlius hs ae Da 51
Mastochilus 56 U ths, Ts YO, Sr 100, 129
australasicus 99, 100, 103.
capitalis . 9
obliquus . 98
pectinigera 98, 122:
polyphyllus 100, 103, 107
quaestionis 99, 100, 103.
subobliquus oc Be 9
Mastochilus (Analaches) australiensis .. 101, 103:
puberilis .. 210199103
Mastochilus (Cetejus) grabowskii 1020
peltostictus 55 10, 108
sodalis’ 74102103
Mastochilus (Pharochilus) dilatatus 98, 103-
nitidulus .. 98, 103.
politus .. 99, 103
fpunctiger}4 99, 103.
Megacephalon maleo a 290:
Megalotis zerda . .. 269, 322.
Megascolecidae .. .. 202, 226:
Megascolecinae .. . 202, 226
Megascolex; 194.
Megascolides 194
Tprashadi ae 202
Malursus ursinus . 270, 336.
Mermis oh 340
Mermithidae .. 340, 341
Metastrongylidae 338
Microchaetinae .. 258
Micropleura 317

Micropleura vivipara
**Micropleurinae
Microthorax
Milvus govinda ..
Mitrorhinus
Monhystera
tMonhysterides ..
Tpiscicola
Moniligastridae ..
Morosophus
Motacilla alba

o-o

N
‘Naididae
Nais |
communis punjabensis
Tgwaliorensis
paraguayensis

"paraguayensis aequalis

pectinata
raviensis ..
tenuidentis
‘Naja tripudians ..
Nandus marmoratus
Nasoproculus
bifidus
Necator
americanus
suillus
_Necatorinae
Neleides
antillarum
Neleidinae
Neleinae
Neleuops
rhodocanthopoides
Neleus
camerani
cognettii ..
festae
Ninoides
Ninus hondurae rosminiae
nobilii
Numida meleagris
Nycticorax griseus

. 273, 314, 332, 334

Index.

Page

317, 318, 319
so Gilt, Guy)
51

275

10, 11, 51
343

342

.. 342, 343
200

315

195
a 196
<< 166,187
= 198
“= I Us)
oe 197
198
199
199

284

.. 7, 22, 30
7, 30

337

Ix
Page

O
Ocnerodrilinae . 226, 258
Ocnerodrilus ae 5 258
Ocnerodrilus (Ocnerodrilus) occidentalis 258
occidentalis arizonae si 258
Octochaetinae .. 223, 226
Octochaetus 194, 228, 237
barkudensis 228
bishambari 237, 240
fermori 228
Tganeshae 238
Tmontanus 234
Tpachpaharensis 239
Tpaliensis 228
*paliensis riparius 231, 233
Tpallidus . 236, 240
Tprashadi 211, 233
surensis 232
Odontotaenius 29, 26
brevioripennis 28
Oeneus Lil
Ogyges 22, 30
Oileoides 1029229230120
Tparvicornis 23, 24
subrecticornis He 2, ALS TOD)
Oileus Er 22, 23, 25
guatemalensis . 57
heros i
ridiculus .. 24, 25
rimator 25
sargi 25
Oligochaeta 191
Omegarius 78, 108
mmmus .. 108
pumilio 108
Ophidascaris 272, 283
filarıa 272, 287
nalae ae 273
Ophrygonius 75 75 86%, 1291
Taequalis 88, 89
aequidens 88, 89
birmanicus 76, 87, 88
eantorl 86, 88
convexifrons 86
dunsiriensis 86
inaequalis 87, 89

TOphrygonius javensis
minor
quadrifer .
singapurae
wallacei

Oxtyx virginianus

Index.

Page

88, 89

de 86
76, 87, 89
87, 89

293

Oxyuridae 297, 302, 303
Oxyuris .. 302, 303
anthropopitheci His „302,303
compar 303
corollatus 303
coronata .. 303
Ozolaimus 303
P

Paradoxurus hermaphroditus bondar 338
niger 338
Parapelopides 16, 93
Symmetricus 94
Parapertinax Bis 5. 5]
Passalinae > 2, 19, 18, 23, TIAL DE
Passalotaenius ae 22, 20
Passalus 10), 2G, GE yl, aye
abortivus 53, 60, 67
affinis + À, 36), 56
angulatus 68
approximatus 70
assimilis . 38
australasicus 100
australis .. 107
barbatus .. 122.18
basalis 15
bicanthatus 16
bicornis 26
binominatus 64, 68
brasiliensis 49
cantori oh 86
Tcatherinae H31995.00

cayor LL
cephalotus 37
compergus 105
convexus 55, 66
cornutus . 28
corticola 40, 41

CTUSSUS

Passalus curtus ..

cylindraceus
dentatus
dilatatus .
edentulus ..
erosus
Tecuadorensis
exaratus
fronticornis
furcilabris
Tglaber
glaberrimus
goryi
gracilis
grandis
guatemalensis
heros
heydeni
hostilis
impressicollis
inaequalis
incertus
incisus
interruptus
interstitialis
jansoni
laevicollis
languidus
latifrons ..
latupennis
manouffi ..
marginatus
moluccanus
monticulosus
morbillosus
morio
mucronatus
nanus
nasutus
naviculator
neelgherriensis
occipitalis
opacupennis
Topacus
palini
parastictus

Page

53, 56, 66
Ne 14
. 112, 116
97, 98

Se 19
53, 64, 68
53, 56, 66
70

a 40
53, 64, 68
58, 67

42, 43

56, 66

89, 92

44, 53, 57, 66

35, 38

107

86, 87

57, 66

le 25
63, 68, 124

53, 58, 67, 124

60, 67

a 63
53, 54, 66
80, 82

36 71
an AAD, A
106

53, 54, 65
53, 60, 67
52, 65

53, 62, 68
110

ate 86
7, 53, 61, 68
be 43
53, 63, 68
74

Passalus pentaphyllus
pertyi
pilifer
planiceps
planus
platyrhinus
politus
jpolli
polyphyllus
Tprominens
punctatissimus
punctato-striatus
punctifrons
quadricollis
quitensis
recticlypeatus
recticornis
rhodocanthopoides
robustus
Trugosus ..
sansibaricus
sinualus ..
spiniger
fspinipes
spinosus ..
striato-punctatus
teres
timoriensis
toriferus
tridens
tropicus
unicorns
Pavo cristatus
muticus
Paxilloides
brasiliensis
Paxillosomus
alfarı
borellii
cameramı ..
Paxillus
brasiliensis
camerani
erenatus .
leachii
parvus

Index.

Page
49
53, 54, 66

RN 99
53, 62, 67

97, 100

57, 66

53, 62, 68

52, 53, 65

53, 61, 67

53, 55, 66, 125
56, 66

53, 65

52, 53, 65

51,

ee Re 7, 48
3, 11, 44, 48, 60, 124
45, 49, 50

48, 50.

: 45, 50, 51
45, 48, 49, 51, 124
7

Paxillus pentaphyllus
robustus ..
Pelamys chiliensis
Pelopides
burmeisteri
dorsalis
gravidus ..
monticulosus
schraderv ..
simplex
symmetricus
tridens
Pelopinae
Pelops
australis ..
impressicollis
triumphator
Pentalobus
barbatus ..
cayor
duplicatus
fur
klugi
parastictus
punctipectus
sansibaricus
Perdix cinerea
Perichaeta stuarti
Perionyx
Talatus
Tfossus
himalayanus
Tigatpuriensis
m’intoshi
millardi
fminimus
fpokhrianus
*pokhrianus affinis
fpullus
frimatus ..
sansibaricus
fshillongensis
fturaensis
Pertinacides
Pertinacinae

Pertinax

Page

45, 48, 49, 51

45, 50, 51, 52

Ar 289
10042 19, 95,121, 123
94, 95

94, 95, 121

10.93, 34, 95

95, 96

94, 95

95, 96, 121

76, 107

107

123

a 9
5, 10), (68), 712
73, 74
iat

74
72, 74
oe i ey He
70, 73, 74
10; 73, 74

1
= ©

194, 204,

bo Ww po bw by
= ND
ROME

=
we

207, 208,

SG Se)
D oe
SS

. 194, 204

Pertinax pertyi

Petrejinae

Petrejoides

Petrejus
archidonae
eurtus
gracilis
henrici
peruvianus
recticlypeatus

spinosus ..
Phalacrocorax fuseicollis
javanicus
Phanocles
Pharochilus ve To, Ole
Phasianus torquatus
Phaulothorax
Pheretima
biserialis .
elongata .
hawayana
heterochaeta
lignicola ..
posthuma
Phlogoenas luzonica
Phoronaesomus ..
Phoroneinae
Phoroneus
jansomi
quadricollis
Phraortes
Physaloptera
alata
cesticillata
colubri
digitata
Physaloptera praeputialis
Physalopterinae
Pirula
burdigalensis
concinna ..
decussata
dussumieri
fieus
gracilis
investigatoris

148, 149,

Index.

Page
54
7,51
22, 30
5l

Or
—

98, 103, 122

ko
bo
bo
bt bw bd bb KO KO NO NI bo
© D ND DD YH WWD oo oo
— $x@) Ike) Ey RS) PRO), BI BUND pee we)

cm
> Ot Or Cr
—_

= 321
178, 187, 188
188

Sc 188

.. 187, 189
187, 188, 189
187, 188, 189
ae 189
187, 188, 190

Pirula pamotanensis
papyratia
reticulata
tessellata
tricarinata

Platyverres
intermedius

Plesthenus
selon
invitus
lottanaa
mandibularis
quadricornis

scutellopunctatus

Pleurarvinae an
Pleurarius

brachyphyllus
Pleurostylus
Plotus melanogaster
Polycanthopus
Polydelphis a
attennata
hexametra
oculata

Tsewelli

Pontodrilus bermudensis

bermudensis ephippiger
Pontoscolex corethrurus Mr
10010022002 302,6%

Popilius 53, 124
yamazonicus 24, 27, 29
brevioripennis 24, 28, 29
cornutus . 28, 29, 124
granulifrons 55 Aal ALG
Tguatemalae 24, 27, 29
marginatus 24, 27, 29:
intergeneus 24, 27, 29»
purulensis oe 46:
recticornis 2, 20,29

28, 29

striato-punctatus

tropicus .
Porrocaecum
augusticolle
crassum
depressum
Tpristis

Page

cab u 188:
187, 188, 189.

ES la, RASE TE)

3 OT IS!)

20 a 188
3, D, 13, 32, 33, 41, 51, 125
on .. D, 34, 42
12, 76, 77, 79, 96, 121, 123
96

96, 97

ee 9.

. (Os Ts SVU

51, 76

5, 12, 51, 76, 78, 82, 84, 120,

121, 125, 126
ee .. D, 82, 84

US UN SU EN
=

[Soy 1S) SS) 1) KS) TRS)
Se ©
Dow we

Or
©

50 24, 28, 29
. 275, 281, 283, 289:
275, 276, 277

275

219,

.. 280, 281.

Index.

Page

Porrocaecum reticulatum .. 278, 279, 289
serpentulus 277, 278, 280
Potamogeton .. oA ar 195
pectinatus en .. 196, 200
Pristina longiseta En 4 199
Pristis antiquorum à jé 281
perotteti .. : 280
Proculejoides 09, LO 22633, 43, 44, 47, 125
championi bie re 10 AT
crassulus .. 7
granulipennis Ae 7
Proculejus 210522, 23, 3], 43, 47, 51, 125
champvoni se hs 47
nudicostis aA an 7
pubicostis ae ca 2 Bl
quitensis .. 22, Dll, 55
SEMAGOI oc ar so EL, BY
CRUG TL GE 31, 32
Proculinae so 2, 1%, 1183, 32, 124, 126
Procululus 1, 83, 4%, 1%
inca ok ae 7
Proculus D. IO, 8%, ce, 22, 1%
becker ff
densipennis 7

goryl ae ao Fe 43
magister .. ae set i
mandibularis er ar 7
mniszechi ss se (LO, 48
opacipennis sis ke 43
Prosoclitus 29, Sl, Dil
quitensis .. a 1 51
Protomocoelus .. 12, 76, 78, 79, 107, 122, 124
australis .. a a: 107
solomonis oe Fos 107
schraderi a a 76
sternbergi he + 9
Pseudacanthinae 2, I, 18, 22, 124, 1126
Pseudacanthus .. Us 2%, 23), BO, AE
bifidus .. de a)
jalapensis D 8 0094530
solidus . aa so 2 BY
ad nidodens a5 297
fpavonis .. 297, 298, 299

Pseudepisphenus 3, 78, 80, 111, 122, 123, 124
perplexus BS Pe TU
Pseudo-heterakidae a Be 309

Psilomus

Ptichopus
angulatus
borellii

Ptychotrichus

Publius me 10, 32,

crassus

spinipes
Python molurus

reticulatus

Ratufa indiea
Rhabdiasoidea
Rhagonocerus
Rhinoceros bicornis
Rhipsaspis
Rhodocanthopinae

Rhodocanthopordes spiniger ..

Rhodocanthopus ..
biolleyi
incertus
DANSE TS
Spinosus ..
Rictularia plagiostoma
Rietularıınae
Rimor
munitus ..
ridiculus ..
Rimoricus
Rondonia
Rusguniella

Saxicola

rubicola ..
Scalmus :
Schizothorax ee
Sciurus indicus .

pygerythrus
tScolecophilus

flumbricicola
Semicyclus

graye

33, 42, 51, 125
4

314

315

285,

266

Me 266
.. 338, 341
338, 339, 340
70:

XIV

Sertorius
agassizi

Setaria

Seuratia

Severus

Simia satyrus

Solenocyclinae

Solenocyclus
antanarivae
approximatus
exaratus ..
grayi
morbillosus

Sonsinia

Soranus ts
depressifrons
imbellis
intergeneus

Spasalus

Sphaerularia

Spinicaudinae

Spiroxyinae

Spiroxys
Tannulata
contorta ..
gangetica

'Spiruridae

Spiruroidea

Spongilla crateriformis

‘Spurius
bicornis
conradi
dichotomus
Stephanocephalus
Streptocara
Strongylidae
Strongyloidea
Strongyluris
brevicaudata :
campanula
fchamaeleonis
elegans
icosiensis
ornata
paronal
sonsinol

Page

33, 34

317

321

ate 266
11..018,2:06,,.126
3:74,69, 70

ue 48
. 340, 341
297

328

; 328
. 328, 329
a 328
. 328, 329
319

.. 7, 24, 26
10, 11, 51
321

331

7 331
294, 296, 297
297
.. 296, 297
294, 295, 296, 297
. 296, 297
297

297

.… 296, 297

|
|

Index.

Page
Strongyluris streptoesophageus 297
Stylaria lacustris > 200
Subulura .. 300, 302
curvata An 302
Tgalloperdieis 300, 301, 302
hallı 302
olympioi .. 302
sarasinorum 300
seurati 302
strongylina 302
Subulurinae .. 300, 302
Succinea He 341
Sus bengalensis .. .. 268, 330
Synesvus 51
Synhimantus 321
T
Taeniocerus 14, 16, 172.320
bicanthatus 16,17
bicuspis 16, 17, 120
deyrollei .. 119)
mastersi 18
platypus . UG), LE
py :maeus LOT
Tanqua 329
; anomala .. 330
tiara 329
Tarquiniinae de oe 76
Tarquinius 3.4, 12, 18.18, SD 122
123, 124
paradoxus Wu!
Tatius 5 78, 108
Testudo elongata 303, 304, 312, 314
forstenll .. 314
parallelus © 314
travancorica 310, 311, 314
Tetrao tetrix ‘293
lagopus 293
urogallus 293
Tetrarachus 51
Tetraracus centralis 7
nobilii 7
Thryptocerus ae 51
Tiberioides 10.18, 84, 121
austeni 85

Tiberioides borealis

kuwerti
Tiberius kuwerti
Tomeutes pygerythrus
Tonaudia
Toxascaris

leonina

transfuga
Toxeutotaenrus
Tragopan satyra
Tragulus javanicus
Trapezochilus

dorsalis

nobilrs

respectabilis
Triaenurgus

solidus
Trichinellidae
Trichinelloidea
Trichocephalus affinis

crenatus

dispar
Trichopleurus
Trichostigmus

glaber

thoreyi

ursulus
Trichostrongylidae
Trichostrongylinae
Trichurinae
Trichuris ovis

suis

trichiura .
Trigaster
Trigastrinae
Trispiculascaris trispiculascaris
Tristorthus

papuanus

puncticollis

tricuspis .. en
Trochalopterum medidionale
Truquius oc =
Trygon (Hypolophus) sephen
Tubificidae 5
Turnix ne
Typhlops braminus =:

330

51
75, 111, 112, 123
113

112
119,103
337

337

330

237

246

a 288
150417

XV
Page

U
Uneinaria = ae 1 337
Undulifer oe ah 22, 23, 25
incisus .. = ». 24,25
Urocissa oceipitalis 319
Ursus malayanus 336
torquatus 270:

V
Valerius ar 51
Varanus 287, 315, 316.
bengalensis 5 329:
flavescens 315, 316, 329:
nebulosus 315, 329
salvator .. .. 269, 287, 315, 329
Vatiniinae 51
Vatinius 51
Vellejus 103
compergus Ge a 105.
Verres 3 sh 0%, 88, att, 20), Bill
camerani as er 7
cavicollis 13, 34, 40, 41, 47
cavilabris 5S ae 8.
corticola .. 13,-34, 41
furcilabris 24, 40, 41
intermedius 41, 42
sternbergianus ce Ae 8
sternipunctus 34, 40, 41
vernicatus aA a 8
Veturius 32, 33, 34, 35, 5l
assimilis . 34, 38, 39:
Tboliviae .. 34, 38, 3%
cephalotus 37, 39
criniceps .. 34, 37, 39
heydeni 34, 38, 39:
isthmicus BE ae 8
peruvianus 8
platyrhinus © oe 36, 39
punctatostriatus = Ae 8
similliimus 36, 39
sinuatocollis by 35
Tsinuatosulcatus so Be (Bin, Slo, Br
sinuatus .. 36, 37, 39
fspinifer .. 34, 36, 39
yunicornis 34, 36, 39

XVI

Vindex
agnoscendus
sculptilis ..
fsynelytris

Vipera russelli .

Vitellinus

Viverricula malaccensis

Vulpes
bengalensis
leucopus .

Vultur monachus

Page

LOI M13. 43, A
UOTE AR A

i 47

45, 47, 125

331

336

270

269, 270, 322, 336
336

275

Index.

Wallago attu
Xiphias gladius

tZanclophorus
Tannandalei
Tkempi

Zosteroth vw

Page

… 309, 310

310, 311, 312, 313
303, 312, 313

CALCUTTA: PRINTED BY SUPDT. GOVT. PRINTING, INDIA, 8, HASTINGS STREET.

a
HAE
AN VE

MEMOIRS

INDIAN MUSEUM

Vol. VII, No. 1.

A Contribution towards the Revision
of the

Passalidae of the World.

onian Insti
‚ns = If. U EN
& YoN

F. H. GRAVELY, D.Sc.
Assistant Superintendent, Zoological Survey of India.

, \ °° F4
“ional Muse! /

CALCUTTA
PUBLISHED BY THE DIRECTOR, ZOOLOGICAL SURVEY OF INDIA.
SUPERINTENDENT GOVERNMENT PRINTING, INDIA
DECEMBER, 1918

Srice Seven Rupees.

À Contribution towards the Revision of the

Passalidae of the World.

BY
Foote GIV AWE ENG DSc.

Assistant Superintendent, Zoological Survey of India.

(With Plate I.)

HEN preparing my “Account of the Oriental Passalidae” (1914c) the lack of a
representative collection of American and African forms prevented me from
considering the relationship to other groups, borne by the Indo-Australian groups to which
the Oriental forms belong. Shortly after the publication of that paper, however, the
Passalidae collected by Mynheer J.R.H. N. Van de Poll were offered for sale. These were
bought by the Trustees of the Indian Museum, who thereby added a particularly fine set of
Indo-Australian species to their collection, together with much material for comparison
from America and Africa. My thanks are due to them for the encouragement which they
have thus given me to extend my investigations, so that they may include a study of the
classification of the whole family.

I have further to thank Mr. G. J. Arrow for assistance in various forms, including the
loan of specimens from the British Museum ; Mr. C. Holman-Hunt for material from the Malay
Peninsula; M. R. Vitalis de Salvaza for material from French Indo-China; Mr. C. F. Baker
for material from the Philippines; and M. Guy Babault and Mr. G. E. Bryant for material
from various localities.

Morphology of the Head.

The study of a general collection of Passalidae, from parts of the world other than the
Oriental Region, at once throws light on the question of the morphology of the anterior part
of the head, a question which seemed most difficult before. It confirms as true for most
species of Passalidae the suggestion, made on p. 337 of my “ Account of the Oriental
Passalidae,” that “ the whole of the upper surface of the anterior part of the head between
the supra-orbital ridges and in front of the frontal ridges is frons, the whole of the clypeus
being doubled beneath this out of sight ;” but it shows that this is not true of all species,
and that the groove which I supposed to represent the suture between the clypeus and
frons is probably situated beyond the lateral extremities of the former plate.

The most primitive surviving forms of Passalidae appear to be included in the genera
Oileoides and Popilius, as these are defined below; and Ovleoides subrecticornis (see fig. 1, 1
on next page) may conveniently be taken as an example of them.

In this species the clypeus is not hidden, but is exposed as an extensive transverse plate
above the labrum. The labrum is attached by a well developed membrane, which extends

beneath it to the lower posterior margin of the clypeus, and not to its anterior margin.
B

2 Memoirs of the Indian Museum. [Vor. VII,

Thus two parts of the clypeus may be recognised, namely, a dorsal free part and a ventral
part closely opposed to the membrane uniting it to the labrum. The posterior part of the
free surface of the ciypeus is sunk in the anterior margin of the frons, the two plates bemg,
however, separated by a well marked suture ; and the posterior angles are united with the
adjoining portions of the frons to form a pair of small tubercles. These tubercles are those

to which the name “ outer tubercles” was given in my previous paper, a name which may
conveniently be retained for them.

Fıe. I.
1. Oileoides subrecticornis (Kuwert). Anterior part of head. The outlines of the plates of the upper
2. Chondrocephalus granulifrons (Bates). - surface indicated by continuous, and those of the lower surface by
3. Passalus affinis, Percheron. dotted, lines.
4. Passalus affinis, Percheron. Median section of upper and anterior parts of head.

a.a.—anterior angles of head. f.v.s.—frontovertical suture.
an.—cavity for insertion of antenna. ı.t.—inner tubercle.
cl.f.—free surface of clypeus. lbr.—labrum.

cl.o.—opposed surface of clypeus.
e.t.—central tubercle.

fr.—trons.

fr. r. frontal ridge.

lbr.m.—membrane between labrum and clypeus.
o.t.—outer tubercle.

s.or.—supraorbital ridge.

v.t.—ventral tubercle.

Having identified the clypeus in forms such as Ovleordes subrecticornis, in which no doubt
with regard to it can arise, the fate of the free surface of this plate in other forms can readily
be followed.

In species belonging to the subfamily Pseudacanthinae (of this paper) the clypeus is
always separated from the frons by a definite suture, though this varies greatly in shape and
in some species the inner tubercles are situated so as to interfere somewhat with its continuity.
In the Proculinae, in a few species of Passalinae, and perhaps also in the Aulacocyclinae, the
clypeus has the same structure as in the Pseudacanthinae. but the suture is absent. In all
other species the free surface of the clypeus is reduced to (at most) a narrow transverse band,
situated below the anterior margin of the head, and terminated laterally by a pair of small
downwardly directed tubercles, the ‘‘ ventral tubercles ” of my previous paper (1g14c, text
fig. 1, p. 181). These “ ventral tubercles ”—really the anterior angles of the reduced free
surface of the clypeus—are extremely persistent, and their presence or absence almost always
enables one to determine whether the anterior margin of the head really conceals this surface
of the clypeus or includes it. To see them clearly, however, it is often necessary to remove
the labrum. The outer tubercles are normaliy situated immediately above or a little to the
outer side of them ; but in some forms these tubercles tend to come nearer together and may

1918.] F. H. Gravety: Passalidae of the World. 3

be situated on the inner side of them. That they still mark the lateral extremities of the
tree surface of the clypeus in these cases is shown by the fact that they are joined by the
sides of the opposed surface of the clypeus. The accompanying diagrams (fig. i) will
help to illustrate the various stages in the suppression of the clypeus. With them may be
compared figs. v, 11-15 and vi, 6-11 on pp. 34 and 45, illustrating in a less diagrammatic
fashion its suppression in Verres and Platyverres and in Paxillus respectively.

The presence, in most American and African as well as Indo-Australian Passalidae, of
definite frontal ridges and of central, inner and outer tubercles is sufficient indication of the
morphological importance of these structures. The outer tubercles, as has just been shown,
are formed in the more primitive species by a fusion of the frons and clypeus about the
posterior angles of the latter. The central tubercle is normally situated at the angle formed
by the union of the two frontal ridges which appear, from the evidence afforded by certain
Indo-Australian species, to mark the position of part of the line separating the frons from
the vertex. The inner tubercles appear to mark the point at which this line, usually quite
imaginary for a short distance beyond this point, bends more or less backwards towards a
suture a little to the inner side of the supraorbital ridges, in which it commonly ends (see
Gravely, 1914¢, p. 184). This suture is easily seen in imperfectly hardened specimens of
various groups, but is less distinct in those which have become thoroughly hard and black.

The homologies of the tubercles found in genera such as Leptaulax and Tarquinius, in
which more than one pair of tubercles is situated on the anterior margin of the head, with
those found in genera with only one pair so situated, are more difficult to determine than
appeared to be the case when Indo-Australian forms only were under consideration.

A comparison of Tarquinius with Pseudepisphenus leaves little room for doubt that the
more medially situated pair of tubercles in the former is homologuous with the pair of inner
tubercles of the latter (see Gravely, 191 4c, pp. 328-329), and in the absence of any evidence
to the contrary it would be natural to assume the same homologies for the tubercles of
Leptaulax, and of genera from the Ethiopian Region with similarly arranged tubercles.

But in the case of Ethiopian genera there is strong evidence in favour of different
homologies. This is best illustrated by reference to the group characteristic of Madagascar, a
group which is separated from the group found on the mainland of Africa by the possession
of a pair of tubercles situated on the frons in the angle made by the fronto-vertical suture
with the anterior margin of the head (see below pp. 68-69).

The most primitive of the Malagasy genera appears to be a new one described below
(pp. 69-70, fig. vill, 1) under the name Malagasalus. In this genus the free surface of the
clypeus, though almost vertical, is fully exposed. The outer tubercles, with a pair of well
developed inner tubercles close behind them, project above it and are separated by a space
equal to scarcely as much as two-thirds of its breadth. In Solenocyclus, the next genus of
the series, the inner tubercles are also well developed and are situated further back from
the outer tubercles ; and an additional pair of marginal tubercles is present immediately
above the ends of the clypeus. The outer tubercles, as pointed out above (p. 2), are formed
in the first mstance about the posterior angles of the clypeus ; and their closer approximation
one towards the other in forms such as Malagasalus leaves space in which the tendency
towards tubercle-formation at this point can manifest itself over again. Presumably, therefore,

4 Memoirs of the Indian Museum. [Voz. VII,

this additional pair of marginal tubercles is formed as a result of this tendency. Solenocyclus
is too closely allied to Pentalobus and Erionomus to admit of any doubt as to the identity
of the homologies of their cephalic tubercles. The cephalic tubercles of certain species of
these however, bear the closest possible resemblance to those of the Indo-Australian
genus Leptaulax ; and there does not appear to be any conclusive evidence to show whether
the homologies of Leptaulax are the same as those of Tarquimus, as previously suggested, or
may not really be the same as those of Hrionomus. In the former case the inner and outer
pairs of marginal tubercles will be the true inner and outer tubercles respectively, and the
rudimentary tubercles sometimes formed on the frontal ridges at the point where these bend
forwards will be secondary structures. In the latter case these rudimentary tubercles will
represent the true inner tubercles, the inner pair of marginal tubercles will be the true outer
tubercles, and the outer pair of marginal tubercles will be secondary structures.

It was suggested in my “ Account of the Oriental Passalidae ” (p. 330) that the Leptau-
lacinae were of comparatively recent origin, and were still spreading from some distributional
centre towards the middle of the Indo-Australian area, into the outlying parts of this
area, suck as the Indian Peninsula and Ceylon. If this is so—as all available evidence
regarding their distribution seems to indicate—it is most unlikely that they are at all
closely related to any but Indo-Australian groups, a suggestion whose correctness is
confirmed by the fact that the anterior lower tooth in the Leptaulacinae is of a form found
only in Indo-Australian groups (see below, p. 9). Presumably, therefore, the origin of
the Leptaulacinae is to be sought among Indo-Australian forms. But there is no definite
evidence to show that they are in any way related to Tarquinius, the ouly known Indo-
Australian Passalid to which they bear even a superficial resemblance. The fact, moreover,
that in most Indo-Australian forms the outer tubercles tend to be separated by a distance
which is less than the width of the clypeus, makes it easy to suppose that the evolution
of the Leptaulacinae may have been on lines parallel to those of the evolution of Ethiopian
forms.

In view of this uncertainty as to the homologies of the cephalic tubercles in the Leptau-
lacinae it seems best to continue to use the terms “imner” and “outer” tubercles in the
connection in which they were used in my previous paper; but in addition to insert the
word “ marginal,” when speaking of the Leptaulacinae, in order to show that in this case
the terms are used in a descriptive sense which is not necessarily in accordance with
morphology.

Flightless Species.

In several groups of American and Indo-Australian Passalidae there are species which
have lost the habit of flight. This tends in all cases to produce certain modifications of
structure :—

1. The union of the lateral and intermediate areas of the metasternum.
2. The union of the elytra in the middle line.
3. The shortening of the elytra and the rounding of all their contours.

4. The reduction of the wings, which appear to become thereby more efficient
stridulating organs.

1918.] F. H. GRAVELY : Passalidae of the World. 5

Flightless species consequently tend to bear a strong resemblance to one another,
especially as regards the general shape of the body, and Bates (1886, pp. 2-3) divided the
American Passalidae into two sections on this character alone (see below, p. 6).

The extent of the modifications differs greatly in different species, and the relation
which they bear one to another is not always the same. Thus in American species the union
of the elytra tends to precede the modification of the metasternum, while in Indo-Australian
species the reverse is the case. Similarly, reduction of the dentition, which seems to be to
some extent associated in American groups with loss of the habit of flight (see below, p- 9)
is only found in Indo-Australian forms (where it reaches 1ts maximum development) among
species which show no indication of this.

In the Indian Pleurarvus brachyphyllus, in which the wings are fully developed, the
union of the elytra is not indicated in the pupa and is imperfect or absent in the newly
hatched adult. This appears to be the case in some other species also, and it seems likely
that the elytra may remain separate throughout life in certain individuals. I have not seen
pupae of any of the more highly modified species.

Genitalia and Sexual Distinctions.

Very little appears to be known of the genitalia of the Passalidae. Sharp and Muir
(1912, pp. 579-580, pl. xliv, figs. 11-13a@) found two forms of male genitalia to exist, one in
which “ the basal piece and the lateral lobes form one piece, either by consolidation or the
suppression of the basal piece ” and one in which “ the tegmen consists of two distinct
pieces, the basal piece and the lateral lobes.” The former they found in the genus
Aubade the latter in all the other genera they examined ; but their observations were
very restricted. I have been able to add to these observations to some extent; but the
results were not such as to warrant any extensive investigation, since all the genitalia
examined proved to be very much alike, except in the Aulacocyclinae. Here both the types
described by Sharp and Muir occur, one in one group of the subfamily and the other in the
other, showing that the difference they found in Aulacocyclus is not a distinctive character
of the Aulacocyclinae as a whole, as suggested in my previous paper (p. 101). The only
other positive result of my investigations was a curious fact which emerged in connection
with species in which the central tubercle varies greatly in size. In these it was found that
the specimens in which it was largest and best developed were females, and not males as

would be expected by analogy with other groups.

Classification, etc., including notes on the structure of the mandibles.

Although the Van de Poll collection is a remarkably fine one, it is by no means complete.
There can, I think, be little doubt that a considerable number of described genera and species
have no separate existence ; but the absence from the collection of such distinctive forms as
Platyverres intermedius, and of other well-known species, affords sufficient proof that the
names of the missing forms are not all to be lightly relegated to synonymy. The number
of new species In so incomplete a collection, on the other hand, seems clearly to indicate
that the Passalidae of the world as a whole are less fully known than are those of the

Oriental Region dealt with in my previous paper.

6 Memovrs of the Indian Museum. Wor. MH:

In order to prevent subsequent confusion I have redescribed, or have at least directed
attention to the distinctive characters of, every species that I have seen, excepting only the-
Oriental ones described in my previous paper; and I have figured a large proportion
of them. This is made essential by the unsatisfactory condition of most of the existing
descriptions, a condition which has united with the incompleteness of the collection before
me to make the compilation of a satisfactory synonymy of the species very difficult, if not
impossible. No attempt has been made to deal with the synonymy of species, the
references given being in all cases to original descriptions or to others on which I may have.
relied in making my determinations.

The forms dealt with in my “ Account of the Oriental Passalidae ” naturally receive less
detailed treatment than the others. The keys to the determination of genera published in
that work are, however, repeated with such alterations as further study has shown to be
desirable ; and, except in some of the smaller genera, keys have been given to the identi-
fication of all species known to me, although in a few cases these are practically identical
with those already published.

The first serious attempts made to divide the Passalidae up into genera were those of
Kaup in 1868-9 and 1871, respectively. In his “ Monographie der Passaliden,” published in
the latter year, he set forth a remarkable conception of the Animal Kingdom, which led him
to postulate a series of subdivisions into co-ordinated series of fives. Believing, as he did,
that none of his five sub-families of Passalidae could contain more than five groups, that no.
group could be composed of more than five genera, and no genus of more than five species,
and believing that corresponding species, genera and groups were to be found respectively in
all or almost all genera, groups and sub-families, his system of classification inevitably led
to a considerable amount of wide separation of closely related genera and species.

In 1886 Bates introduced a number of changes into Kaup’s classification of the American
species. He brought together in one section all those in which the elytra are relatively short,
and more or less protuberant in the middle of the base, and into another all the rest—.e.,
those in which the elytra are moderately long with broadly emarginate base—subdividing each
of these sections primarily according to the lengths of the antennal lamellae. Unfortunately
for this classification the shape of the elytra is correlated with the modification of the wings
for stridulation and the loss of the power of flight, changes which not only appear to have
taken place in some of the most highly specialized forms of several different American groups,
but are also found among widely separated Indo-Australian genera (see below, p. 125); and.
the lengths of the antennal lamellae rarely seem to have much phylogenetic importance,
being more or less variable in a number of genera, perhaps in all (see also Gravely, 19146,
pp. 180 and 182).

Kuwert’s elaborate “‘ Passaliden dichotomisch bearbeitet ” is still more lacking in any
sense of phylogenetic values, and disregards in addition the facts of geographical distribution.

Zang and Arrow have done much useful work in the direction of clearing up various items
of the confusion thus produced, and my “ Account of the Oriental Passalidae ” has, I hope,
helped to reduce the classification of the forms with which it deals to order; but Kuwert’s
remains the most recent monograph of the Passalidae of the world. In my “ Account of the
Oriental Passalidae ” I have recorded all references known to me relating to Oriental genera.

1918.] F. H. Gravety: Passalidae of the World. 7

and species. The following appear to be the only forms not referred to either in Kuwert’s
-work or there : —
A.— AMERICAN Forms.
Coniger, n. gen. with Rimor ridiculus, Kuwert, as type, Zang, 1905c, p. 232.
Epiphoroneus, n. gen. with Passalus occipitalis, Eschscholtz, as type (Phoroneinae of Kuwert),
Arrow, 1907, pp. 459-460.
Eumelus nasutus, Arrow, 1907, p. 459.
Nasoproculus, n. gen. with Passalus heros, Truqui, as type, Zang, 1905c, p. 226.
es bifidus, Zang, 1905¢, p. 232,=Orleus heros, part, Kaup, nec Passalus heros, Truqui.
Neleides antillarum, Arrow, 1907, p. 452.
Neleus camerani, Pangella, 19055, pp. 9-11.
5, cognettii, Pangella, 19055, pp. 13-15.
» festae, Rosmini, 1902, p. 8.
Ninus hondurae var. rosminiae, Pangella, 1905a, pp. 11-12.
„ nobili, Pangella, 19055, pp. 5-7.
Paxillosomus alfari, Pangella, 1905a, pp. 9-11.
borelliz, Pangella, 19055, pp. 3-4.
camerani, Rosmini, 1902, pp. 4-5.

Paxillus parvus, Casey, 1897, pp. 644-645.
Petrejus archidonae, Arrow, 1907, p. 450.
à henrici, Rosmini, 1902, pp. 6-7.
= peruvianus, Arrow, 1907, pp. 456-457.
» spinosus, Arrow, 1907, pp. 457.
Proculejoides crassulus, Casey, 1897, pp. 642-643.
A granulipenms, Zang, 1905a, pp. 229-231.
Proculejus nudicostis,! Bates, 1886-1890, p. 383.
Procululus, n. gen. with P. inca, n. sp. as type (Petrejinae of Kuwert), Zang, 19050, pp. 225-227.
Proculus beckeri, Zang, 1905a, p. 315.
,, densipennis, Casey, 1914, p 374
„ mandrbularis, Casey, 1914, p. 374.
5, magister, Casey, 1897, pp. 641-642.
Ptichopus borelliz, Rosmini, 1902, p. 10.
Publius spinipes, Zang, 1905a, pp. 231-232.
Rhodocanthopus biolleyi, Pangella, 1905a, pp. 3-4.
Rimor munitus, Casey, 1897, pp. 643-644.
Soranus depressifrons, Bates, 1886-1890, pp. 384-385.
„ mbellis, Casey, 1897, pp. 645-646.
Spurius conradi, Rosmini, 1902, p. 2.
„ dichotomus, Zang, 1905a, pp. 227-229.
Tetraracus centralis, Arrow, 1907, p. 458.
x nobilii, Rosmini, 1902, pp. 5-6.
Triaenurgus solidus, Arrow, 1907, pp. 452-453.
Verres camerani, Pangella, 1905a, pp. 7-9, text fig.

1 Zang, who knew it from the description only, placed this species in the genus Proculejoides (1905a, p, 229).
_According to Arrow this is incorrect (1907, p. 450). I have not seen a specimen, but the description shows it to have
-the tridentate mandibles and hairiess elytra of the genus Pseudacanthus as this is defined below.

8 Memoirs of the Indian Museum.

Verres cavilabris, Casey, 1897, pp. 646-647.
,, sternbergianus, Zang, 1905a, p. 315.
» vernicatus, Casey, pp. 647-648.
Veturius isthnucus, Arrow, 1907, pp. 453-454.
3 peruvianus, Arrow, 1907, p. 455.
3 punctatostriatus, Arrow, 1907, pp. 454-455.

B.—ETHIoPIAn Forms.

Didymus congoensis, Arrow, 1907, p. 463.

ae crassus, Arrow, 1907, p. 465.

» Curvilineatus, Arrow, 1907, pp. 462-463.

», laevisternus, Arrow, 1907, pp. 463-464.

nH latipunctus, Zang, 1905a, pp. 315-316.

55 ruwenzoricus, Arrow, 1907, pp. 464-465.
Erionomus platypleura, Arrow, 1907, p. 461.
Eumelosomus affinis, Arrow, 1907, pp. 465-466.

aloysii sabaudiae, Pangella, 1906, p. 1-

C.—Inpo-AUSTRALIAN FoRMS.

Analaches bicavis, Zang, 1905a, pp. 241-242.
> brachymetopus,! Zang, 1905a, pp. 30-31.
=A dubius, Heller, 1910, pp. 20-21.
ss infestus, Heller, 1910, p. 20.
ce laevigatus,? Zang. 19034, p. 339 ; and 1905a, pp. 29-30.
5 laticauda, Zang, 1905a, pp. 240-241.
33 paraplesius, Zang, 1905a, pp. 238-240.
punctithoraz,” Zang, 1903a, p. 339 ; and 1905a, pp. 28-29.
Aulacocyclus rouxi, Heller, 1916, pp. 352-35.
Cetejus acutangulus, Heller, 1910, pp. 18-19.
>» injans, Heller, 1910; p. 18.
,, schenklingi, Heller, 1910, pp. 19-20, text-fig. B.
„ sodalicus, Zang, 19065, p. 25.
Episphenoides pectiniger, Heller, 1910, pp. 17-18, text-fig. A.
Gonatas cetioides, Zang, 1905a, p. 316.
Hyperplesthenus glaber, Gravely, 1913, pp. 106-107, text-fig. 2A.

[ VoL.

VII,

Kaupioloides, n. gen. with Kaupiolus trigonophorus, Zang as type, Gravely, 1913, pp. 103-105, text-

inex, I
Kaupiolus trigonophorus, Zang, 1905a, p. 316.
Labienus (2?) gracilis, Heller, 1910, pp. 16-17.

1 Subsequently transferred to the genus Cerejus (Zang, 19054, p. 238, footnote).

? Originally named Epilaches owing to Kuwert’s confusion of the two generic names (Zang, 1905a, p. 24) ; subseque:itly

transferred to the genus Cetejus (Zang, 1905a, p. 238, footnote).

1918.] F. H. GRAvVELY : Passelidae of the World. 9

Mastochilus capitalis,} Blackburn, 1900, pp. 209-210.
subobliquus, Tryon, 1892.

Pelops triumphator, Zang, 19045, pp. 182-184.

Plesthenus mandibularis, Heller, 1900, p. 11, pl. fig. 5.

: scutellopunctatus, Zang, 1903a, p. 339.
Protomocoelus sternbergi, Zang, 1905a, pp. 236-238.
Tristorthus papuanus, Heller, 1910, p. 15, pl. figs. 15-154.
Tristorthus puncticollis, Heller, 1916, p. 353.

In view of the striking correlation which has been shown to exist, in certain Indo-
Australian Passalidae, between classification and distribution (Gravely, 19145), it is natural,
when seeking for sound principles on which to base the classification of the familv as a
whole, to consider first whether it may not be possible on purely morphological grounds to
regard the American and Ethiopian Passalidae respectively as series independent both of
one another and of Indo-Australian forms.

It will be remembered that among the Indo-Australian Passalidae the Aulacocyelinae
stand by themselves, widely separated from all other forms (Gravely, 1914¢, pp. 191-192).
They are also widely separated from American and Ethiopian forms and need not be further
considered in this connection. In all other Indo-Australian Passalidae, except certain highly
asymmetrical ones with reduced dentition, the lowest terminal tooth is directed inwards
and is situated beside the anterior lower tooth, which is broadly triangular as seen from
above and tends to be flattened above and below. In American and Ethiopian forms, on
the other hand, none of which are asymmetrical, either the terminal teeth are situated in a
line one above another; or the anterior lower tooth (on the right side at least) is more
columnar and tends to be flattened in front and behind, or may be concave behind much
as it the Aulacocyclinae, or both characters may be present. These characters are more
marked in some species than in others, but in greater or less degree they are of universal
application ; and they are associated with various other, often more striking, differences
of more limited application.

The reduction of the dentition takes place mainly in highly specialized genera of the Indo-
Australian and American series, genera whose relationships are, as a rule, clearly indicated by
other characters. Among the former it is confined to the more highly asymmetrical genera,
and among the latter it is found chiefly in genera composed wholely or in part of flightless
species. It is easy to follow the manner in which the reduction is carried out in the former ;
but in the latter it is often more difficult, the reduction being more abrupt. In the Indo-
Australian series it is always due to the fusion of the anterior lower tooth with the lowest
terminal tooth (see Gravely, 1914¢, text-fig. 7, p. 314). In the American and Ethiopian
series, with the possible exception of the genus Proculus, it seems always to be due to
the fusion of the two lowest terminal teeth, though this is apt to be accompanied by a
dorsoventral flattening of the anterior lower tooth, which thus loses its distinctive shape.

The reduction of the dentition in the American and Ethiopian series may most
conveniently be illustrated by reference to the genera of the Proculejoides group of Passalinae

t Mastochilus capitalis, Blackburn=Episphenoides quzestienis, Kuwert. See Zang, 1905c. p. 222.

10 Memoirs of the Indian Museum. [Vor. VII,

(fig. ii, 1-3). Im Chondrocephalus the dentition is normal, though the right anterior
lower tooth is less distinctly columnar than usual. In Vindex the dentition is reduced,
but the double origin of the lower of the two remaining terminal teeth may often be
seen, especially in the right mandible of unworn specimens of V. agnoscendus. The anterior
lower tooth of the left side is more conical in this genus, with dorso-ventral rather than
antero-lateral compression, though that of the left side is still much broader than that of the
right: and there is a more or less indistinct tubercle at the base of this tooth on both
mandibles. In Proculejoides all trace of the double nature of the two remaining terminal
teeth is lost, the anterior lower teeth of the two sides become still more alike, and the
tubercles at their base become more conspicuous.

Ofthe genera with modified dentition in other families, Proculejus has teeth exactly like
those of Proculejoides, and the only species of Publius known to me has teeth not unlike

those of Vindex, but has no trace on either side of the
third terminal tooth and has more markedly
| 2 if asymmetrical anterior lower teeth. The teeth of

Proculus, the most striking of all these genera, are,

D however, more puzzling. Superficially they usually
resemble those of Proculejoides ; but between what
\ 2 A, would be the anterior lower tooth in that genus and

the large tubercle at its base there is ordinarily a scar,

lef: by the breaking away ofa columnar tooth which
seems only to persist in imperfectiy hardened
\ x Y specimens. This columnar tooth (see fig. 11, 4) has

the form of the type of anterior lower tooth charac-

teristic of the American series ; and it 1s impossible
to be sure whether the tooth in front of it forms the
4 5 n most anterior part of a single but highly complex

anterior lower tooth, or ıs a lowest terminal tooth,

Fie: II. reduced in size and situated behind and on the inner

1. Chondrocephalus granulifrons (Bates). side of the middle and upper terminal teeth as in the
2. Vindex agnoscendus (Percheron).

3. Proculejoides championi (Bates). Indo-Australian series.
4. Proculus mniszechi, Kaup.

The parts of the mandibles in front of and
excluding the movable tooth, illustrating modi-
fications in the dentition of American Passalidae.

The more typical American and Ethiopian forms can be distinguished from each other
by the presence of one or more pairs, respectively, of marginal tubercles between the
anterior angles of the head ; but this distinction does not at first sight appear to be of
universal application, since at least two genera, Mitrorhinus and Stephanocephalus, which have
been accepted as American, have the additional tubercles characteristic of African genera.
Zang, moreover, notes records of the American genus “ Passalus ” (= Popilius of the present
paper) from the Congo, Senegal and Madagascar; and in the Van de Poll collection several
specimens of Pentalobus sansibaricus bear the record “ Bolivia ”; while of two specimens of
a new species of Brionomus, described below under the name L. trichostigmoides, one bears

1918. ] F. H. GRAVELY : Passalidae of the World. 11

the record “ Dar-es-Salaam ” and the other “ S. Catherina, $. O. Brazil (Staudinger).” If
these records are correct, and the species to which they refer really do occur both in America
and in Africa or Madagascar, the occurrence in America of genera belonging to the group
typical of Africa would be very natural. But in the absence of direct personal evidence by
a collector I find it easier to believe them to be due to mistakes in labelling, numerous
though they are. The position of Mitorhinus and Stephanocephalus therefore requires
special consideration.

Judging from Kaup’s figure (1871, pl. vi, fig. 3) Pussalus punctifrons, Dejean, the type
of the genus Mitrorhinus, would seem to be from America and to belong in all probability,
to the genus Passalus as this is defined below; but Kaup identifies the species with
Percheron’s Passalus cayor, a species the type of which is recorded as coming either from
Senegal or from Brazil. As Percheron’s figure (1835, pl. v, fig. 2) shows this to be an insect
of the African type, the former is doubtless the correct locality. There can, I think, be little
doubt that cayor belongs in reality to the African genus Pentalobus ; and I suspect that
both Kaup and Kuwert have coafused prolongations of the anterior angles of the head
with the tubercles on the inner side of these angles characteristic of African forms.
Probably this has also been the case with the genus Stephanocephalus, the only species
figured, Percheron’s Passalus hostilis (1841, pl. Ixxvu, fig. 4), whose locality was not known
to its author, being in all probability African, although Kaup and Kuwert have applied the
name to an American species, and have placed other American species in the same genus
with it. For the present, then, the form of the anterior margin of the head may be
regarded as separating American from Ethiopian groups ; but further evidence on the point
is much to be desired, especially as the aberrant genus Ptichopus is to some extent
transitional between the two.!

Another character by which the American and Ethiopian groups may be separated is
afforded by the posterior plate of the prosternum. In the former this 1s usually more or
less narrowed behind, and is often pointed ; whereas in the latter it is always more or less
parallel sided and broadly truncate. This plate has the African character in most species
of the American genus Paxillus, but I know of no other exceptions to the rule, although the
distinction is not equally well marked in all species.

The Ethiopian genera fall conveniently into a single subfamily Solenocyclinae. The
American genera, on the other hand, fall into three groups, each in my opinion of sufficient
size and distinctness for recognition as a separate sub-family.

The first of these, to which the name Pseudacanthinae may be given, is distinguished
chiefly by the presence of a well marked clypeo-frontal suture. Traces of this suture are
found also in the species described below under the name Chondrocephalus quinquecornutus,
and it is more or less complete in Vindex agnoscendus. The former of these species may be
regarded as in this character transitional between the two most primitive genera ot the
Pseudacanthinae and Passalinae respectively ; and the latter agrees in every other character
with the Passaline genus Vindex and not with any Pseudacanthine genus. In spite of these
exceptions the presence of the suture in question remains the most distinctive single

1 See also below, p. 13, footnote !.

12 Memoırs of the Indian Museum. [Vor. VII,

character running through the Pseudacanthinae. The second American sub-family may be
called the Proculinae. It includes forms in which the outer tubercles are rudimentary or
absent ; the clypeus is almost always exposed, though never separated from the frons by a
distinct suture. The third and last American sub-family, the Passalinae, includes forms
in which the outer tubercles are well developed and usually very widely separated, the
clypeus being hidden in almost all species.

The classification of the Indo-Australian Passalidae was recently revised (Gravely,
1914C, pp. 191-204, 316-318 and 328-330). They fall into two distinct sections, of which
one forms the sub-family Aulacocyclinae. The results of my previous work on the
classification of the other section have been summarized as follows (1914c, pp. 336-337):—
“ The second section has been sub-divided into the Pleurarvus, Aceraius, Macrolinus,
Kaupioloides, Protomocoelus, Hyperplesthenus, Gnaphalocnemis, Plesthenus, Gonatas,
Tarquinius and Leptaulax groups. Of these groups the first three and the last two
appear to be of most importance, and have been provisionally ranked as subfamilies,
the remainder being put together into a single subfamily which takes its name from the
genus Gnaphalocnemis. Their rank cannot be finally settled without reference to American

and African species.’

The number of subfamilies thus recognized is undoubtedly too great in comparison with
the classification of American and African forms outlined above, and may advantageously
be reduced to two. One of these, the Leptaulacinae, remains as defined in my previous
paper. The second, which may be called the Macrolininae, includes all the rest.

The above enumerated subfamilies of Passalidae may now be defined thus :—

/ The distal ends of the anterior coxae projecting beyond the
surface of the intercoxal portion of the prosternum, which is
entirely hidden except in aberrant Chinese and Japanese
forms, where it is exceptionally strongly keeled; the usual
paired cephalic tubercles absent, the middle lower tooth
1 almost always fixed U 2 an .. Aulacocyclinae, p. 13.

The distal ends of the anterior coxae about on a level with the
general surface of the intercoxal portion of the prosternum,
which is visible between them ; at least one pair of the usual
paired cephalic tubercles present in most species ; the middle

\, lower tooth always jointed on to the mandible .. = de o0 2.
The anterior lower tooth more or less columnar, at least on the
right side, being compressed before and behind or even
concave behind ; or all three terminal teeth directed forwards,
and arranged in a line one above another ; or the dentition
5 incomplete ; the head always symmetrieal (American and

3 African forms) ve aS 36 Le os ae Bo
"he anterior lower tooth conical, compressed above and below ;
the lowest terminal tooth directed inwards and set a little
behind the other two; the head often asymmetrical—always

V so in forms with reduced dentition (Indo-Australian forms) .. as ce 6.

1918.] F. H. GravELY: Passalidae of the World. 13

No secondary tubercles or angular processes present on the

anterior margin of the head (or behind the elypeus when this

is exposed) between the outer tubercles and anterior angles ;

the posterior plate of the prosternum almost always much

narrowed, often pointed, behind (American forms) Be 5 5 4.
At least one pair of secondary tubercles or angular processes

present on the anterior margin of the head between the outer

tubercles and the anterior angles ; the posterior plate of the

prosternum more or less nelle sided, broadly truncate

behind (African forms) . ae : .. Solenocyclinae, p. 68.
( The clypeus exposed and separated from the frons ie a distinct
4 suture ; .. Pseudacathinae, p. 22.
The clypeus Hase with the fone aa re len m oo Le 5.

The outer tubercles obsolete or absent, the clypeus almost
always exposed’ Be Se da .. Proculinae, p. 32.

The outer tubercles distinct, the clvpeus rarely exposed .. Passalinae, p. 43.

of tubercles,* though these may be compound in structure ;

the antennae rarely with less than four well developed

lamellae ; never with both these characters as .. Macrolininae, p. 76.
The anterior margin of the head always with two pairs of

| The anterior margin of the head rarely with more than one pair

simple tubercles; the antennae always with three well
developed lamellae only .. 0 3 .. Leptaulacinae, p. 111.

Subfamily AULACOCYCLINAE.

Three of the genera defined in my previous account of this subfamily (1914c, pp. 192-3)
appear, in the light of further material, to be unnecessary. Two of these—Caulifer and
Auritulus—are monospecific, and may advantageously be merged in Aulacocyclus and
Cylindrocaulus, respectively. The third, Tristorthus, may also be merged in Aulacocyclus.
The best known species of Tristorthus is T. tricuspis, Kaup, from New Caledonia, a Species
shown by its short antennal lamellae to be most nearly allied to the Australian species
of Aulacocyclus. With this are associated firstly, two other New Caledonian species,
apparently distinguished largely by differences in size, and in my opinion doubtfully

1 The processes especially characteristic of the Solenocyclinae are situated immediately above the ventral tubercles,
Others may also, however, be present, and in all Malagasy forms a pair is more or less distinctly developed immediately
on the inner side of the fronto-vertical suture. The most primitive Malagasy genus, Malagasalus, lacks the former pair of
processes, and the above definition is applicable only by reason of its possession of the latter. The African and

Malagasy groups are composed, broadly speaking, of parallel series of genera separated largely by the presence or absence of
the latter pair of processes. I do not know of any African genus paralleling Malagasalus ; but if one exists the above
definition can hardly be expected to apply to it. See also above, pp. 10-11.

2 Except in one species of Vindex (below, pp. 43 & 47); imperfectly fused in one species of Chondrocephalus (below
pp. 43-45, fig. vi, 1). See also above p. 11.

3 The clypeus is completely hidden only in the genus Platyverres, though in the transitional species Verres corticola
it is hardly apparent. In both these species the inner tubercles are situated on the anterior margin of the head and may
readily be mistaken for outer tubercles, though a comparison with other species of Verres, and especially with V.
cavicollis, at once settles their true homology. The heads of the species in question are shown in fig v, p. 34.

4 Only in the genus Tarquinius, which has six well developed antennal lamellae.

14 Memoirs of the Indian Museum. (Vor. VER.

distinct ; secondly, a species which Zang (1905¢, p. 226) has shown not to be the Malaysian
species for which Kuwert took it, but probably also a form of fricuspis; and thirdly, a
species felderi, Stoliczka, from Amboina and Ceram, with a simply pointed central tubercle:
and long antennal leaflets like those of the Moluccan species of Aulacocyclus. The genus
Tristorthus is thus seen to consist only of a New Caledonian species (or group of species)
and a Moluccan species, allied respectively to different groups of the genus Aulacocyclus, in
which I therefore propose to place them.

The genera of Aulacocyclinae may now be redefined thus :—

The middle lower tooth fixed ; the aedagus composed of two

consecutive pieces, the tegmen and median lobe ;! the central

tubercle always present near the middle of the upper surface

of the head .. 50 of ae 58 3: Se 2.
1< The middle lower tooth moveable ; the aedagus composed of

three consecutive pieces, the tegmen being composed ofa basal

piece and lateral lobes ; the central tubercle absent or occupy-

ing the whole of the antero-median part of the upper surface

of the head .. = à = Le 52 ae 4.
The mentum with a strong median el a .. Comacupes, p. 14.
2} The mentum not keeled .. an af | ifs x 3.
The scars on the pronotum large, more or less ramified

or S-shaped; the central tubercle broad and low, never

pedunculate (Oriental forms only) .. 00 .. Taemocerus, p. 16.

The scars on the pronotum not very large, crescentic or circular ;

the central tubercle taller, usually pedunculate (mostly not

Oriental) a: 58 ate de .. Aulacocyclus, p. 17.

The central tubercle present, often very strongly elevated.
| apex fused with the anterior margin of the head; the a

separate aes fs ae a .. Ceracupes, p. 21

The central tubercle absent ; the elytra united .. .. Cylindrocaulus, p. 21.

Genus COMACUPES, Kaup, 1871, p. 17.
Type, Passalus cylindraceus, Perty, 1831, p. 36, fig. 3.

Comacupes cylindraceus (Perty).

Passalus cylindraceus, Perty, 1831, p. 36, fig. 3
Comacupes cylindraceus, Gravely, 1914c, p. 207, text-fig. 2, pl. xi, figs. 5-6a.

Specimens from the Malay Peninsula (Pahang), Singapore, North Nias (many from Hilr
Madjedja and one from G. Madjeja), Middle Nias (Dyma and Kalim Bungo), Sumatra
(Bedagei interior, ca. 600 ft.), Java and Borneo (Mt. Marapok). Mr. C. Holman-Hunt
has sent me for examination a specimen from Rawang in the Malay Peninsula. Length

24°5-27°5 mm.

1 These names for parts of the male genital tube are those adopted by Sharp and Muir (1912, pp. 481-483 and
484-485).

1918.] F. H. GRAvELY : Passalidae of the World. 15

Comacupes masoni, Stoliczka.
Comacupes masont, Stoliczka, 1873, pp. 151-2.
Comacupes masoni, Gravely, 1914c, p. 207, pl. xi, figs. 4-4a.

One specimen from an altitude of about 600 ft. in the interior of Bedagei, Sumatra,
and three without locality label. They all resemble the Sumatran specimen referred to in
footuote 2 of p. 269 of my “ Account of the Oriental Passalidae.” Without further
specimens from the Malay Peninsula, however, it is impossible to say whether the Malay
and Sumatran races of the species are distinct. Length 29-30 mm.

Comacupes stoliczkae, Gravely.
Comacupes stoliczkae, Gravely, 1914c, p. 206, pl. xi, figs. 3-3a.
Two specimens from Djember, Besoek, Java; two from Mt. Marapol- Borneo ; and two
without locality label. M. Babault has sent a specimen 24:5 mm. long from Medan,
Sumatra, for identification. Length 23:7-25'0 mm.

Comacupes basalis (Smith).
Passalus basalis, Smith, 1852, p. 18, pl. 1, fig. 5.
Five specimens from Mindoro and other parts of the Philippine Islands. Length
33°7-36°3 mm.
Comacupes cavicornis, Kaup.

Aulacocyclus cavicornis + laevicornis, Kaup, 1868a, p. 6.
Comacupes cavicornis, Kaup, 1871, p. 19.
Comacupes cavicornis, Gravely, 1914c, pp. 204-205, pl. xi, figs. 1-20.
Several specimens from Singapore, Java (Tjibodas, Telaga Bodas in Garoet, Preanger)
and Borneo (Pontianak). Length 24-5-26-0 mm.
These additional specimens show that the local races described in my ‘‘ Account of the
Oriental Passalidae ’’ (pp. 204-206) are not constant, and must be treated as one.

The species of Comacupes may be identified thus :—

The mesosternum strongly punctured all over; the abdominal
sterna with at least a few hair-bearing punctures in fresh

1 Specimens... ee bi 56 56
The mesosternum unpunctured except at the sides, the abdomen

unpunctured and hairless 50 56 .. C. fovercollis, Kuwert.
The lower margin of the overhanging portion of the central

tubercle short, or ascending obliquely to meet the upper

margin, which is always horizontal; the tubercle usually

truncate or concave anteriorly, not sharply pointed, narrower,

keeled or rounded above .. = 56 30 ae ac 3.
The lower margin of the overhanging portion of the central

tubercle usually rather long, always horizontal, the anterior

part of the upper margin descending obliquely to meet it in a

more or less acute angle ; the upper surface usually more or

x Jess distinctly grooved longitudinally, or excavate ae a ae 4,

Memoirs of the Indian Museum. [Vor. VII,

The central tubercle not strongly elevated, truncate or concave
in front, very variable

| The whole anterior end of the central tubercle raised well above

. cylindraceus, p. 14.

the supraorbital ridges, not truncate or concave in front C. mason, p. 15.
The central tubercle marked above with, at most, a more or less

4‘ distinct longitudinal groove

dE a 2 5% es 5.
The central tubercle strongly excavate above =

C. cavicornis, p. 15.

5 The central tubercle broad above, pedunculate

C. basalis, p. 15.
| The central tubercle narrow, not pedunculate

C. stoliczkae, p. 15.

senus TAENIOCERUS, Kaup, 1871, p. 20.
Type, Passalus bicanthatus, Percheron, 1841, pp. 41-2, pl. Ixxix. fig. 5.

Taeniocerus bicanthatus (Percheron).

Passalus bicanthatus, Percheron, 1841, pp. 41-42, pl. Ixxix, fig. 5.
Taeniocerus bicanthatus, Gravely, 1914c, pp. 208-209, pl. x1, figs. 7-75.

One specimen said, probably erroneously, to come from Ceylon ; others from Penang,
Singapore, Bintam Island, and Borneo (a number of specimens from Mt. Marapok in Dent
Province, northern spurs of Mt. Kina-Balu, Labuan and Pontianak). Mr. C. Holman-Hunt
has sent a specimen from Rawang, Malay Peninsula, for examination. Length 26-0-29:5 mm.

Taeniocerus platypus, Kaup.
Aulacocyclus platypus, Kaup, 1868a, p. 5.
Taeniocerus platypus, Kaup, 1871, p. 21.

Numerous specimens from Sumatra (Bedagei Interior, ca. 600 ft.), Java (Boeloe Lawang
in Pasoeroean) and Borneo (Mt. Kina-Balu).

M. Babault has presented us with specimens.
from Medan, Sumatra.

Length 16-3-19:5 mm.

Taeniocerus pygmaeus, Kaup.
Aulacocyclus pygmaeus, Kaup, 1868a, p. 5.
Taeniocerus pygmaeus, Kaup, 1871, p. 20.
Taeniocerus pygmaeus, Gravely, 1914c, pp. 209-210, pl. x1, figs. 8-84.

One specimen each from the Malay Peninsula and Sumatra (Bedagei Interior, ca. 600 ft.)
and several from Borneo (two from Mt. Marapok, Dent Province). Mr. C. Holman-Hunt

has sent specimens from Kuala Kansar, Malay Peninsula, for examination. Length 14-2-
15°9 mm.

Taeniocerus bicuspis, Kaup.
Aulacocyclus bicuspis, Kaup, 1858a, ». 5.
Taeniocerus bicuspis, Kaup, 1871. pp. 21-22.
Taeniocerus bicuspis, Gravely, 1914e, pp. 210-211, pl. xi, figs. 9-9a.

Four specimens trom Tukvat, Darjiling District, and one from “ India.” Length 18-5—
21-1 mm.

1918.] F. H. GRAVELY : Passalidae oj the World. 17

The species of Taeniocerus may be identified thus :—
The upper surface of the central tubercle about twice as long as
broad, flat, punctured, bordered by a very distinct horse-shoe-
shaped ridge which is open in front .. ae .. T. bicanthatus, p. 15.
1< The upper surface of the central tubercle relatively broader as a
rule, less flattened, unpunctured. the marginal ridge often in-
distinct or absent across the middle-line behind as well as in
front fe 22 pe se A at 3a DR
( The anterior tibiae very broad Ms oc -. IL. platypus, p. 16.

2 : ne
=] The anterior tibiae slenderer

The anterior margin of the canthus meeting the side of the head
at a considerable distance behind the anterior angle; the
5 external angle of the canthus obtuse = -. T. pygmaeus, p. 16.
The anterior margin of the canthus meeting the side of the head
| at a very short distance behind the anterior angle ; the
external angle of the canthus sharper a -. T. bicuspis, p. 16.

Genus AULACOCYCLUS, Kaup. 18684, p. 4.
Incl. Taemocerus [part] +Caulifer, Kaup, 1871 + Tristorthus, Kuwert. 1806, p. 220.

Type. Passalus edentulus, MacLeay, . 826, p. 430.

Aulacocyclus glabriusculus, Kuwert.
Aulacocyclus glabriusculus, Kuwert, 1897, pp. 280 and 282.

A number of specimens from Aru Island. Length 25:4-27-3 mm.

The antennal lamellae are long and slender; the central tubercle, which closely resem-
bles that of A. edentulus (fig. i, 7), is erect at base, oblique or almost horizontal above,
where it is medially grooved, the apex scarcely or not at all bent downwards. The elytral
grooves are shallow, and are scarcely punctured either above or below.

Aulacocyelus perlatus, Kaup.
Fig. IIL, 1, p. 18.
Aulacocyclus perlatus, Kaup, 1868a, p. 7.

Four specimens from New Guinea, three of them being from Stephansort, Astrolabe Bay.
Length 19:5-21'6 mm.

The antennal lamellae are long and slender, as in the preceding species. The central
tubercle is decumbent, and more or less distinctly bent downwards at the apex. All the
grooves of the elytra are strongly punctured.

Aulacocyclus parryi, Kaup.
Aulacocyclus parryi, Kaup, 1868a, p. 8.
Represented by specimens from Ceram, Halmaheira (Dodinga), Ternate and Singapore.
Length 23-2-25°3 mm.
The antennae and elytra resemble those of the preceding species. The central tubercle
resembles that of A. glabriusculus and A. edentulus, but is melined to be slightly thickened
on the under side just at the apex, giving it a somewhat more hooked appearance.

18 Memoırs of the Indian Museum. [ Vor. :VII,

Aulacocyelus aruensis, Kuwert.
Fig. III, 2.
Aulacocyclus aruensis, Kuwert, 1897, p. 282.

Two specimens from each of the following localities :—New Guinea, Jobi, Aru, Ceram.
Length 21-25 mm.

The antennae and elytra resemble those of the two preceding species. The central
tubercle somewhat resembles that of A. deyrollei, but is stouter and less elevated in front,
and has the apex perhaps a little more distinctly overhanging than is usual in that species.

Aulacocyclus felderi (Stoliczka).

tier UL, Sh,
Comacupes felderi, Stoliczka, 1873, p. 152, footnote.
Comacupes felderi, Arrow, 1907, p. 447.

One specimen from Amboina, and one from Honitetoe, West Seran! (? Ceram). Length
21 mm.

This species ditfers from all others that are known in having the central tubercle
ungrooved, erect, laterally compressed and simply pointed. The antennal lamellae are long

Se
A oe

Fie. III.
The central tubercle and supraorbital ridge of species of Aulacocyclus, from the left side x 4.
1. A. perlatus, Kaup. 5. A. tricuspis, Kaup.
2. A. aruensis, Kuwert. 6. A. erruns, Blackburn.
3. A. felderi (Stoliezka). 7. A. edentulus (MacLeay).
4. A. mastersi, MacLeay. 8. A. teres (Percheron).

and slender. The elytral grooves are scarcely punctured either above or at the sides, but
are decidedly stronger than in A. glabriusculus.

Aulacocyclus mastersi, MacLeay.
Fig. III, 4.
Aulacocyclus mastersi, MacLeay, 1871, p. 174.
Taeniocerus mastersi, Kuwerc, 1897, p. 275.

Several specimens from Queensland (Port Denison and Clarence River), N. S. Wales
(Richmond River) and Victoria. Length 21:7-27-5 mm.

The antennal lamellae are somewhat stout but moderately long. The central tubercle is
laterally compressed and comparatively low. It is somewhat variable, and may be scarcely
more elevated than in T'aeniocerus bicuspis ; usually, however, it is somewhat higher and
the apex may even be faintly overhanging. The fine puncturing on the head is usually in-
conspicuous or absent. All the elytral grooves, except perhaps the first pair, are very
tinely punctured ; none of them are very deep.

* = Exp. Martin III 92.” Concerning the distribution of this species see Arrow, 1907, p. 447.

1918.] F. H. GRaveLy : Passalidae of the World. 19

Aulacocyclus deyrollei, Kaup.

Aulacocyclus deyrollei, Kaup, 18684, pp. 7-8.
Taeniocerus deyrollei, Kuwert, 1897, p. 275.

One specimen 25:5 mm. long, in the Indian Museum collection. I have also examined
specimens in the British Museum.

A. deyrollei differs from A. masters: only in the much greater breadth of the central
tubercle and in the more conspicuous puncturing of the anterior part of the head.

Aulacocycius tricuspis, Kaup.
Fig. IIL, 5.
Aulacocyclus tricuspis, Kaup, 1868a, p. 7.

Several specimens from New Caledonia and one from Woodlark Island. Length 22-2 5
mm.

This species is easily recognised by the tridentate form of the central tubercle. which
resembles that of Comapcupes cavicornis, except that it is always broadest between the
paired dorsal denticles. The antennal lamellae are much shorter than in A. deyrollei. The
punctures in the grooves of the elytra are almost uniformly coarse.

Aulacocyclus errans, Blackburn.
Fic. III, 6.
Aulacocyclus collaris, Blackburn, 1896, p. 233.

Both sexes of this species appear to occur in two forms distinguishable from each other
only by their size.

Large form.—Several specimens from Queensland. Mr. H. Schroder has presented us
with specimens from the New England District of New South Wales. Length 27-30 mm.

Small form.—Numerous specimens from Queensland, including four from Cooktown,
three from the Melvor River and one from Cardwell. Length 19:6-23-0 mm.

The antennal lamellae are somewhat shorter than in A. deyrollei. The central tubercle
rises almost vertically and is then bent over forwards, the antero-ventral surface gradually
and the postero-dorsal abruptly, the dorsal surface being straight and more or less
horizontal. The anterior enlargements of the pronotal marginal grooves are somewhat
more pronounced than in other species of the genus. The grooves of the elytra are
even more coarsely punctured than in A. tricuspis.

Aulacocyelus edentulus (MacLeay).
Fig. III, 7.
Passalus edentulus, MacLeay, 1826, p. 439.

A number of specimens from Queensland (Brisbane), New South Wales (Sidney) and
Victoria. Length 23-30 mm.

The central tubercle is more strongly elevated than in :{. errans and less abruptly bent
forwards. The anterior enlargements of the pronotal marginal groove are not very
pronounced. The elytral grooves are less strongly punctured, especially dorsally.

Aulacocyclus teres (Percheron).
Fig. III, 8.
Passalus teres, Percheron, 1841, pp. 39-40.
A number of specimens from Queensland, New South Wales and Victoria (Melbourne).
Length 30'8-40:0 mm.
D 2

20 Memovrs of the Indian Museum. (Wer, WUE,

A. teres is much larger than A. edentulus ; its central tubercle is even more elevated,
and is somewhat more abruptly bent forwards—scarcely as abruptly, however, as in A.
collaris. The elytral grooves are very shallow and are obscurely punctured at the sides only.

The above-mentioned species of Aulucocyclus may be separated thus :—
The antennal lamellae long and slender ; ee and Moluccan
species ee ue : ; oc er 2.
1< The antennal lamellae shorter, at most note long a

| slender ; Australian and New Caledonian species (? also from

Woodlark island) de es Br a an 6.

| The central tubercle grooved above, its apex more or less
9 overhanging .. 38 a ae ; 5% aM 3.
The central tubercle not grooved above, its apex erect. es

compressed and simply pointed 56 se PAT Ciden pale
The grooves of the elytra shallow and scarcely punctured .. A. glabriusculus, p. 17.
The grooves of the elytra deeper and more or less strongly
punctured... I De Re 53 ae a 4.
The central tubercle depressed, with slender overhanging apex A. perlatus, p. 17.
42 The central tubercle less depressed, with stouter or scarcely

overhanging apex # de A a de an 5.

or less hook-like si os 21. .. A. parryr, p. 17
The apex of the central tubercle short and stout, scarcely
overhanging .. 3E a ats .. A. aruensıs, D. 18.

The central tubercle very short, its apex scarcely or not over-

6) hanging : oe “ie oo
The central tubercle more strongly elevated, with overhanging
apex Be a sie ee a Ba an 8.

The central tubercle laterally compressed ; the anterior part of

the head not conspicuously punctured ti A. mastersi, p. 18.
| The central tubercle very broad ; the anterior part of the ren

somewhat strongly punctured ae Be .. A. deyroller, p. 19.

)
The apex of the central tubercle well developed, usually more

( The central tubercle pointed at apex as seen from above, with
8 a pair of small tubercles bes:de the mid-dorsal groove behind A. tricuspis, p. 19.

The central tubercle normal, bifid at apex as seen from above ahs ne 9.

The central tubercle moderately elevated and somewhat abruptly
| bent forwards ; the elytral grooves very coarsely punctured .. A. errans, p. 19.
9) The central tubercle more strongly elevated and less abruptly
| bent forwards ; the elytral grooves not very strongly punctured 50 are 10.

( The central tubercle gradua'ly curved forwards; the elytral
aK erooves strongly impressed and distinctly punctured .. A. edentulus, p. 19.
| The central tubercle somewhat abruptly bent forwards ; the

| elytral grooves shallow and indistinctly punctured do Zale IAB; JD. 19

1918.] F. H. GRAVELY : Passalidae of the World. 2]
Genus CERACUPES, Kaup, 1871, p. 16.
Type, Passalus fronticornis, Westwood, 1842, pp. 124-125.

Ceracupes fronticornis (Westwood).

Passalus fronticornis, Westwood, 1842, pp. 124-125.
Ceracupes fronticornis, Gravely, 1914c, p. 212, pl. xi, fig. 12.

Three specimens from 5,000-7.000 ft., Ruby Mines District of Upper Burma ; two from
Renong, Siam, and several without recorded locality. M. Vitalis de Salvaza has sent for
examination a specimen from ca. 4,000 ft., Chapa, near Lao Kay, Ht. Tonkin ; and Mr. J.
Coggin Brown has presented us with a specimen from Loi Tawng Kyaw, Tawng Peng State,
N. Shan States, Upper Burma, 5,500-7,000 ft.; and one from Man Pat, Mongmit State,
5.200 ft., Ruby Mines District, Upper Burma.

Length 20-23 mm.

This species is less distinct from C. austeni and C. arrowi than I supposed when drawing
up the key to the species (1914c, p. 319), the shape of the central tubercle being very
variable. The distinction given in that key between C. austen: and the present species
appears to hold good, though the distal bifurcation of the central tubercle of the latter is
sometimes very weak. The most distinctive difference between C. arrow: and the present
‚species is found in the elytra, whose grooves are deeply impressed and coarsely punctured
in the latter, but shallow and less strongly punctured in the former.

The species of Ceracupes may be identified thus :—

The grooves of the elytra, and their punctures, somewhat
Î shallow RR aa a: ae .. CO. arrowı, Heller.

The grooves of the elytra, and their punctures, very deep
The apex of the horn formed by the fusion of the central
tubercle with the anterior maraın of the head more or less
distinetly bifid ae Le 56 .. | Cs fronticornis, p. 21.

The apex of this horn acute 5 as .. OC. austen, Stoliczka.

Genus CYLINDROCAULUS, Fairmaire, 1880, p. 164.
Incl. Auritulus, Zang, 1905.

Type, Cylindrocaulus bucerus, Fairmaire, 1880, p. 164.

The species of this genus may be identified thus :—

The front coxae almost contiguous; the canthus extending

about half way across the eye; the supra-orbital tubercles

flattened, expanded at the apex, truncate 36 .. C. paialis (Lewis).
The front coxae widely separated ; the canthus extending sall

the way across the eye ; the supra-orbital tubercles slender

| andpointed .. te > 58 .. ©. bucerus, Fairmaire.

22 Memoirs of the Indian Museum. [Vou. VII,
Subfamily PSEUDACANTHINAE.

The most extensive genus of this subfamily, and one of the most primitive, appears to.
be Popilius, a genus which, as limited by previous authors, has proved most difficult to define
satisfactorily. None of the characters hitherto used for this purpose appear really to have
more than specific value. Especially variable and untrustworthy, in many cases even as
a specific character, is the shape and size of the central tubercle. Some of the most definite
characters are found in the clypeus, but even these do not as a rule afford satisfactory
generic distinctions. It seems necessary, therefore, to smk the names Heliscus (=Soranus),
Odontotaenius (= Passalus, auct. nec Fabricius) and Passalotaenius as synonyms of Popilius.
Similarly Conger, Rimor and Rimoricus may be sunk as synonyms of Oileus. And
both Popilius and Orleus must be redefined.

The plastic and primitive genus Popilius forms a starting point to which the remaining
genera of American Passalidae may be traced back, the more primitive species of all the
remaining subfamilies having the clypeus similarly exposed, although there is usually no
clypeofrontal suture, and the more highly specialized having it hidden.

In the subfamily Pseudacanthinae itself three separate lines of evolution may be
recognized. In one, which includes a new genus Ovrleoides, and the genera Oùleus and
Undulifer, the sides of the metasternum are broadly hairy. In another, which is represented
only by the genus Spurius, the central tubercle is absent. In the third the elytra are
united and the wings are of use for stridulation but not for flight.

The more primitive members of the third group have the three terminal teeth of the
mandibles distinct as in Popilius. I have only seen three such species ; these belong to
the genera Pseudacanthus, Triaenurgus and Nasoproculus, in none of which are the sides of
the elytra hairy. It will be convenient to unite these three genera, and with them should
probably go the genera Ogyges, Prosochtus and Truquius, which have hairless elytra —
unfortunately their mandibles have not been described. Petrejoides should perhaps come
here also, but Kuwert’s definition is inconclusive, and I have nothing else to go by. The
name may equally well be synonymous with Proculejus or even with Proculejoides.

The genera Proculejus,! Prosochtus, and Eriopterus should likewise, in all probability,
be united into one genus Proculejus, differing from Pseudacanthus in having only two teeth
at the apex of the mandible instead of three and in having hairy sided elytra. The
reduction in the number of teeth on the mandible probably takes place by the union
of the two lowest terminal teeth. The anterior lower tooth appears broad and bidentate in
this genus on both mandibles, instead of bidentate on the left and unidentate on the right as.
in allied forms of the preceding genus.

The genera of Pseudacanthinae may be defined thus :—

The elytra separate, their vertical anterior part hghtly concave = ye 2.

1) The elytra united along the middle line, their vertical anterior
part lightly convex at Bie = 54 oe oe 6.

1Except P. quitensis, Kaup (see below p. 51).

1918.1 F. H. GRAVELY : Passalidae of the World. 23

The sides of the metasternum broadly hairy and punctured
throughout .. : ie 2 3 Bh ae 3.

24 The hair-bearing punctures of the metasternum confined to the

lateral areas and extreme anterior parts of the anterior

intermediate areas

3 | The clypeus transversely trapezoidaı . a .. Otleoides, p. 23.
| The clypeus transversely linear, straight or undulating ao = Na 4.
( The clypeus straight or lightly concave in the middle line .. Oileus, p. 25.
4 The elypeus strongly convex in the middle and on either side,
strongly produced backwards at the two points uniting the
three curves thus formed .. ae ER .. Undulifer, p. 25.
Al The central tubercle absent ae Mr -. Spurvus, p. 26.
4 The central tubercle present Popilius, 5. 26.

The mandibles tridentate at apex, Ae left one w ith a Parle
bifid tooth between the apex and the movable tooth, the
right one with a unidentate tooth in this position (fie. iv, 13,
6 p. 24) ; the sides of the elytra hairless 3 Pseudacanthus, p. 30.
The mandibles bidentate at apex, both of them with a broadly
bifid tooth between the apex and the moveable tooth (fie. iv,
16. p. 24); the sides of the elytra hairv 3 -. Proculejus, p. 3l.

Genus OILEOIDES, n. gen.

Metasternum with a broad band of hair-bearing punctures on each side; clypeus
expanded as in the more primitive species of Popilius. Otherwise like Popilius.

Type, O. parvicornis, n. sp.

Oileoides parvicornis, n. sp.
Rio IV, tp

Two specimens from the Cauca Valley, Columbia. Length 27-28-5 mm.

The antennae have moderately long lamellae. The labrum is punctured and hairy, with
distinctly concave anterior margin and convex sides, the latter somewhat convergent
behind. Both mandibles have well developed upper, terminal and anterior lower teeth.
The last-mentioned tooth is simple on the right side ; on the ieft it is double, sometimes
with the anterior of the two parts into which it is (vertically) divided itself divided to a
less extent horizontally. The mentum is smooth in the middle, hairy and punctured
laterally ; its scars are not very strongly marked. The clypeus is extensive and flat like
that of Spurius bicornis and dichotomus ; itis lightly procurved as in the latter species.
The frontal ridges are obsolete, and the central tubercle, though situated on the crest of
a straight transverse ungrooved ridge formed by the union of the two parietal ridges, is
very small. Between the central tubercle and the obsolete frontal ridges is a triangular
area which is somewhat dull and strongly roughened, all the rest of the surtace of the head

‘being smooth and glossy.

24 Memoirs of the Indian Museum. (Vor. VII

The pronotum is transverse. The anterior margin is practically straight, the sides
lightly and the posterior margin more distinctly convex. The scars are a little uneven, but
are not definitely punctured. The prosternum is pointed behind.

The scutellum is smooth or distinctly but sparsely punctured. In the specimen in which
the scutellum is unpunctured there is a strongly marked median groove; in the other this
groove is much broken. The mesothoracic episterna are glossy and coarsely punctured all
over, except in the posterior angles, which are matt and have less distinct or no punctures
but may be more or less rugose. The mesosternum is more or less smooth and glossy all
over—less so at the sides than in the middle.

Nae
Seo
Vs
N
Tic. IV.
Pseudacanthinæ ; specific characters in the upper surface of the head x 4.
1. Otleoides parvicornis, Gravely. 9. Popilvus guatemalae, Gravely.
2. Oileus ridiculus (Kuwert). 10. Popilius tropicus (Percheron).
3. Unduhfer incisus (Truqui). 11. Popilius brevioripennis (Kuwert).
4. Spurius dichotomus, Zang. 12. Pseudacanthus solidus (Arrow)
5. Popilius recticornis (Burmeister). 13. Psexdacanthus bifidus (Zang).
6. Popilius marginatus (Percheron). 14. Pseudacanthus jalapensis, Bates.
7. Popilius amazonicus, Gravely. 15. Proculejus sartori, Kaup.
8. Popilius intergeneus (Bates). 16. Proculejus pubicostis, Bates.

The lateral areas of the metasternum are narrow. These, the anterior intermediate
areas, and the outer margins of the posterior intermediate areas are punctured and hairy.
The rest of the plate is smooth, hairless and glossy, as are also the posterior coxae and
the abdominal sterna. The grooves of the elytra are somewhat strongly punctured,
especially laterally, but the punctures are small and round. The middle and hind tibiae
are without spines except at the apex.

Oileoides subrecticornis (Kuwert).
Soranus subrecticornis, Kuwert, 1897, p. 296.
Three specimens from the Cauca Valley, Columbia. Length 22-24 mm.
O. subrecticornis differs from O. parvicornis in its somewhat smaller size, its shorter
antennal lamellae, its straighter clypeus, its somewhat more strongly developed central

1918.] F. H. GRAveLy : Passalidae of the World. 25

tuberele and frontal ridges, which may be united and never have a roughened area between
them, and its prothorax which is strongly punctured in and close round the scars.

Genus OILEUS, Kaup, 1869, p. 3.
Incl. Coniger, Zang, 1905 ce; Rimor, Kaup, 1871 ; Rimoricus, Kuwert, 1897.
Type, Passalus rimator, Truqui, 1857, p. 266 (see Arrow, 1907, pp. 447-449.)
Species of this genus appear to be much larger than are those of the last, and further
have always, so far as is known, a large decumbent central tubercle with free apex, instead
of a feebly developed one, as well as a linear instead of an expanded clypeus.

Oileus ridiculus (Kuwert).
ie, JOY, Ae
Rimoricus ridiculus, Kuwert, 1897, p. 287.

Three specimens from Guatemala. Length 34-35 mm.

The lamellae of the antennae are extremely short, being equal to not more than two of
the immediately preceding joints in length; the pronotum is punctured in and around
the scars ; the scutellum is roughened all over with obsolete punctures; the mesosternal
scars are hairy; and the grooves of the elytra are very finely punctured. In one specimen

the mesosternum is coarsely punctured.

Oileus sargi (Kaup).
Rimor sargii, Kaup, 1871, pp. 119-120.
One specimen from Guatemala. Length 30 mm.
The antennal lamellae are very long, being as long as about four of the immediately
preceding joints ; the pronotum is unpunctured ; the scutellum is strongly punctured in
the middle ; and the mesosternal scars are not hairy.

Oileus rimator (Truqui).
Passalus rimator, Truqui, 1857, p. 266.

One specimen from Omilteme, Guerrero, 8,000 ft., lent by the British Museum. Length
378 mm.

This species is closely allied to O. sargi, from which it differs in having the clypeus
lightly biconvex, the anterior margin of the pronotum biconcave and the mesosternal scars
hairy. |

Genus UNDULIFER, Kaup, 1869, p. 6.

Type, Passalus incisus, Truqui, 1857, pp. 266-267.

Undulifer incisus (Truqui).
Fig. IV, 8.
Passalus incisus, Truqui, 1857, pp. 266-267.
One specimen from Cordova, Mexico, presented by the British Museum. Length 20 mm.
Undulifer incisus is closely related to Oileus ridiculus and sargi. The antennae of the

single specimen before me are broken, but from the one remaining lamella I conclude that
E

26 Memoırs of the Indian Museum. [Vor. VII,

the lamellae were very long and slender as in the Jatter of these species. The clypeus has the
form characteristic of the genus. The central tubercle is narrower and less separated from
the surface of the head than in O. ridiculus and O. sargi. The pronotum is more extensively
punctured round the scars than in either of these species. The mesothoracic episterna are
polished throughout. The meso- and metasterna are extensively covered with hair-bearing
punctures, especially laterally. The grooves of the elytra are quite as strongly punctured
as in O. ridiculus.

Genus SPURIUS, Kaup, 1871, p. 75.
Type, Passalus bicornis, Truqui, 1857, p. 317.

| Spurius bicornis (Truqui).
Passalus bicornis, Truqui, 1857, p. 317.

One specimen from Mexico and two from Guatemala. Length 17°5-18°0 mm.

Spurius dichotomus, Zang.
Fig. IV, 4, p. 24.
Spurius dichotomus Zang, 1905a, pp. 227-229.

One specimen from Guatemala, and one unlabelled preparation of the head and
appendages. Length 21 mm.

This species may readily be distinguished from the preceding by its somewhat larger
size ; by the smaller size of the conical processes representing the parietal ridges, which are
oblique instead of transverse ; and by the shape of the clypeus, which is procurved instead of
straight.

Genus POPILIUS, Kaup, 1871, p. 75.

Incl. Heliscus, Zang, 1905 (=Soranus, Kaup, 1871, preoccupied) ; Odontotaenius, Kuwert,
1896 (=Passalus, auct. nec Fabricius, see Zang, 19056, pp. 224-225); Passalotaenius,
Kuwert, 1896.

Type, Passalus marginatus, Percheron, 1835, pp. 89-90, pl. vu, fig. 1.

Popilius recticornis (Burmeister).
Fig. IV, 5, p. 24.
Passalus recticornis, Burmeister, 1847, pp. 508-509.

Four specimens from Mexico. Length 18-20 mm.

P. recticornis is the smallest species of its genus known to me. The antennal lamellae
are short. The clypeus is strongly procurved. The frontal ridges are obsolete, but the
central tubercle, which is directed forward, is very strongly developed, the apex being free.
Apart from some of these characters, and the smoothness of the metasternum
characteristic of the genus, P. recticornis closely resembles the above described species of
the genus Ouileoides. The mesothoracic episterna are, however, less extensively punctured
and uniformly glossy, and the prosternum is truncate behind. The punctures round the
scars on the pronotum are usually more numerous even than in O. subrecticornis. |

1918.] F. H. Gravety: Passalidae of the World. 27

Popilius marginatus (Percheron).
Fig. IV, 6, p. 24.
Passalus marginatus, Percheron, 1835, pp. 89-90, pl. vu, fig. 1.

Three specimens from Farinas, Bolivia. Length 21-22 mm.

Popilius marginatus is of about the same size as Oileoides subrecticornis or a little
smaller, but it has longer antennal lamellae, its straight clypeus is less extensive
antero-posteriorly and consequently less flat, there is a strong median keel or pair of keels
between the small central tubercle and the obsolete frontal ridges, the pronotum is more
thickly punctured above the scars and its median groove is complete, the scutellum
is strongly punctured at least near the middle line in front. The metasternum resembles
that of P. recticornis and other species of Popilius ; it may have a few punctures in the
posterior intermediate areas. The abdominal sterna are punctured in the angles of the
scars. In other respects P. marginatus resembles O. subrecticornis.

Popilius amazonicus, n. sp.
Hi MINE Wa py 24

One specimen from the Amazon, Peru. Length 20 mm.

P. amazonicus resembles P. marginatus in general appearance, but the tubercles and
ridges of the head resemble rather those of P. intergeneus, the pronotum is very sparsely
punctured near the scars, the pointed posterior extremity of the prosternum is very slender,
and the scars of the abdominal sterna are more extensively punctured.

Popilius intergeneus (Bates).
Fig. IV, 8, p. 24.
Soranus (?) intergeneus, Bates, 1886, p. 21.

Numerous specimens from Guatemala. I am indebted to Mr. Arrow for their identi-
fication by comparison with Bates’ type. Length 20‘0-21°5 mm.

This species is intermediate in size between the two preceding. From P. marginatus
it differs in having the antennal lamellae short, the anterior margin of the clypeus slightly
convex and often indented in the middle line, the frontal ridges stronger and on an average
more directly united with the central tubercle, the pronotum punctured in the anterior
angles as well as above the scars, the prosternum truncate instead of pointed behind, the
mesothoracic episterna uniformly glossy except for an unpunctured matt oval patch a little
below and in front of the unpunctured posterior angles, the scars of the abdominal sterna
more extensively punctured and the grooves of the elytra more strongly punctured.

Popilius guatemalae n. sp.
Hig. IV, 9, p. 24.

One specimen from Guatemala. Length 28'353 mm.

This species differs from P. intergeneus in its much larger size, straight margined
clypeus, smaller and more arcuate frontal ridges, grooved parietal ridges, incomplete
pronotal median groove, much less extensively punctured pronotum (the punctures being
confined to the neighbourhood of the scars), pointed posterior end of prosternum, almost

28 Memoirs of the Indian Museum. (Vor. VII,

unpunctured scutellum (the punctures being confined to a pair of longitudinal lines), normal
‘mesothoracic episterna with matt and indistinctly punctured posterior angles, smoother
abdominal sterna, and more finely punctured elytral grooves.

Popilius tropicus (Percheron).
Fig. IV, 10, p. 24.
Passalus tropicus, Percheron, 1835, pp. 97-99, pl. vii, fig. 4.

Five specimens from Mexico, and one without locality record. Length 25(4)-28(9) mm.

Kaup recognizes three varieties of this species, based on the sculpture of the upper
‚surface of the head, and quotes Truqui to the effect that the central tubercle is very variable.
One of the specimens before me has a much more strongly developed central tubercle than
‘the others. It is a female and all the others are males!.

The clypeus appears always to be distinctly recurved, and I doubt whether the frontal
ridges, which are often absent, are ever very strongly developed. They are straighter than
in P. guatemalae. The parietal ridges are not grooved, and the scutellum may be irregularly
punctured. In other respects this species resembles P. guatemalae-

Popilius brevioripennis (Kuwert).
Fig. IV, 11, p. 24.
Odontotcenius brevioripennis, Kuwert, 1897, p. 290.

One specimen from Mexico, distinctly smaller than the type (23 instead of 26 mm.
long) and with numerous punctures above the scars. The number of these punctures is,
however, very variable in the closely allied species, P. striatopunctatus, and the difference in
size is within the usual limits of variability. The specimen of P. brevioripennis differs from
the single female specimen of P. tropicus before me in having longer antennal lamellae,
a strong median convexity of the anterior margin of the clypeus, a slightly trunctate
posterior prosternal plate, mesothoracic episterna like those of P. intergeneus, and more
strongly punctured elytral grooves.

Popilius striato-punctatus (Percheron).

Passalus striato-punctatus, Percheron, 1835, pp. 101-102, pl. vi, fig.

Numerous specimens from Mexico, one from British Honduras and two from Nicaragua.
‘Length 25-28 mm.

The size of the central tubercle is variable ; it is smaller in males than in females, but
the difference is not a sharp one. P. striatopunctatus differs from T. brevioripennis in
its larger size, its more strongly developed central tubercle and slightly less indistinet
frontal ridges. The posterior end of the prosternum is variable.

Popilius cornutus (Fabricius).
Passalus cornutus, Fabricius, 1801, p. 256.

Numerous specimens from Florida, two from Mexico and one from each of the following
places :—New York, Illinois, Delaware (Wilmington), Ohio and Haiti. Length 29-37 mm.

1 In one the genitalia were found to be damaged, and I was unable to determine the sex with certainty-

1918.] F. H. Gravety: Passalidae of the World. 2s

The central tubercle is very variable in shape and size, and is larger in females than in
males. P. cornutus differs from P. striatopunctatus in its larger size, somewhat shorter
antennal lamellae, evenly arched anterior margin of clypeus, pedunculate and larger
central tubercie, unpunctured pronotum, prosternum usually more broadly truncate behind.
mesothoracic episterna with a broad matt band parallel to but not in contact with the
lower margin, unpunctured posterior intermediate areas of metasternum, and more finely
punctured elytral grooves.

The above species of Popilius may be distinguished from one another thus :—

( The clypeus more or less broadly flattened and trapezoidal

1< The clypeus transversely linear or distinctly prominent in the

=~!

middle line

À The central tubercle elongate, with free forwardly directed apex P. recticornis, p. 26.
The central tubercle small, its apex not free sis Klo 5% IN 3.

/ The antennal lamellae long, equal to about four of the

5 immediately preceding joints in length as a “he = 4.
The antennal lamellae short, equal to about two of the
immediately preceding joints in length is st oe sé 5,

The frontal ridges short and curved, together forming almost
a semi-circle .. La sé 00 .. P. marginatus, p. 27.
4 : ‘ - RE:
The frontal ridges long and approximately straight, meeting in

a slightly obtuse angle .. Ae + .. P. amazonıcus, p. 27.

The central tubercle situated in, or a little in front of, the
middle of the head; the frontal ridges well developed, long
and approximately straight, meeting in an obtuse angle .. P. intergeneus, p. 27.
The central tubercle situated a little behind the middle of the
head ; the frontal ridges weaker or absent 52 wis 5% a 6.

[ The clypeus straight ; the frontal ridges forming a semi-circle in
| front of the central tubercle ; the parietal ridges grooved .. P. quatemalae. p. 27.
6) The clypeus lightly recurved; the frontal ridges absent, or
extending straight to the obsolete frontal tubercles from
either side of the base of the central tubercle at about a
| right-angle to each other ; the parietal ridges keeled
The clypeus linear, parallel-sided, lightly recurved ; the central + P. tropicus, p. 28.
tubercle sometimes with free apex, but never pedunculate
| The clypeus with at least the posterior margin straight; or
\ the central tubercle pedunculate : 5% 50 Da 0% 8.
{ The anterior margin of the clypeus with a strongly developed
median convexity, the central tubercle not pedunculate ra Ye Be 9.
8) The anterior margin of the clypeus at most lightly convex
medially, the central tubercle pedunculate so .. P. cornutus, p. 28.
( The central tubercle of moderate size, its apex scarcely free .. P. brevioripennis, p- 28.
9+ The central tubercle large in both sexes (especially the female),
( its apex entirely free CE ai oo .. P. striato-punctatus, p. 28.

30 Memoirs of the Indian Museum. (Wom, Wile

Genus PSEUDACANTHUS, Kaup, 1869, p. 9.

Incl. Nasoproculus, Zang, 1905¢.;% Ogyges, Kaup, 1871 ;? Petrejoides, Kuwert, 1806 :
Triaenurgus, Bates, 1886; ? Truquius, Bates, 1886.

Type, Passalus mexicanus Truqui, 1857, pp. 315-316.

Pseudacanthus bifidus (Zang).
Fig. IV, 13, p. 24.
Nasoproculus bifidus, Zang, 1905c, p. 232.

Six specimens from Mexico. Length 37-41 mm.

The antennal lamellae are long, being equal to three or four of the immediately preced-
ing joints. The anterior margin of the labrum is deeply cleft ; the sides of the labrum are
straight and parallel. The clypeus is strongly arched on either side of a still stronger
median concavity. The central tubercle resembles that of the genus Ovleus in general form.
There are no frontal tubercles or ridges. The parietal ridges are obsolete. The pronotum
is unpunctured ; its median groove is very strong, but does not quite reach the anterior
margin. The scutellum is strongly punctured on either side of the middle line. The
mesothoracic episterna are punctured above and near the anterior margin only ; they are
glossy, except for a matt oval patch behind the lower margin and a matt streak in the
posterior angle. The mesosternum is glossy except in the scars, which are matt; it
is unpunctured. The metasternum is hairless, and is unpunctured except in the
posterior intermediate areas; the lateral areas are linear and smooth. The abdominal
sterna are smooth. The grooves on the elytra are finely punctured laterally.

Pseudacanthus solidus (Arrow).
Fig. IV, 12, p. 24.
Triaenurgus solidus, Arrow, 1907, pp. 452-453.

Described from a (damaged) cotype, 39 mm. long, from Chuipache, Quezaltenango,
Guatemala, lent by the British Museum.

The anterior margin of the labrum is lightly concave. The clypeus is straight, and bears
a pair of small tubercles, each situated about half way from the middle line to the tubercle
in which it terminates laterally. The parietal ridges, though somewhat small, are quite.
distinct. There are two or three punctures in or near the scars on each side of the
pronotum. The matt areas of the mesothoracic episterna are even more restricted than in
P. bifidus. The mesosternum is punctured and hairy laterally behind. The lateral areas of
the metasternum are broad ; both these areas, and the intermediate areas in front of and
behind the usual group of coarse hairless punctures on the posterior intermediate areas,
are covered with hair-bearing punctures. The abdominal sterna are smooth. In other
respects this species resembles P. bifidus.

Pseudacanthus jalapensis, Bates.
Fig. IV, 14, p. 24.
Pseudacanthus jalapensis, Bates, 1886, p. 9, pl. 1, figs. 10-10a.
One specimen from Oajaca, S. Mexico. Also a cotype from Jalapa lent by the British-
Museum. Length 22:0-23:5 mm.

1918.] F. H. GrAvELY : Passalidae of the World. 31

P. jalapensis is very much smaller than either of the preceding species of the genus,
and its antennal lamellae are much shorter in proportion than in those species. The clypeus
is very lightly concave (practically straight) and has no tubercles except at its lateral
extremities. A pair of small frontal tubercles is situated a little behind it, each about a third
-of the way from the end to the middle line. The frontal and parietal ridges are obsolete.
The central tubercle is very large and resembles in form that of the two preceding species.
The pronotum resembles that of P. solidus. The scutellum is strongly punctured near the
middle line, especially behind. The mesothoracic episterna have one or two matt areas close
to the lower margin, the rest of the surface being glossy ; they are punctured above and
along the anterior margin. The mesosternum is polished except along the lateral margins
where it is dull. The metasternum has hair-bearing punctures only beside the middle coxae ;
the lateral areas are linear and somewhat rough; the general surface is glossy and
unpunctured. The abdominal sterna are slightly roughened in the broad outermost parts of
‘the scars only. The elytra are somewhat coarsely punctured in the grooves.

Genus PROCULEJUS, Kaup 18685, p. 13.

Incl. ? Eriopterus, Kuwert, 1806 ; ? Prosochtus, Bates, 1886.

Type, Proculejus truquui, Kaup, 18685, pp. 16-17.

Proculejus pubicostis, Bates.
Fig. IV, 16, p. 24.
Proculejus pubicostis, Bates, 1886, p. 5, pl. 1, figs. 4-4a.

Two specimens from Mexico, 30-31 mm. long.

The antennal lamellae are moderately long, being about equal to three ofthe immediately
‚preceding joints in length. The labrum is parallel-sided, and is strongly concave in front.
The clypeus is very lightly convex (practically straight) ; it bears an obscure tubercle at
either end, and just behind and on the inner side of these tubercles is a pair of small frontal
tubercles, which the obsolete frontal ridges do not clearly reach. The central tubercle is very
large, it is massive and rounded at the base, but the long forwardly directed apex is
somewhat slender. The parietal ridges are obsolete. The pronotum bears only a few fine
hair-bearing punctures below the scars; the strongly impressed median groove scarcely
reaches the anterior margin. The scutellum is strongly punctured on either side of the
middle line, especially behind. The mesothoracic episterna are punctured above and in
front ; they are glossy except for an extensive matt band, which extends downwards from a
little above the posterior angle of each towards the ventral angle. The mesosternum is
polished except in the scars, which are deep and narrow. The metasternum resembles
that of Pseudacanthus jalapensis. The abdominal sterna are smooth. The elytra are
‘somewhat coarsely punctured above, very coarsely punctured at the sides; the last
three ribs above the margin are finely punctured and hairy.

32 Memoirs of the Indian Museum. [Vor. VII,

Proculejus sartori, Kaup.
Fig. IV, 15, p. 24.
Proculejus Sartor, Kaup, 18685, p. 17.

One specimen without locality record. Length 12:5 mm.

The labrum is concave in front, but not so strongly as in P. pubicostis. The clypeus is
straight or lightly concave, and is united with the well developed frontal tubercles a little
distance from each end. The frontal tubercles are united by well developed frontal ridges
to the central tubercle, which is situated a little in front of the middle of the head; this
tubercle is small and upwardly directed, and has no free apex. The scutellum is sparsely
and somewhat finely punctured, especially in front. The mesothoracie episterna are
punctured except in the posterior angles, and are glossy except for an oval patch behind the
lower margin. In all other respects this species resembles the last, except that the
mesosternal scars are shallower and that the elytral grooves are if anything even more
coarsely punctured.

Proculejus truquii, Kaup.
Proculejus Truquii, Kaup, 18686, pp. 16-17.

One specimen from Mexico, 31 mm. long.

Proculejus truquii is much broader in proportion to its length than is either of the two
preceding members of the genus. The head is ornamented much as in P. sartori, but the
frontal area is somewhat narrower and the central tubercle is situated more in the middle.
The pronotum resembles that of the two preceding species. The scutellum bears a few
obscure punctures. The grooves of the elytra are very much broader and more coarsely
punctured than in either of the preceding species, being about as wide as the ribs at the sides,
and about half as wide above, the punctures in each groove being much broader than the
spaces between them. In all other respects P. truquii resembles P. sartori.

Subfamily PROCULINAE.

Almost all members of this subfamily have the clypeus exposed as in the
Pseudacanthinae ; but it is fused with the frons instead of separated from it by a distinct
suture. Faint traces of this suture, as of the outer tubercles, may perhaps be recognized in
the genus Arrox, which is probably the most primitive genus of the family. From this genus
the extensive genus Veturius may easily be derived ; and this appears to have given rise to
two divergent lines of descent, culminating respectively in the flightless genera Platyverres
and Proculus. The former, in which the clypeus is completely hidden, is connected to
Veturius through the genus Verres, in which all stages of the disappearance of the clypeus
may be traced (compare fig. v, 11-15, p. 34). The latter, in which the specialized
features associated with flightless species! reach their highest development, has the clypeus
exposed, and is connected to Veturius by the genus Publius, a genus which only differs
from species of Veturius with the two lowest terminal teeth fused by its fused elytra and
reduced wings.

1 See above pp. 4-5.

1918.] F. H. GRAVELY : Passalidae of the World. 33-

The genus Procululus, Zang (19054, pp. 225-227, type, P. inca) is probably allied to
Publius and Proculus, though in no other genus of Proculinae are the lateral areas of the
metasternum known to be hairless—a character suggesting possible affinities with
Proculejoides, in the subfamily Passalinae. It resembles Publius in having normal
antennae, and Proculus in having egg-shaped elytra. The mandibles appear to resemble
those of the last named genus.

The genera of Proculinae known to me may be defined thus :—

| The elytra separate, their vertical anterior part lightly con-
1 cave oe ss ae 00 a 5.0 .. 2.
| The elytra united along the middle line, their vertical anterior

part more or less distinetly convex . ae a 56 coul 4.

from above ; the clypeus always exposed and more or less

horizontal; the anterior margin of the pronotum usually

biconcave a si a nr
The anterior lower tooth of the left mandible trıdentate as seen
trom above ; the clypeus steeper, or hidden and rudimentary ;

the anterior margin of the pronotum straight, or lightly

| The anterior lower tooth of the left mandible bidentate as seen

biconvex Le AF a a so GS, ip 20,

Rudimentary outer tubercles present ; the anterior margin of

the pronotum straight De vr os ALON, Dee
The outer tubercles absent; the anterior margin of the
pronotum more or less sinuous or with a stronely sinuous

marginal groove is ae 58 -. Veturius, p. 35.
The clypeus entirely hidden; the mandibles tridentate
distally = Br Ss .. Platyverres, p. 41.
| The clypeus broadly exposed ; the mandibles usually bidentate
distally = + de = = 38 Be 5.
The lamellae of the antennae of moderate length ; the anterior
lower tooth of the left mandible broader than that of the
right ; the sides of the elytra approximately straight and
parallel ; the metasternum with distinct lateral areas .. Publius, p.42.
The lamellae of the antennae abnormally long and slender.
the three together being narrower than any one of them is
long ; the mandibles symmetrical ; the sides of the elytra
strongly rounded ; the metasternum without distinct lateral

areas Proculus, p. 42.

Genus ARROX, Zang, 1905), p. 155.
—Sertorius, Kaup, 1871, preoccupied.

Type, Sertorius agassiz, Kaup, 1871, p. 114.

34 Memovrs oj the Indian Museum. [Vor.. VII,

Arrox agassizi (Kaup).
Fig. V, 1.
Sertorius Agassizi, Kaup, 1871, p. 114.

One specimen from Nicaragua, 30 mm. long. Both mandibles have three well developed
terminal teeth ; the anterior lower tooth of the right side is more or less columnar, that of
the left is broadly bidentate as seen from above, but the anterior denticle is itself divided
into two horizontally. The upper surface of the head though glossy is irregular, not smooth

TG ave

Proculinae ; specific characte s in the head and prothorax x 4.
c.A., clypeus angles ; I.T., inner tubercles ; 0.T., outer tubercles.

1. Arrox agassizi (Kaup). 9. Veturius boliviae, Gravely.

2. Publius crassus (Smith). 10. Veturius heydeni, Kaup.

3. Veturius sinuatosulcatus, Gravely. 11. Verres furcilabris (Eschscholtz).
4. Veturius spinifer, Gravely. 12. Verres sternipunctitus, Kuwert.
5. Veturius sinuatosulcatus Grav ly. 13. Verres cavicollis, Bates.

€. Veturvus unicornis, Gravely. 14. Verres cortecola (Truqui).

7. Veturius criniceps, Kuwert. 15. Platyverres intermedius (Kaup).
8. Veturius assimilis (Weber).

as in species of the allied genera Veturius and Verres ; and the clypeus is somewhat convex
above. On either side of the clypeus is a rudimentary outer tubercle, but there is no ridge
between these tubercles and the larger inner tubercles. The pronotum has a very few
coarse punctures only in the slightly enlarged anterior extremities of the narrow marginal
groove ; Its median groove scarcely reaches the anterior margin. The scutellum is thickly
punctured on either side of a smooth middle line. The mesothoracic episterna are glossy
and closely covered with strong coarse punctures above, matt and more faintly and sparsely

oe
Qt

1918. F. H. GRAVELY : Passalidae of the World.

punctured and hairy below. The mesosternum is dull, unpunctured and hairless all over.
The metasternum is punctured and hairy in the lateral and anterior intermediate areas
only ; the lateral areas are slightly broadened behind. The abdominal sterna are smooth
and glossy, except the first pair of scars which are somewhat rough. The elytra are
sparsely hairy between the shoulders; their grooves are somewhat strongly punctured:
The middie tibiae each have a small spine before the end.

Genus VETURIUS, Kaup, 1871, p. rio.
Incl. Pleurostylus, Kaup, 1871 (see Arrow, 1907, pp. 449-450).
Type. Passalus heydeni, Kaup, 18686, p. 27.

Veturius sinuatosulcatus, n. sp.
Fig. V, 3 & 5.

One specimen from Chaco, 40 mm. long.

The labrum is hghtly concave in front, lightly convex at the sides. The mandibles
(which are somewhat worn) appear to have been very like those of Arrox agassizi, but the
two lowest terminal teeth have been fused, probably at the base only, and somewhat widely
separated from the uppermost terminal tooth, especially on the left side. The head is smooth,
with a strongly elevated central tubercle flanked by straight and somewhat forwardly directed
parietal ridges. The central tubercle unites in front with a slender, finely pointed, V-shaped
ridge or pair of ridges from which the obsolete frontal ridges arise at an obtuse angle to each
other. The inner tubercles are well developed, but the frontal ridges do not reach
them.

The sides of the pronotum are strongly concave, but are overhung by the strongly
convex upper borders of the marginal grooves. The anterior margin is practically straight,
but is bordered by a strongly sinuous marginal groove which, though much narrowed near
the middle line, is scarcely broken. The pronotum is highly convex behind the median
curve of this groove, which is enlarged on either side of the convexity. At the sides the
groove is broad behind as well as in front, not narrow behind as in V. sinuatocollis, Kuwert.
The median groove is deeply impressed and practically complete.

The scutellum is coarsely punctured in the anterior angles and on either side of all but
the extreme posterior part of the middle line, bands meeting in the posterior angle to form
an {-shaped figure being left unpunctured. The mesothoracic episterna are glossy and
punctured above and along the anterior margin, matt and unpunctured behind. The
mesosternum bears matt bands in the middle line and along the sides, meeting in a broad
matt area in the anterior angle to form an 7 -shaped figure. Between these bands it is
glossy. Ibis unpunctured and hairless throughout. The metasternum is punctured and
hairy only in the anterior parts of the anterior intermediate areas and in the lateral areas ;
the lateral areas are moderately broad behind. The abdominal sterna are glossy. The
grooves of the elytra are distinctly punctured, especially laterally ; the shoulders are
without hair. There is a strong spine a little beyond the middle of the middle and hind

tibiae.
F2

36 Memoirs of the Indian Museum. [Vor. VII,

Veturius spinifer, n. sp.
Fig. V, 4, p. 34.
? Veturius sinuatus, Kuwert, 1898, p. 172 (nec Eschscholtz).

One specimen from Columbia ; two from Venezuela ; and three from Santa Catherina,
S. E. Brazil. I have selected one of the Brazilian specimens as the type of the species.
Length 42-45 mm.

Veturius spinifer agrees closely with Kuwert’s description of V. sinuatus, but lacks the
hairy shoulders of Eschscholtz’s species. It 1s closely allied to V. sinuatosulcatus but differs
in having more distinct frontal ridges, which proceed almost direct from the central to the
inner tubercles at about a right angle to one another ; in having anormal pronotum with
practically straight sides, sinuous anterior margin, marginal groove broadly incomplete
and not enlarged in front, and no anterior median convexity ; in having the scutellum
somewhat less regularly punctured; and in having the mesosternum more or less matt all
over. There are (? always) some hair-bearing punctures in the anterior angle of the
mesosternum. The anterior intermediate areas of the metasternum are almost covered
with hair-bearing punctures. The two lowest terminal teeth are smaller than the upper
one and are fused at the base, but free distally.

Veturius platyrhinus (Hope).
Passalus platyrhinus, Hope, 1845, p. 28.
Veturius platyrhinus, Kaup, 1871, pp. 111-112.
Three specimens from Chiriqui, Panama, and one from Brazil. Length 48 mm.
This species differs from the last only in its larger size, and in the absence of spines from
the middle and hind tibiae.
Veturius unicornis, n. sp.
Fig. V, 6, p. 34.
Two specimens from the Peruvian Amazon and one from the Upper Amazon. Length
Sr >
The two lowest terminal teeth of both mandibles are completely fused. The inner
tubercles and frontal ridges, as well as the outer tubercles, are entirely absent; the
central tubercle is distinct, but is less strongly elevated than in the three preceding species.
The anterior margin of the pronotum is almost straight, and the marginal groove is narrow
throughout. The scutellum is almost or quite unpunctured. The mesosternum is glossy
except for a narrow band along each side and across the middle behind the anterior angle.
The hairy portions of the anterior intermediate areas of the metasternum are very restricted.
The grooves of the elytra are somewhat more distinctly punctured than in any of the
three preceding species. In other respects this species resembles V. spinifer.

Veturius simillimus, Kuwert.
Veturius simillimus, Kuwert, 1898, p. 167.
Eight specimens from Bahia, Brazil, 35-40 mm. long.
The mandibles resemble those of V. wnicornis. The head resembles that of V. spinifer,
or may be somewhat more slender, in which case the frontal ridges meet in a more acute

1918.] F. H. GRAVELY : Passalidue of the World. 37

angle. A well-marked tubercle is sometimes present between the frontal tubercles. The
-scutellum is strongly punctured, except over a more or less well developed posterior median
band and in the lateral angles. The whole of the mesothoracic episterna are punctured ;
in the matt areas the punctures bear hairs, but not elsewhere. The mesosternum bears a
pair of glossy patches in an 1-shaped matt figure as in V. sinuatosulcatus ; these patches
bear a number of hair-bearing punctures, which become thicker on the matt surface
anterior and lateral to them, but the median matt band is hairless and unpunctured.
The metasternum is almost as extensively smooth as in V. wmicornis. In all other respects
V. simillimus resembles V. spinifer.

Veturius cephalotus (Saint-Fargeau and Serville).
Passalus cephalotus, Saint-Fargeau and Serville, 1825, p. 20 (nec Kuwert).

One specimen from Cayenne, one from Surinam, and five from the Peruvian Amazon.
Length 35-40 mm.

Veturius cephalotus was first described from Cayenne, and our specimen from that
locality bears a label showing that it was identified by Kaup. The species in our collection
which most closely resembles Kuwert’s V. cephalotus appears to me to be V. sinuatus, and one
of these has been determined as V. cephalotus by Kaup, who regarded the two as identical.
Kuwert’s V. sinuatus is probably the species described above as V. Bu fer although this
lacks the hairy shoulders of the true sinuatus of Eschscholtz.

V. cephalotus differs from V. simillimus chiefly in the absence of the spines above the
ends of the middle and hind tibiae, and in the uniformly punctured and hairy anterior
intermediate areas of the metasternum. In addition, the two lowest terminal teeth are
usually distinct at the apex on the right mandible and sometimes also on the left ; the
marginal grooves of the pronotum are inclined to be narrower in the anterior angles and
more strongly punctured behind them; and the median matt band of the mesosternum is
inclined to be broader, and the hair-bearing punctures to be more definitely concentrated
into marginal bands.

Veturius sinuatus (Eschscholtz).
Passalus sinuatus, Eschscholtz, 1829, pp. 25-26.
?Veturius cephalotus (nec sinuatus), Kuweit 1898, p. 168.

Seven specimens from Brazil, including one from Rio Grande do Sul, one from Bahia,
and two from Blumenau. Length 37-44 mm.

V. sinuatus is closely allied to V. cephalotus, but the two lowest terminal teeth, though
fused at the base, are distinct distally on both mandibles; the frontal ridges are often
obsolete ; the matt posterior angles of the mesothoracic episterna are hairless and
unpunctured ; the metasternum resembles that of V. simillimus ; and the elytra have a tuft

-of hair on the shoulders.
Veturius criniceps, Kuwert

ie WS Uo [Oa eee
Veturius eriniceps, Kuwert, 1898, p. 170.
Two specimens from Chiriqui, Panama, 32 mm. long.
The head differs from that of V. simillimus in having the central tubercle less elevated
-and set further forward, with the result that the frontal ridges meet in a very obtuse angle.

38 Memoirs of the Indian Museum. Vor Wu

The mesosternum bears matt bands arranged as in V. sinuatosulcaius, but has in addition a
band of hair-bearing punctures on either side of the whole length of the middle-line. The
anterior lateral and intermediate areas of the metasternum are entirely covered with
hair-bearing punctures. The elytra have a tuft of hair on the shoulders; they are less
glossy than in other species, although the head and pronotum are normal in this respect.
The middle but not the hind tibiae have a spine before the end. Otherwise V. criniceps
resembles the much larger V. simillimus in structure.

Veturius assimilis (Weber).
Rio. Vers pao
Passalus assimilis, Weber, 1801, p. 81.

One specimen from Rio de Janeiro, 26 mm. long.

The mandibles resemble those of V. simillimus and V. eriniceps, but the head is much
narrower than in these species, and the central tubercle is set much further back than in
the latter, being if anything somewhat further from the inner tubercles than these are from
each other. The frontal ridges are absent except close to the central tubercle, where they
are directed straight towards the inner tubercles. The scutellum is strongly punctured on
either side of a broad median band. The mesosternum resembles that of V. simillimus,
but has fewer punctures. The grooves of the elytra are very strongly punctured. The
middle and posterior tibiae have no spines before the end. In other respects this species
resembles V. simillimus.

Veturius boliviae n. sp.
His M9 psa

Five specimens from Chaco, Bolivia, 31-34 mm. long.

Veturius boliviae is very like V. cephalotus, but is smaller, has three well developed
terminal teeth on both mandibles, has very few punctures on the scutellum and the matt
parts of the mesothoracic episterna, has hair-bearing punctures on either side of the middle
line of the mesosternum as well as in front and at the sides and sometimes diffused over
other parts, the middle line being matt narrowly or not at all, and has more or less
impunctate grooves on the elvtra.

Veturius heydeni, Kaup.
Fig. V, 10, p. 34.
Passalus Heydenii, Kaup, 18685, p. 27.
? Nec. Veturius heydeni, Kuwert, 1898, p. 169.

One specimen from Mexico, 33 mm. long and very broad in proportion ; determined by
Kaup.

The frontal ridges and inner tubercles of this species are obsolete, and the short parietal
ridges are dwarfed by the large central tubercle. In other respects the structure of this
species resembles that of V. boliviae, except that the marginal grooves of the pronotum are
narrower, that there is scarcely any hair near the middle line of the mesosternum and none
between this and the marginal bands, which alone are matt, and that the elytra are more
distinctly punctured.

1918.] F. H. GRAVELY : Passalidae of the World.

The abovementioned species of Veturius may be recognized thus :—

The mesosternum hairless and unpunctured except, as a rule,
in the anterior angle af ae se
The mesosternum with hair-bearing punctures behind or at the

side , as well as in front

|
|
The anterior margin of the pronotum practically straight, ale
the anterior marginal groove sinuous; this groove almost
complete across the middle line, where the pronotum is some-
what strongly convex immediately behind it :
The pronotum normal, its anterior margin sinuous with widely
broken marginal groove N. Le se
The frontal ridges and inner tubercles present ; the two lowest
terminal teeth of both mandibles fused at base only
The fronta' ridges and inner tubercles absent ; the two lowest
_ terminal teeth of both mandibles completely fused
( The middle tibiae with spines above their ends
The middle tibiae without spines above their ends ..
The two lowest terminal teeth of both mandibles Ganley
fused (spines present above the ends of the middle tibiae ; the
5 elytra hairless at the shoulders) : nie
The two lowest terminal teeth fused at base one in unworn
specimens, on the right and usually also on the left mandible
The two lowest terminal teeth of the left mandible completely
fused (no spines above the ends of the middle tibiae ; the
elytra hairless at the shoulders)
The two lowest terminal teeth distinct at apex in unworn speci-
\ mens on both mandibles. BR
( The elytra with a tuft of hair on each shoulder
7% The elytra without any such tufts, (the middle tibiae without
\ spines above the end) à à =
The central tubercle normally | at least as far from the
inner tubercles as these are from one another ; the elytra
| normal ; the middle tibiae without spines above the end
The central tubercle set unusually far forwards, being nearer to
the inner tubercles than they are to each other ; the elytra
less glossy tha the head and thorax ; the middle las with
spines above the end Bs 2
The central tubercle situated at least as far from the mner
tubercles as these are from one another ; the obsolete frontal
ridges meeting in an acute angle ate
The central tubercle situated nearer to the inner note than
these are to each other ; the frontal ridges meeting in an
obtuse angle er = N 5
The central tubercle normal, not free at apex ; “he frontal]
ridges moderately distinct : A
The central tubercle slender, with free apex ; the frontal ridges
| cbsolete

V. sinuatosulcatus, p. 35.

—

. unicornis, p. 36.
V. spinifer, p. 36.
V. platyrhinus, p. 36.

7. simillimus, p. 36.

-—

. cephalotus, p. 37.

V. sinuatus, p. 37.

7. criniceps, p. 37.

V. assimilis, p. 38.

7. boliviae, p. 38.

V. heydent, p. 38.

40 Memoirs of the Indian Museum. AO PRES

Genus VERRES, Kaup, 1871, p. 114.
Type, Passalus corticola, Truqui, 1857, p. 310.

Verres furcilabris (Eschscholtz).
RO MES SE
Passalus furcilabris, Eschscholtz, 1829, p. 25.

Three specimens, of which two are from Para, Brazil. Length 40-44 mm.

The anterior margin of the labrum is very deeply incised. The mandibles each have
three well developed terminal teeth. The central tubercle is massive, the inner tubercles
and parietal ridges are obsolete. The median groove of the pronotum is practically
complete. The marginal grooves are abruptly terminated and deeply impressed near the
anterior angles. The scutellum is more or less densely punctured, except in the middle line
and the anterior angles. The mesothoracic episterna are glossy and densely punctured,
except in the posterior angles, which are extensively matt. The mesosternum is
unpunctured and hairless (except in the anterior angle); it may be wholly matt or
partially glossy. The metasternum is covered with hair-bearing punctures only in the
anterior parts of the anterior intermediate areas and in the lateral areas, but there are
some larger hairless punctures near the posterior margin. The abdominal sterna are smooth.
The elytra are hairless; their grooves are strongly punctured, especially laterally. The-
middle and hind tibiae are without spines before the end.

Verres sternipunctatus, Kuwert.
Fig. V, 12, p. 34
Verres sternipunctus, Kuwert, 1898, p. 174.

Three specimens from Nicaragua, 33-38 mm. long.

I am doubtful whether this species is really distinct from V. #ageni, Kaup; if not,
the name hageni must stand.

The labrum is less deeply incised than in the preceding species, and the parietal ridges
are distinct. The scutellum may be entirely covered with punctures. The mesosternum
is entirely matt, with a few hair-bearing punctures on either side of the middle line and along
the outer margins. The anterior intermediate areas of the metasternum are entirely
covered with hair-bearing punctures. The grooves of the elytra are less strongly punctured |
than in the preceding species. which this one resembles in characters not mentioned.

Verres cavicollis, Bates.
Fig. V, 13, p. 34.
Verres cavicollis, Bates, 1886, p. 24, pl. 1, figs. 20-20a, nec. Kuwert (see Arrow, 1907, p. 455).

One specimen from Guatemala, 37 mm. long.

The labrum is deeply incised, as in V. furcilabris, and is very strongly depressed behind
the incision. The mandibles resemble those of other members of the genus. The inner
tubercles are well developed, and the part of the head in front of them is short and almost
vertical. The central tubercle has a slender free apex, and the parietal ridges, though small,
are complete. The median groove of the pronotum does not nearly reach the anterior-

1918.] F. H. Gravzıy: Passalidae of the World. 41

margin. The marginal grooves are rudimentary, except in the anterior angles where they
form a pair of large circular pits punctured on the inner side. The scutellum is almost
entirely covered with punctures. The mesothoracic episterna resemble those of other
members of the genus. The mesosternum is matt and is entirely covered with hair-bearing
punctures except near the middle line behind. The metasternum is covered with
hair-bearing punctures, except the central area and the inner parts of the posterior
intermediate areas, which bear a number of hairless punctures behind. The abdominal
sterna are smooth. The elytra and legs resemble those of V. sternipunctatus.

Verres corticola, (Truqui).
Fig. V, 14, p. 24.
Passalus corticola, Truqui, 1857, p. 310.

Numerous specimens from Guatemala and one trom Mexico. Length 31-36 mm.

The labrum is slightly concave in front. The mandibles resemble those ot other
members of the genus. The central tubercle is fused with the pair of short parietal ridges to
form a massive protuberance, in front of which the more or less distinct frontal ridges
extend towards the inner tubercles at a somewhat obtuse angle to one another. The inner
tubercles are distinct as in V. cavicollis, but are situated on the anterior margin of the head
with only the angles of the clypeus visible in front of, or rather below, them. The angles of
the clypeus are somewhat more widely separated than the inner tubercles, and a pair of
rudimentary outer tubercles may perhaps be recognized in a pair of more or less tumid
areas connecting them. The addition to fig. v, 14, illustrates these tubercles and the
clypeus angle on the left side, being drawn on a larger scale than the main figure, and from
amore anterior position. The prothorax and mesothorax resemble those of V. furcilabris,
except that the scutellum is more sparsely punctured. The metasternum and elytra
resemble those of V. sternipunctatus. The abdominal sterna and legs resemble those of
other members of the gerus.

The above mentioned species of Verres may be recognized as follows :—

The clypeus extensive, oblique ; the inner tubercles obsolete
14 The clypeus smaller, almost vertical, or rudimentary ; the
inner tubercles distinct .. ae a as of wy de
The labrum very deeply cleft ; the parietal ridges obsolete . V. furcilabris, p. 40.
The labrum less deeply cleft ; the parietal ridges short but
distinct Se 5 : oy

| The labrum very deeply cleft ; the clypeus distinct; the apex
|

7. sternipunctatus, p. 40.

of the central tubercle free; à pair of large circular pits in

the anterior angles of the pronotum .. ce .. V.cavicollis, p. 40.
The labrum not deeply cleft; the clypeus hidden and

rudimentary ; the apex of the central tubercle not free ;

the pronotum normal... - 2: .- V. corticola, p. 41.

Genus PLATYVERRES, Bates, 1886, p. 9.
Type, Verres intermedius, Kaup, 1871, p. 115.

-42 Memoirs ot the Indian. Museum. Vor; VIE

Platyverres intermedius (Kaup).
Fig. V, 15, p. 34.
Verres intermedius, Kaup, 1871, p. 115.
One specimen (with worn mandibles) from Omilteme, Guerrero, lent by the British
Museum. Length 42-5 mm.

Platyverres intermedius is closely allied to Verres corticola. The labrum is, however,
somewhat more deeply excavate ; the clypeus is entirely hidden even at the angles ; the
frontal ridges are broadly arched and are more or less confluent half way between the central
tubercle and the anterior margin of the head ; the anterior ends of the marginal grooves
of the prothorax are less deeply impressed ; the lateral areas of the metasternum, though

punctured and hairy, are narrow throughout ; the elytra are united and are more coarsely
punctured in the grooves.

Genus PUBLIUS, Kaup, 1871, p. 7o.
? Incl. Procululus, Zang, 19054.

Type, Passalus crassus, Smith, 1852, p. 14.

Publius crassus (Smith).
Fig. V, 2, p. 34.
Passalus crassus, Smith, 1852, p. 14.

One specimen from Bogota, Columbia, 43-5 mm. long.

The antennal lamellae are short, being equal to about two of the immediately preceding
joints in length. The labrum is broader behind than in front ; its anterior margin is lightly
concave, its sides are lightly convex. Both mandibles are bidentate distally ; the left one
has a broad bifid tooth between these teeth and the moveable tooth; the right one has a
simple conical tooth in this position. The frontal ridges are obsolete ; the frontal tubercles
are broad and short and somewhat rounded ; the central tubercle has a slender but not
very long free apex, which is less depressed than that of Proculejus pubicostis. The
pronotum is unpunctured, and its scars are indistinct ; its median groove does not nearly
reach the anterior margin. The scutellum bears a few large punctures near the middle
line in front. The mesothoracic episterna are extensively matt and unpunctured in the
posterior angles, being glossy and punctured elsewhere. The mesosternum is glossy,
except in the scars, which become very large and almost meet in the middle line in
front. The metasternum is glossy, unpunctured and hairless except beside the middle
coxae and in the linear lateral areas. The abdominal sterna are smooth. The grooves
on the elytra are very faintly punctured.

Genus PROCULUS, Kaup, 18680, p. 8.
Incl. Cyphoproculus, Kuwert, 1896.
Type, Passalus goryi, Melly. 1833, pl. Ii

1918.] F. H. Gravety: Passalidae of the World. 43

Proculus goryi (Melly).
Passalus goryi, Melly, 1833, pl. lvi.
Two specimens from Vera Paz, Guatemala, 67-72 mm. in length.
Proculus gory: may readily be distinguished from the other two species before me by
the obsolete upper tooth of the mandibles, normally flattened mentum, and glossy elytra.

Proculus opacipennis (Thomson).
Passalus opacipennis, Thomson, 1857, pp. 420-421, pl. xxi, fig. 4.
Three specimens from Ecuador and two from Guatemala, 51-56 mm. in length.
Proculus opacipennis has a long and slightly curved upper tooth on each mandible,
a normally flattened mentum, and matt elytra.

Proculus mniszechi, Kaup, 1868.
Proculus mniszechi, Kaup, 18685, pp. 11-13.
Eight specimens from Guatemala (mostly from Vera Paz), one from Ecuador, and one
from San Pedro Sula, Honduras. They vary from 53-69 mm. in length.
Proculus mniszech has an acute and well developed upper tooth on each mandible, glossy
elytra, and a mentum with the inner margin of the forwardly directed lateral processes
turned almost at right angles to the rest of the plate in a ventral direction to form a pair

of smoothly rounded lobes.

Subfamily PASSALINAE.

The subfamily Passalinae as represented in the collection before me includes five
clearly defined genera, and a large assemblage of species separated from one another by
various combinations and modifications of characters so graded as thoroughly to obscure
their true relationships one to another.

In the first three genera the clypeus is always exposed and the antennae always
have three lamellae. The first of them, Chondrocephalus, only differs from the primitive-
Pseudacanthine genus Popilius in having no suture between the clypeus and the frons
and no hair on the lateral areas of the metasternum. C. quinquecornutus is to some
extent transitional between the two genera, having definite traces of the suture; but as
these are not very distinct and as the lateral areas of the metasternum are hairless it
seems to me to make, on the whole, a better Chondroceynalus than Popilius.

The second genus, Vindex, contains one species, V. agnoscendus, in which the
clypeo-frontal suture is distinct throughout as in the Pseudacanthinae; and but for its
flattened form and separate elytra with hairless sides this species might have been held to
indicate the relationship of its genus to Proculejus instead of to Chondrocephalus ; for
Vindex possesses the main peculiarities of the dentition characteristic of Proculejus
(see above, p. 10).

One species of Vindex, described below for the first time, has the elytra united as
they are in the next genus Proculejoides. But whereas this species retains the flattened
form characteristic of other species of Vinder, Proculejoides has assumed the more massive
form ordinarily assumed by flightless species of all groups.

44 Memoirs of the Indian Museum. IVor. VIL

The fourth genus, Paxillus, is distinguished from all others by having more than three
well developed lamellae on each antenna. In some species the clypeus is exposed and
in others it is hidden ; but all are closely related to one another and as they are not very
numerous all the genera which have been established for them may conveniently be united
into one. It is probable that a considerable proportion of the species that have been
described are invalid (see Arrow, 1907, p. 443).

The remaining species, though undoubtedly numerous, probably require proportionally
still more drastic reduction of their numbers. It seems to be impossible to separate
them into groups having the same value as the genera mentioned above. Consequently
I propose to regard almost all of them as constituting a single large and plastic genus
Passalus.

The only exception is the genus Ptichopus, the last and in some respects the most
highly specialized genus of the subfamily. It may readily be recognized from all others
by the remarkable structure of its mandibles.

The genera of Passalinae known to me may be separated thus :—

( The clypeus clearly exposed; the antennae with not more

than three well developed lamellae 14 ie BS a 2.
: The clypeus hidden! ; or, the antennae with more than three

| well developed lamellae .. ise % Er de Be 4.
ne The dentition normal ; the clypeus more or less horizontal Chondrocephalus, p. 44.
= The dentition reduced ; the clypeus more or less vertical .. a ae 3.

Much flattened insects with the inner and outer tubercles
| almost in a straight line on the upper part of the anterior

> margin of the head, and projecting forwards above the clypeus Vindex, p. 46.
| Robust insects with the cephalic tubercles normally situated Proculejoides, p. 47.
‘ The antennae with more than three well developed lamellae ;
| the clypeus exposed or hidden ARE .. Paxillus, p. 48.
4) The antennae with not more than three well developed
| lamellae ; the clypeus hidden’ AR Me er He “is 5.
The dentition normal 8 Ge Le .. Passalus, p. 51.
The upper terminal tooth very large and acute, directed
> forwards; the middle one ıudımentary or absent; the
lowest one very small, directed inwards Ae .. Ptichopus, p. 68.

Genus CHONDROCEPHALUS, Kuwert, 1896, p. 221.
Type, Popilius granulifrons, Bates, 1886, p. 12, pl. 1, figs. 13-134.
Chondrocephalus quinquecornutus, n. sp.
ie WAL, Al,

Two complete specimens from Guatemala and one dissected head. Length 17 mm.
The lamellae of the antennae are moderately long and slender. The anterior lower
tooth of the left mandible is broad and more or less distinctly bidentate, that of the right

' Exposed to some extent in Passalus guatemalensis ; see below, p. 57.

1918. | F. H. GRAVELY : Pussalidae of the World. 45

mandible slender and scarcely ıf at all bidentate. The anterior margin of the labrum is
straight, the sides all lightly convex. The parietal ridges are somewhat short. The central
tubercle is laterally compressed ; its base extends forwards to the point at which the frontal
ridges diverge towards the stout conical inner tubercles, from which tubercles they bend
abruptly outwards to end in the similar outer tubercles—a character in which this species
differs from C. granulum, Kuwert, to which it appears to be more nearly related than to
any other species yet described. The transverse course of the ends of the frontal ridges,
combined with the more irregular (though glossy) surface of the whole area in front of the
inner and outer tubercles combine in some specimens to give this area an appearance
of being definitely cut off from the rest of the head, as is the clypeus from the frons in the
genus Popilius. The anterior margin of the elypeus is more or less distinctly notched.

eg
|

Fie. VI.
Passalinæ (except Passalus) ; specifie characters in the upper surface of the head X 4.

1. Chondrocephaius quinquecornutus, Gravely. 7. Paxillus brasiliensis (St. Farg. and Serv.).
2. Chondrocephalus cordiger, Gravely. 8. Paxillus pentaphyllus (Beauvois).

3. Chondrocephalus purulenis (Bates). 9. Paxillus leachri, MacLeay.

4. Chondrocephalus granulifrons (Bates). 10. Paxıllus robustus (Percheron).

5, Vindex synelytris, Gravely. 11. Paxillus crenetus, MacLeay.

6. Paxillus camerani (Rosmini).

The angles of the pronotum are rounded, the posterior more so than the anterior.
The sides and marginal grooves of the pronotum are coarsely and irregularly punctured ;
there are no distinct scars. The scutellum bears a few punctures near the middle. The
mesothoracic episterna are glossy throughout, and are coarsely but somewhat sparsely
punctured except in the posterior angles, which are smooth. The mesosternum is smooth
and glossy, except the lateral margins which are matt. The metasternum is hairless ; its
lateral areas are narrow and slightly roughened throughout ; there are a few large punctures
on the posterior borders of the intermediate areas. The abdominal sterna are glossy. All
the grooves of the elytra are coarsely punctured. The middle and posterior tibiae are
armed with about two spines each before the apex.

Chondrocephalus cordiger, n. sp.
Fig. VI, 2.

One specimen from Guatemala, 18-3 mm. long.

The antennae, mandibles and labrum resemble those of C. quinquecornutus. The
parietal ridges are somewhat longer than in that species. The frontal ridges extend straight
from the anterior extension of the central tubercle to the small outer tubercles. Not far
from the central tubercle they are united by a curved groove, immediately beyond which
is a pair of almost obsolete inner tubercles. The somewhat heart-shaped area enclosed
between this groove and the frontal ridges 1s smooth and glossy, like the surface of the

46 Memoirs of the Indian Museum. No. WANT.

head without and behind the ridges ; in front of this groove and between the ridges it is
matt.

The marginal grooves of the pronotum are more finely and evenly punctured than
in P. quinquecornutus. The coarse punctures on the sides of the pronotum are less
numerous than in that species, and a pair of punctured scars are evident ; there are a few
coarse punctures on the dorsal surface as weli as at the sides.

The scutellum is without punctures. The mesothoracic episterna, mesosternum
and abdominal sterna resemble those of P. quinquecornutus ; the metasternum is more
extensively punctured in the posterior intermediate areas. The punctures in the grooves
of the elytra, and the spines on the middle and posterior tibiae, are not quite so
pronounced as in that species.

Chondrocephalus purulensis (Bates).
Fig. VI, 3, p. 45.
Popilius purulensis, Bates, 1886, p. 13.

One specimen from Guatemala, 22°8 mm. long.

The anterior margin of the labrum is slightly concave, and the sides are slightly
convergent behind. There are no inner tubercles. The frontal ridges, which are flattened
above, are straight and meet in a more obtuse angle than in either of the preceding species ;
they and the clypeus are glossy, the whole of the triangular area bounded by them being
matt. In other respects the head resembles that of C. cordiger, as do also the antennae
and mandibles.

The pronotum resembles that of C. cordiger in form, but the scars are more pronounced
and there are no coarse punctures on the general surface, either at the sides or nearer the
middle. The scutellum bears a group of punctures on either side of the middle line ; in
other respects the mesothorax resembles that of the two preceding species. The
metasternum bears a few punctures in the inner angles only of the posterior intermediate
areas. The abdominal sterna, elytra and legs resemble those of C. cordiger.

Chondrocephalus granulifrons (Bates).
Fig. VI, 4, p. 45.
Popilius granulifrons, Bates, 1886, p. 12.

Numerous specimens from Guatemala, 26-2-32-2 mm. long.

C. granulifrons differs from C. purulensis in having the frontal ridges (which meet in a
right angle) less flattened above, with the inner tubercles more or less imperfectly developed
and sometimes united. The small area behind and between the inner tubercles is glossy,
that between the inner and outer tubercles matt, and the clypeus more or less rough and
glossy. The puncturing on the scutellum is variable. The spines on the middle and hind
tibiae are stronger.

Genus VINDEX, Kaup, 1871, p. 78.

Type, Passalus agnoscendus, Percheron, 1841, p. 22, pl. xxviii, fig. 2.

1918.]. F. H. Gravety : Passalidae of the World. 47

Vindex agnoscendus (Percheron).
Passalus agnoscendus, Percheron, 1841, p. 22, pl. Ixxviii, fig. 2.

Seven specimens from Mexico, 20-22:4 mm. long.

This species may readily be recognized from either of the other two members of the
genus known to me by its free elytra, with coarsely, but not transversely punctured lateral
grooves. It is also characterized by the presence of a distinct trace of the lowest terminal
tooth of the right mandible, and of a distinct clypeofrontal suture. The inner tubercles
are situated upon this suture as in certain species of Pseudacanthus and Proculejus, and
the suture is most readily seen between them, where it replaces the ridge found in this
position in other species.

Vindex sculptilis, Bates.
Vindex sculptilis, Bates, 1886, p. 13.

Numerous specimens from Guatemala, 20:3-23°0 mm. long.

In this species there is no definite trace of the lowest terminal tooth on either mandible,
and there is no clypeofrontal suture. The elytra are free, and the punctures in their lateral
grooves are very large and transverse.

Vindex synelytris, n. sp.
Hig. VI, 5, p. 45.

Three specimens from Guatemala, 23°8—25°0 mm. long.

The head and mandibles resemble those of V. sculptilis. The elytra are united in
the middle line, though their form is unmodified and the wings are well developed ; the
lateral grooves are coarsely but not transversely punctured ; their dorsal grooves are more
finely punctured than in either of the other two species.

Genus PROCULEJOIDES, Kuwert, 1806, p. 221.
Type, Proculejus champion, Bates, 1886, pp. 5-6, pl. i, figs. 5-54.

Proculejoides championi (Bates).
Proculejus champion, Bates, 1886, pp. 5-6, pl. 1, figs. 5-5a.

Numerous specimens from Guatemala, which show that the species is very variable
in size (length 23-5-32°0 mm.), and that the ridges bounding the frontal area are variable,
both as to form and distinctness. |

The mandibles closely resemble those of Proculejus. The clypeus, though exposed, is
bent downwards as in Verres cavicollis and is far from conspicuous, not even being
prominent laterally as in that species. Both outer and inner tubercles are distinct, but
the frontal ridges are obsolete in front of the latter. The prothorax resembles that of
Chondrocephalus granulifrons except for its greater convexity. The scutellum is not
distinctly punctured. The mesothoracic episterna and mesosternum resemble those
of C. granulifrons except for a ventral matt patch on each of the former, The

48 Memoirs of the Indian Museum. _ [Voz. VII;

metasternum is unpunctured and has smooth hairless narrow lateral areas. The
abdominal sterna are glossy. The outermost grooves of the elytra are very indistinctly
punctured ; the dorsal grooves are unpunctured. The middle tibiae each bear two, and.
the posterior one, strong spine before the apex.

Genus PAXILLUS, Macleay, 1819, p. 105.

Incl. Paxilloides, Kuwert, 1896 ; Paxillosomus, Kuwert, 1896 ; Spasalus,
Kaup, 1868.

Type, Paxillus leachri, MacLeay, 1819, p. 106 (Paris edition, p. 20).

Paxillus camerani (Rosmini).
Fig. VI, 6, p. 45.
Paxillosomus camerani, Rosmini, 1902, pp. 4-5.

One specimen from the Upper Amazon ; 16 mm. long.

The anterior margin of the labrum is straight; the sides are lightly convex. The
last five joints of the antennae are lamellate, the middle lamella being distinctly the
longest, though this scarcely exceeds the one immediately preceding it by as much as
does the corresponding lamella of P. pentaphyllus. The mandibles each have three
distinct terminal teeth, of which the middle one is perhaps slightly nearer to the one
below it than to the one above it. The left anterior lower tooth is broader than the
right and is probably bidentate when unworn as in other species. The central tubercle
and parietal ridges are small. The frontal ridges extend from the central tubercle at a
very obtuse angle to each other towards the outer tubercles, their course being slightly
curved. The inner tubercles are distinct and are nearer to the outer than to the central
tubercle. The outer tubercles, which are bluntly conical, are little larger than the inner ;
they do not overhang the angles of the clypeus, which project horizontally in front of
them, terminating the lightly concave anterior margin of the clypeus. The area between
the frontal ridges is punctured in front of the inner tubercles, and smooth behind them.

The pronotum is coarsely punctured laterally except (? always) in the neighbourhood
of the scars. The sides and anterior margin of the pronotum are straight, with the angles
between them slightly prominent. The posterior margin and angles are rounded; the
latter are densely covered beneath with long yellowish hair. The marginal groove is fine ;
the median groove is almost complete. The posterior plate of the prosternum is broadly
truncate behind, being little narrower behind than in front. There are a few indistinct
scattered punctures on the scutellum. The mesothoracic episterna are glossy above, and
matt, except beside the anterior margin, below ; they are punctured except in the posterior
angles. The mesosternum is glossy, with deep punctured scars. The lateral areas of the
metasternum are narrow throughout, rugose, and very finely hairy ; a row of punctures
extends along the inner side of the posterior intermediate areas. The abdominal
sterna are polished except in the scars, which are matt. The elytra are densely and
somewhat extensively hairy at and below the shoulders; their lateral grooves are
coarsely, their dorsal grooves more finely, punctured.

1918.] F. H. GRAVELY : Passalidae of the World. 49

Paxillus brasiliensis (Saint-Fargeau and Serville).
Tie, WAL, Te jae 44),
Passalus brasiliensis, Saint-Fargeau and Serville, 1825, p. 21.
Paxilloides brasihiensis, Kuwert, 1898, p. 181.

Three specimens from Bolivia, one from Bogota, one from Yucatan and two without
locality labels. Length 18-21 mm.

I follow Kuwert with regard to this determination ; the original description (quoted
by Guérin, 1828, p. 90) being altogether inconclusive.

The anterior margin of the labrum is lightly concave as a rule. The ends of the five:
antennal lamellae form a straight line when furled, the middle lamella being scarcely if at
all longer than the penultimate one. The anterior margin of the clypeus may be lightly
concave, or may be lightly convex close to (but on either side of) the middle line, where
in this case it is faintly notched. The frontal ridges arise at right angles to one another
and are practically straight ; the extent of the transverse anterior punctured area between
them is variable. The posterior angles of the pronotum bear only a small patch of
short hair beneath. The mososternal scars may be smooth or rugulose, glossy or matt .-
The lateral areas of the metasternum are hairless. The strength of the puncturing of the
dorsal grooves of the elytra is somewhat variable. The shoulders of the elytra are
somewhat hairy, but are not densely covered with long pile as in the preceding species. In:
other respects the present species resembles the last.

Paxillus pentaphyllus (Beauvois).
Fig. VI, 8, p. 45.
Passalus pentaphyllus, Beauvois, 1805, p. 2, pl. i, fig. 2.

One specimen from Mosquito 26°3 mm. long.

The anterior margin of the labrum is somewhat more strongly convex than in the-
preceding species. The antennae resemble those of P. cameram, but the ends of their
lamellae when furled form a somewhat more strongly curved line. The outer tubercles.
are long and slender, and are directed forwards above the angles of the clypeus which,
however, are exposed beneath them. The area between the outer tubercles is glossy and
unpunctured. In other respects this species resembles the preceding one, except that
the posterior intermediate areas of the metasternum are more extensively and very coarsely
punctured.

Paxillus leachii, Macleay.
Fig. VI, 9, p. 45.
Paxillus leachii, MacLeay, 1819, p. 106 (Paris edition, p. 20).

Many specimens from Guatemala, Iquitas (Upper Amazon), Bahia, Rio de Janeiro,
Para, Esperito-Santo (Brazil), Bolivia, Nicaragua, British Honduras, Mexico, and
Montevideo. Length 16°0-21-5 mm.

This species differs from the last only in having the anterior margin of the labrum less
strongly concave, in having the ends of the antennal lamellae in a straight line when the

antennae are furled, in having coarse punctures. between the outer tubercles of the head,
H

50 Memoirs of the Indian Museum. [Vor. VII,

in having the lower side of the prothorax still less hairy, and in having the posterior
intermediate areas of the metasternum more finely punctured.

Paxillus robustus (Percheron).
Fig. VI, 10, p. 45.
Passalus robustus, Percheron, 1835, pp. 35-36, pl. in, fig. 1

Several specimens from different localities in Brazil (Santa-Catherina, Bahia, Rio de
Janeiro and Esperito-Santo). Length 132-180 mm.

This species and the next are less stronglv flattened than any of the preceding members
of the genus; they have the posterior plate of the prosternum strongly tapered behind
and have the clypeus entirely hidden ; the outer tubercles are variable in size, but are
prebably always somewhat long and slender in unworn specimens, and the frontal ridges
are obsolete between them and the inner tubercles, though well developed between the
inner tubercles and central tubercle. The anterior angles of the head are sometimes
rather strongly produced and acute. The sides of the pronotum are more extensively,
and the grooves of the elytra more coarsely punctured than in P. leachi, which this species
resembles in other respects, except that the shoulders of the elytra are entirely hairless.
The tibiae of the intermediate (and to a less extent the hind) legs often bear several strong
spines on the outer side, especially in small specimens.

Paxillus crenatus, MacLeav.
Fig. VI, 11, p. 45.
Paxillus crenatus, MacLeay, 1819, p. 10; (Paris edition, p. 20).

Several specimens from the Upper Amazon, Guadaloupe, Surinam and Mosquito.
Length 17'0-19:5 mm.

This species differs from the last only in having the frontal ridges broadly arcuate,
instead of meeting abruptly, in having the anterior extremities of the marginal grooves
of the pronotum broader and deeper, and in having the metasternum and elytra somewhat
less strongly punctured.

The above mentioned species of Paxillus may be recognized from one another, as
follows :-

J) tern flattened insects, with the clypeus more or less exposed de
| More robust insects, with the clypeus entirely hidden %,
‘ The outer tubercles short and blunt, not directed forwards
above the clypeus = I = 20 er Se 3.
J The outer tubercles long and slender in unworn specimens,
| directed forwards above the angles of the clypeus which
\ they tend to obscure .. : or aie se i aE

The frontal ridges meeting in a nde obtuse angle ; the
posterior angles of the pronotum, and the shoulders af the

| elytra, densely covered beneath with long yellowish hair P. camerani, p. 48.
®\ The frontal ridges meeting in a right angle ; the posterior angles
of the pronotum, and the shoulders of the elytra, less hairy

i beneath ie Le se 36 .. P. brasihensis, p. 49.

1918. | F. H. Gravety: Passalidae of the World. 51

The ends of the antennal lamellae in a strongly curved line
when furled ; the anterior margin of the head unpunctured P. pentaphyllus. p. 49.
The ends of the antennal lamellae in a straight line when furled ;

|
| the anterior margin of the head coarsely punctured between
the outer tubercles as a a .. BP.leachn, p. 49.
( The frontal ridges straight, arising at right angles to each
N other es me Is oe .. P. robusius, p. 50:

The frontal ridges together forming a crescent .. D PScrenatüs, ps OU:

Genus PASSALUS, Fabricius, 1792, p. 240.

— Neleus nec Passalus, Kaup, etc. (see Zang, 1905¢, pp. 224 and 226).

Incl. *Epiphanus+ Eumelus+ Lucilius+ Mitrorhinus!+"Neleides+*Ninus (=Scalmus, Zang)
+ "Pertinax+"Petrejus+"Phoroneus (=Macrolobus, Zang) + Rhagonocerus+*Rhodocan-
thopus + Siephanocephalus! + *Vatinius (=Zosterothrix, Zang), Kaup. Also *Apone-
leides+ Cassius+ Epipertinax+*Flavius (=Lasioperix, Zang), + Lophocephalus+ Manlius
+ Microthorax (=Phaulothorax, Zang) + *Morosophus+*Neleuops + Ninoides + Oeneus
+ Parapertinax + *Pertinacides + Phanocles (=Psilomus, Zang) + *Phoronaeosomus +
Polyacanthopus+*Ptychotrichus (—EÆEpipleurothrix, Zang) +Severus+Synesius+*Tetr-
aracus+ Thryptocerus (= Alococerus, Zang) + Toxeutotaenius+*Trichopieurus+ Valerius,
Kuwert. Also *Æpiphoroneus, Arrow.

Type, Lucanus interruptus, Linnaeus, 1767, p. 560.”

It seems probable, and has been assumed in the above synonymy, that all genera
belonging to Kuwert’s subfamilies Rhodocanthopinae, Neleidinae, Pertinacinae, Pleurariinae
(except Pleurarius which belongs to the Indo-Australian Series), Phoronaeinae, Petrejinae,
Vatiniinae and Neleinae should be included in this complex and heterogeneous genus ;
but I have been able to confirm this by reference to specimens only in the case of the
genera marked with an asterisk (*). The genus Prosochtus, Bates, has been omitted
because I think, judging from Bates’ description, which is all that Kuwert also had to go
upon, that Bates was probably right in regarding the species for which he founded it as
closely allied to the genus Proculejus.

One of Kaup’s species of Proculejus, P. quitensis, was rightly removed by Kuwert to
one of his subfamilies here included in the genus Passalus ; but the genus Prosoclitus, in
which he placed it, is probably allied to if not identical with Proculejus, not Passalus.
P. quitensis differs from all other species of Passalus known to me in having the elytra
united ; but, if this character is to be regarded as in itself sufficient to warrant the
separation of the species possessing it into special genera, new genera will be required
for one of the four described species of Vindex, and for each of two species that otherwise
fall into two widely separated divisions of the genus Macrolinus. This multiplication of
small genera seems to me undesirable, although I have found it convenient to retain the
already existing small flightless genera Platyverres and Publius, instead. of uniting them

1 See above, pp. 10-11.
2 This is the reference usually given, but is not the earliest description. See below, p. 63, footnote.

H 2

52 Memoirs of the Indian Museum. Dom WI,

with Verres and Veturius respectively, as would have been mere consistent. Proculejus
quitensis consequently becomes Passalus quitensis.

In spite of the complex manner in which various specific characters are intermingled
the species of Passalus can be arranged in a series leading up from forms with the margin
of the head between the outer tubercles straight or faintly notched in the middle line, the
inner and outer tubercles usually more or less widely separated, and the lateral areas of
the metasternum usually narrow, smooth and hairless, to forms with the margin of the head
abruptly emarginate in the middle line, the inner and outer tubercles in contact with one
another, together forming an oblique edge, and the lateral areas of the metasternum broad,
punctured and hairy. The following descriptions have been placed in this order.

Passalus nanus (Kuwert).
Rhodocanthopus nanus, Kuwert, 1898, p. 139.

One or two specimens from each of the following localities—Guatemala, Ecuador,
and the Cauca Valley in Columbia. Length 15-3-17-7 mm.

This species is very like Paxillus robustus, but its antennae have only three well
developed lamellae, its frontal ridges are apt to be more denticulate (especially in Columbian
specimens), its eyes are smaller and less prominent, its pronotum and elytra are much more
coarsely punctured, the punctures in the lateral grooves of the elytra being more or less
distinctly transverse, and its intermediate and hind tibiae are still more strongly spined.
The epipleura are unpunctured and hairless as in Paxillus robustus.

Passalus rugosus, n. sp.
Fig. VIL, 1.

Three specimens from the Cauca Valley, Columbia. Length 21:0-21:8 mm.

The central tubercle is relatively nearer to the anterior margin of the head than in
P. nanus, the frontal ridges consequently diverging in a more obtuse angle. The sides of
the pronotum are very coarsely punctured, but there are no coarse punctures nearer the
middle as in C. nanus, nor are there any on the central area of the metasternum as in that
species. The grooves of the elytra are even more strongly punctured than in C. nanus, the
punctures in the lateral grooves being distinctly transverse. The epipleura are punctured
and hairy. In other respects this species resembles P. nanus.

Passalus punctato-striatus, Percheron.
Fig. VII, 2.
Passalus punctato striatus, Percheron, 1835, pp. 78-79, pl. vi, fie. 1.

A large number of specimens from Guatemala and Mexico, some of the latter being
from Oaxaca; a few from San Salvador, Honduras (San Pedro Sula), Nicaragua, Columbia
and Surinam. Length 20:3-28-3 mm., relative breadth extremely variable ; the distance
between the inner and outer tubercles, and other characters, also variable to some extent.

This species differs from the last only in having the frontal area slightly larger, the
pronotum and the grooves of the elytra less coarsely punctured, the epipleura unpunctured

1918.] F. H. GRAVELY : Passalidae of the World. 53

and hairless, and the middle and hind tibiae usually unspined or nearly so.! The
difference is most marked as regards the punctures in the dorsal grooves of the elytra, those
in the pair of grooves nearest the suture being almost impunctate in most specimens of the
present species.

Fic. VII.
Passalus spp. ; specific characters in the upper surface of the head x 4.

1. P. rugosus, Gravely. 13. P. interstitialis, Eschscholtz.
2. P. punctato-striatus, Percheron. 14. P. spinipes, Gravely.

3. P. rhodocanthopoides (Kuwert). 15. P. abortivus, Percheron.

4. P. morio, Percheron. x 16. P. mucronatus, Burmeister.
5. P. latifrons, Percheron. 17. P. quadricollis Eschscholtz.
6. P_ pertyi (Kaup). 18. P. occipitalis, Eschscholtz.
7. P. quitensis (Kaup). 19. P. nasutus, Percheron.

8. P. catherinae, Gravely. 20. P. polli, Gravely.

9. P. eucadorensis, Gravely. 21. P. punctatissymus, Eschscholtz.
10. P. curtus (Kaup). 22. P. opacus, Gravely

11. P. prominens, Gravely. 23. P. glaber, Gravely.

12. P. guatemalensis (Kaup). 24. P. erosus, Truqui.

Passalus rhodocanthopoides (Kuwert).
Ties WAUL 3%
Neleuops rhodocanthopoides, Kuwert, 1898, pp. 142-143.
Several specimens from Peru (Cumbasa and the Amazon region). Length 24-25 mm.
A somewhat flatter insect than the preceding, with the inner tubercles situated
almost vertically behind the outer tubercles but separated from them by a well marked
concavity. The posterior intermediate areas of the metasternum are coarsely punctured,

1 One of the Mexican specimens has numerous small spines on these tibiae, and the Columbian specimen has them
numerous and very strong. The latter specimen has the elytra more coarsely punctured than any other that I have

seen and may prove to belong to a different species.

54 Memovrs of the Indian Museum. Vor. Vie

the lateral areas are hairy and punctured, as are also the epipleura and the shoulders and
anterior half of the lower margin of the elytra. In other respects this species resembles
the preceding one.
Passalus morio, Percheron.
Fig. VIL, 4, p. 53.
Passalus morio, Percheron, 1835, pp. 83-84, pl. vi, fig. 4.

Sixteen specimens from Brazil (Espirito-Santo, Bahia, Blumenau, Rio). Length
10-2775 mm.

This species is somewhat more convex than either of the two preceding. The length
of the antennal lamellae is somewhat variable.

The head is very like that of P. rhodocanthopoides, but the outer tubercles are very
obtuse, the anterior margin is somewhat thickened, the frontal area behind this thickening
is flat with no special depression between the inner and outer tubercles, and the inner
tubercles are situated much nearer together than the outer. The punctures on the
pronotum are usually confined to the scars and marginal grooves, but may be absent from
the former or may extend beyond them. The mesosternal scars are represented only by
triangular matt areas which are not sunk below the level of the surrounding surface. The
posterior intermediate areas of the metasternum are as a rule strongly punctured ; the
lateral areas are more or less linear, smooth or slightly roughened, with or without
hair-bearing punctures. The epipleura are hairless and unpunctured, the puncturing of the
elytra in this and other respects resembling that of P. punctato-striatus. The armature.
of the middle and hind tibiae is variable, never as strong as in P. rugosus.

Passalus latifrons, Percheron.
Fig. VII, 5, p. 53.
Passalus latifrons, Percheron, 1841, pp. 32-33, pl. Ixxix, fig. 1.

Three specimens from Surinam, length 30:6-32-1 mm.

P. latifrons is very like P. morio, and will perhaps prove to be no more than a local race
of it. The outer tubercles are more prominent and consequently less obtuse ; the inner
tubercles are less distinct ; the frontal ridges diverge at a more obtuse angle and extend
more distinctly beyond the latter towards the former. The anterior angles of the pronotum
are produced forwards to form a somewhat obscure but distinctly acute projection. The
punctures in the pronotal scars and on the posterior intermediate areas of the metasternum
are few in number in all our specimens, and the lateral areas of the metasternum are smooth
and hairless. The mesosternal scars are entirely absent.

Passalus pertyi (Kaup).
Fig. VII, 6, p. 53.
Pertinax pertyi, Kaup, 1869, p. 22.
Two specimens without locality labels. Length 39:0-41:5 mm.
The frontal area is smaller in all directions than in P. latifrons and more densely
punctured in front. The inner tubercles are obsolete and the frontal ridges do not reach
the outer tubercles. The anterior angles of the pronotum are strongly rounded. ‘The

1918. ] F. H. Gravety: Passalidae of the World. 55

posterior intermediate areas of the metasternum are more extensively punctured than
in P. latifrons, which the present species resembles ın other respects.

Passalus convexus, Schönherr.
Passalus convexus, Schönherr, 1817, p. 333, and appendix pp. 142-143.

Ten specimens from Cumbase (Peru), Tejuca, Upper Amazon, and Amazonas.
Length 38-0-43:7 mm.

This species, like the last, is closely allied to Passalus latifrons, from which it differs
only in its larger size, in not having the frontal ridges continued beyond the inner tubercles,
in having less prominent outer tubercles, and in having the anterior angles of the pronotum
‘more or less rounded.

Passalus quitensis (Kaup).
Fig. VII, 7, p. 53.
Proculejus quitensis, Kaup, 1871, pp. 63-64.

Described from a cotype from Quito, lent by the British Museum. Length 32 mm.

In addition to its rounded pronotum and fused and rounded elytra Passalus quitensis
differs from P. convexus in having the inner tubercles directly behind and much nearer
to the outer tubercles, in having curved instead of straight frontal ridges, in having well
developed mesosternal scars, and in having the posterior intermediate areas of the metas-
ternum strongly and extensively punctured.

Passalus affinis, Percheron.
Passalus affinis, Percheron, 1835, pp. 72-73, pl. v, fig. 5.

Several specimens from Cuba, three from St. Domingo and one from Haiti. Length
-37°5-42;0 mm. Relative breadth very variable.

The frontal area resembles that of P. quitensis, but is more closely and extensively
punctured in front, while the inner and outer tubercles are contiguous, together forming
a pair of more or less oblique ridges on the anterior margin of the head, as in P. interruptus,
etc. The mesosternal scars are well developed; the posterior intermediate areas of
the metasternum are coarsely and extensively punctured. The lateral areas of the
-metasternum, the epipleura and the shoulders of the elytra are punctured and hairy. In
other respects this species resembles P. convexus.

Passalus catharinae, n. sp.
Fig. VIL, 8, p. 53.

One specimen from Santa Catharina and one from Chaco. Length 31-33 mm.

The head is very like that of P. affinıs, but the frontal ridges and all the tubercles are
more strongly elevated, the broad and rectangular or obtuse apex of the central tubercle
slightly overhanging the frontal area, which is more or less sparsely punctured. There
are a few strong punctures above the pronotal scars. The posterior intermediate areas
of the metasternum bear a few coalescent punctures along the inner margin ; the lateral
areas are linear, hairless and unpunctured. The elytra resemble those of P. affinis except
that the dorsal ridges are less and the lateral more coarsely punctured, the three grooves

56 Memoirs of the Indian Museum. [Vor. VII,

nearest the suture being unpunctured, the fourth containing more or less obsolete
punctures, the fifth and sixth containing strong round punctures, and the seventh, eighth
and ninth containing larger and more or less transverse punctures. In other respects
this species resembles P. affinis.

Passalus recticlypeatus (Kuwert).
Petrejus recticlypeatus, Kuwert, 1898, p. 202.

Four specimens, without locality record. Length 23-3-24:7 mm.

This species is very like the preceding one, but is smaller and has the apex of the central
tubercle acute, free and directed forwards above the finely roughened but unpunctured
frontal area, the inner tubercles being obsolete or absent. The posterior intermediate
areas of the metasternum are almost or quite unpunctured ; the lateral areas are more or
less rugose and bear a few fine hairs. The lateral grooves of the elytra, though much more
strongly punctured than the dorsal, are less strongly punctured than in Passalus catherinae,
which this species resembles in other respects.

Passalus eucadorensis, n. sp.
Fig. VII, 9, p. 53.

One specimen from Ecuador, 20:4 mm. long.

In this species the cephalic tubercles and ridges resemble those of P. catherinae, except
that the central tubercle is weaker ; otherwise the insect resembles P. recticlypeatus, except
that the anterior margin of the head is slightly prominent in the middle line, and that the
metasternum and elytra are entirely hairless.

Passalus curtus (Kaup).
Fig. VII, 10, p. 53.
Petrejus curtus, Kaup, 1869, p. 38.

One specimen from the Cauca Valley, 23:3 mm. long.

The free apex of the central tubercle is longer than in P. recticlypeatus, but the frontal
ridges are less strongly elevated and become obsolete in front. The frontal area is
irregularly marked with coarse punctures. The sides of the pronotum and the posterior
intermediate areas of the metasternum are somewhat more extensively punctured. The
mesosternal scars and the lateral areas of the metasternum are finely punctured and
hairy. The epipleura are without, and the shoulders of the elytra almost without,
hair-bearing punctures. The puncturing of the dorsal grooves of the elytra is about
as coarse as that of the lateral grooves. In other respects this species resembles
P. recticlypeatus.

Passalus gracilis (Kaup).
Petrejus gracilis, Kaup, 1869, p. 38.

One specimen from Columbia, 16-5 mm. long.
A much smaller species than the preceding, with more flattened central tubercle,

Pas

conical forwardly directed parietal ridges, somewhat more distinct frontal ridges, smooth

1918.] F. H. GraveLy : Passalidae of the World. 57

unpunctured frontal area, hairless mesosternal scars, metasternal lateral areas and elytra
shoulders, and less coarsely punctured dorsal grooves on the elytra.

Passalus prominens, n. sp.
Innes, WANT, al io, 58:

One specimen from the Peruvian Amazon, 22 mm. long, and somewhat convex.

The anterior margin of the head is shghtly prominent as in P. eucadorensis, forming
an obtuse angle in the middle line ; but the frontal area is larger, being quite half as long
as it is wide in front, and its general surface is smooth and somewhat sparsely marked with
large punctures instead of being finely roughened all over. The central tubercle is laterally
compressed, with an antero-posteriorly obtuse apex situated behind the junction of the
frontal ridges. The frontal tubercles are obsolete. in other respects this species
resembles P. gracilis, except that the anterior angles of the pronotum are more acute and
the posterior angles (like the hind part of the posterior intermediate areas of the
metasternum) are somewhat densely punctured, the anterior ends of the marginal grooves
being densely punctured and strongly enlarged.

Passalus guatemalensis (Kaup).
Inne, WIT, NZ 106 GB,
Oileus guatemalensıs, Kaup, 1869, p. 6.

Three specimens from Nicaragua, 19'6-20'3 mm. long.

Passalus guatemalensis differs from all other species of Passalus known to me in having
a vertical and more or less conspicuously exposed clypeus. It should perhaps be regarded
as the type of a distinct genus allied to Vindex, but such a course would hardly be justi-
fiable at present.

P. quatemalensis is a somewhat flatter insect than the last, with a more or less
punctured obtuse-angled frontal area, the frontal tubercles being situated about half way
between the less prominent central and more prominent outer tubercles. The margin of the
head between the outer tubercles is straight, with or without a median notch. The sides of
the pronotum are somewhat rounded, and the anterior angles are not acute; the median
groove 1s complete, and the anterior ends of the marginal grooves are scarcely enlarged or
punctured ; the scars are transverse and punctured, with a longitudinal band of punctures.
above them. In other respects this species resembles P. prominens, except that the
punctures on the posterior intermediate areas of the metasternum are more widely

dispersed.
Passalus incertus, Percheron.

Passalus incertus, Percheron, 1841, pp. 27-28, pl. Ixxviii, fig. 4.

Six specimens from the Cauca Valley, and one from Venezuela. Length 19°4-24°3 mm.

A slender and somewhat flattened insect, differing from the last only in having the

clypeus entirely hidden, the median notch of the anterior margin of the head sometimes
I

58 Memoirs of the Indian Museum. Vom WILL,

very pronounced, the frontal area more or less rugose but unpunctured, the frontal ridges
strongly arched, the inner and outer tubercles nearer together, the anterior ends of the
marginal grooves of the pronotum more strongly dilated and punctured, the sides of the
pronotum more extensively punctured in small and less in large specimens, the posterior
intermediate areas of the metasternum less strongly and extensively punctured, the grooves
of the elytra somewhat more strongly punctured, and the shoulders and epipleura
punctured and hairy. The spines on the middle tibiae are variable in number ; in none of
our specimens are they very strong.

Passalus interstitialis, Eschscholtz.

Fig. VII, 13, p. 53.
Passalus interstitialis, Eschscholtz, 1829, pp. 18-19.

One or more (often numerous) specimens from each of the following localities :— Mexico,
Guatemala, Honduras (San Pedro Sula), Panama (Chiriqui), Cuba, Surinam, Peru (Cumbase
and Amazon), Brazil (Amazonas, Pernambuco, Bahia, Rio and Blumenau), Bolivia (Cordico
and Farinas) and Paraguay. Length 24 34 mm.

The anterior margin of the head is much more extensively notched in the middle line
than in the preceding species, and the notch is bounded by a pair of very prominent
tubercles. These tubercles are fully as strong as the outer tubercles, which are somewhat
small and are more or less obscured by the inner tubercles, the latter being very long and
projecting from a point contiguous to and a little behind and on the outer side of them.
From the inner tubercles the imperfectly denticulate frontal ridges extend in a straight
line backwards and inwards to meet in a right or slightly obtuse angle. The anterior part
ot the frontal area is more or less punctured.

The pronotum resembles that of the preceding species in shape and is more or less
extensively punctured at the sides, but the anterior ends of the marginal grooves are not
expanded. The mesothorax is normal. The posterior intermediate areas of the
metasternum are closely and coarsely punctured ; the lateral areas are moderately broad
and are covered with hair-bearing punctures. The dorsal grooves of the elytra are
strongly but finely punctured, the lateral grooves are coarsely punctured. The epipleura
shoulders and anterior half or two-thirds of the outermost rib of the elytra are thickly
covered with hair-bearing punctures. The middle tibiae are armed with one stout spine.

Passalus glaberrimus, Eschscholtz.

Passalus glaberrimus, Eschscholtz, 1829, p. 20.

Four speeimens, of which three are from Brazil (Blumenau and Rio), and one bears
no record. Length 20 mm.

Passalus glaberrimus is very like P. incertus, but has the anterior margin of the head
more definitely notched than is usual in that species, the sides of the pronotum usually
more extensively punctured, the anterior angles of the pronotum somewhat more acute
and the elytra devoid of hair-bearing punctures.

1918.] F. H. Gravety: Passalidae of the World. 59

Passalus spinosus (Kuwert).

Rhodocanthopus incertus, Kuwert, 1898, p. 140.

Two specimens from Chiriqui, Panama, 20:3 mm. long. A somewhat more robust
insect than the preceding.

The frontal ridges are straight and meet in a right angle ; the inner tubercles are situated
about half way from the central to the outer tubercles and there is a pair of small secondary
tubercles between them and the latter ; the anterior margin of the head is broadly notched
in the middle ; the anterior part of the frontal area is moderately strongly punctured. The
pronotum resembles that of P. incertus in shape, but is only punctured in and close to the
scars and in the marginal grooves, whose anterior ends are smaller. The mesothorax
resembles that of P. incertus. The metasternum differs from that of P. incertus only in
having the lateral areas somewhat broader and less smooth behind. The elytra are
hairless ; their four dorsal grooves are about as strongly punctured as in P. incertus ; the
next four are marked with very large transverse punctures, the transverse ridges between
which tend to become obsolete behind. In the two remaining grooves these ridges
are obsolete throughout, and at the extreme posterior end the longitudinal ridges become
obsolete also, the remaining surface being matt. The middle and hind tibiae are armed

with three or four strong spines.

Passalus spinipes, n. sp.
Fig. VII, 14, p. 53.

One specimen from Nicaragua, 22:7 mm. long.

This species is closely allied to P. spinosus, but the inner tubercles and the secondary
tubercles in front of them are less pronounced, the sides of the pronotum bear a longitudinal
band of punctures above the scars, the posterior intermediate areas of the metasternum are
less extensively punctured, the lateral areas are extremely narrow throughout, the
punctures in the grooves of the elytra are finer, those in the lateral grooves though

moderately coarse not being transverse.

Passalus spiniger (Bates).
Rhodocanthopus spiniger, Bates, 1886, pp. 15-16, pl. 1, figs. 16-16a.

One specimen from Columbia, 22 mm. long.

This species is very like the last two, but the anterior margin of the head is less broadly
and perhaps more sharply notched, the frontal area is scarcely as long or as distinetly
punctured, there are no secondary tubercles between the inner and outer tubercles, the
sides of the pronotum are more coarsely and extensively punctured, the lateral areas of the
metasternum are intermediate between the two in width, and the puncturing of the grooves
of the elytra is somewhat coarser than in P. spinosus above and less coarse (scarcely if at all
transverse) at the sides, all the transverse ridges being distinct and broadly elevated as

in P. spinvpes.

‘60 Memoirs of the Indian Museum. Von, WAUL,

Passalus abortivus, Percheron.
Ines, WAUL, 215, 10, 83.
Passalus abortivus, Percheron, 1835, pp. 87-89, pl. vi. fig. 7.

Three specimens from the Amazon, of which at least two are from Peru. Length
28:0-28°3 mm. long. One specimen with deformed antennae.

This species differs from all other species of Passalus known to me in sometimes having
a distinct lamella in front of the three ordinarily found on the antennae of species of this
genus. This lamella, however, is not fully developed as are the additional lamellae found
in the genus Pazillus, but is either much slenderer than those following it as well as only
about half their length, or is quite short and inconspicuous.! The anterior margin of the
head bears a pair of small but well developed tubercles on either side of the median notch,
which is consequently much more pronounced than is ever the case in Paxillus. The
cephalic tubercles and ridges closely resemble those of Passalus spiniger. The pronotum
resembles that of P. spiniger in shape, but is unpunctured except in the scars and marginal
grooves. An oval patch a little below the middle of the mesothoracic episterna, the
mesosternal scars, the lateral areas of the metasternum, and the lower parts of the
shoulders of the elytra (but not the epipleura) are covered with hair-bearing punctures.
The posterior intermediate areas of the metasternum are coarsely punctured behind and
on the inner side. The elytra are strongly and uniformly punctured. The middle and hind
tibiae bear from one to three spines which are stronger on the former than on the latter.

Passalus jansoni (Bates).
Phoroneus jansoni, Bates, 1886, p. 18, pl. i, figs. 17-17a.

One specimen from Nicaragua, 32 mm. long.

This species is very like the last, but the triconcave margin of the head between the
outer tubercles is thickened and lightly convex as a whole; the frontal area and the
pronotum (even in the scars and anteriorly weak marginal grooves) are unpunctured ;
the mesothoracic episterna, the mesosternal scars, the metasternum and the elytra are
hairless, the second and third being entirely unpunctured, and the last being punctured
in the lateral grooves only ; and there are no very distinct spines on the middle and hind
tibiae.

Passalus mucronatus, Burmeister.
Fig. VII, 16, p. 53.
Passalus mucronatus, Burmeister, 1847, pp. 488-489.

One specimen from Columbia and one from Guatemala. Length 24 mm. :

The head and pronotum resemble those of the preceding species in outline, but the
central tubercle is much elongated with free decumbent apex, the frontal ridges are feebly
developed, the anterior margin of the head is not thickened, and the sides of the pronotum,
including the scars and anteriorly enlarged marginal grooves, are strongly punctured. The

+ In one of our two specimens in which the antennae are not deformed it is slender and about half the length of
the others ; in the other it is scarcely if at all different from the enlargement often found in the same position in other
‘species.

1918.] F. H. GRAVELY : Passalidae of the World. 61

mesothorax is normal. There are a number of coarse punctures on the posterior
intermediate areas of the metasternum, the lateral areas being linear, smooth and hairless.
The anterior intermediate areas of the metasternum, and the shoulders of the elytra with
the anterior parts of the epipleura, are covered with hair-bearing punctures, the shoulders
of the elytra being densely hairy. All the grooves of the elytra are distinetly punctured,
the lateral scarcely more strongly than the dorsal. The middle and hind tibiae are
without distinct spines.

Passalus quadricollis, Eschscholtz.
Fig. VIL, 17, p. 53.
Passalus quadricollis, Eschscholtz, 1892, pp. 21-22.
Phoroneus quadricollis, Kaup, 1871, p. 102.

Two specimens from Brazil, one of them in the collection of M. Guy Babault of Paris.
Length 33 mm.

The central tubercle is set further back than in P. mucronatus and the free apex extends
forwards horizontally at right angies to the massive base by which it is raised high above
the large and coarsely rugose frontal area. The frontal ridges and inner tubercles are more
strongly developed than in P. mucronatus. The pronotum resembles that of P. mucronatus,
except that the punctures are confined to the scars and marginal grooves, those in the
former being of very large size. The mesosternal scars are matt, but are not depressed.
The inner angles of the posterior intermediate areas of the metasternum are marked
with very large and more or less coalescent punctures ; the anterior intermediate and lateral
areas of the metasternum and the shoulders of the elytra are covered with hair-bearing
punctures. The grooves of the elytra are marked with shallow punctures which are very
broad in the lateral grooves. The middle tibiae bear a stout spine on the outer side.

Passalus occipitalis, Eschscholtz, 1829.
Fig. VIL, 18, p. 53.
Passalus occipitalis, Eschscholtz, 1829, p. 21.

One specimen 33 mm. long. Locality not recorded.

This species is closely related to the last, but the massive central tubercle is set still
further back and has no free horizontal apex. The frontal ridges diverge at about a right
angle and then bend forwards and run parallel to one another as far as the inner tubercles,
which are situated not very far behind the outer. The anterior margin of the head, though
straight as a whole, is a little irregular ; it is not distinctly notched in our specimen ;! the
whole anterior part of the frontal area is thickly covered with large punctures. The whole
of the thorax resembles that of P. quadricollis, except that the mesothoracic scars are to
some extent depressed, and the punctures in the inner anges of the posterior intermediate
areas of the metasternum are much smaller. The elytra are punctured as in that species
in the lateral grooves, more finely or not at all in the dorsal ones. The legs resemble
those of that species.

1 See also, however, Arrow, 1907, pp. 459-460.

62 Memoirs of the Indian Museum. [Vor. VII,

Passalus nasutus, Percheron.
Bje-aVI21955p253:
Passalus nasutvs, Percheron, 1835, pp. 90-91, pl. vi, fig. 8.

One specimen from Parana, 24°4 mm. long.

This species resembles the preceding, but is much smaller ; the central tubercle is conical,
free distally and directed forwards and a little upwards; the pronotum has less acute
anterior angles and somewhat more numerous punctures in the scars ; the epipleura and
shoulders of the elytra are densely covered with hair-bearing punctures; the grooves of
the elytra are punctured much as in P. quadricollis.

Passalus polli, n. sp.
Fig. VII, 20, p. 53.

One specimen from Joinville, 34-5 mm. long.

Passalus polli is much larger than P. nasutus and has a smaller central tubercle
situated somewhat further forwards, whose apex is scarcely free. The frontal ridges are
small and extend almost directly outwards, then arching slightly forwards to end in
the inner tubercles, which are equidistant from the central and outer tubercles; the
whole surface of the head in front of the inner tubercles is rugose and glossy. The
anterior ends of the marginal grooves are somewhat curved but scarcely expanded. The
mesosternal scars are depressed and covered with moderately large, indistinct, coalescent
punctures. The posterior intermediate areas of the metasternum are closely covered with
coarse punctures; the lateral areas are very broad, especially behind, and are covered
with hair-bearmg punctures. The puncturing of the grooves of the elytra resembles that
of P. quadricollis, but is somewhat deeper laterally; the shoulders, epipleura and
anterior half of the mb immediately above each of the thickened lateral margins are
covered with hair-bearing punctures. The middle and hind tibiae are each armed with one
stout spine.

Passalus toriferus, Hschscholtz.
Passalus toriferus, Eschscholtz, 1829, pp. 17-18.

Three specimens from Brazil, one from Yucatan, and one said (no doubt incorrectly)
to come from 8. Australia. Length 28-34 mm.

This species differs from the last only in having the anterior part of the head less rugose
(the frontal area sometimes punctured), the inner tubercles situated close behind and slightly
on the outer side of the outer tubercles, the sides of the pronotum strongly and coarsely
punctured at least near the scars, and the extreme anterior part of the eighth groove of
the elytra hairy, The central tubercle is very variable and may be distinctly or not at all
free distally.

Passalus punctatissimus, Eschscholtz.
Ms WANE, Ally jd. 58:
Passalus punctatissimus, Eschscholtz, 1829, pp. 19-20.

A number of specimens from the Peruvian and Upper Amazon, two from Rio and
one from Blumenau. Length 26-5-30:0 mm. Also one specimen 34:2 mm. long, said to

1918.] F. H. Gravety: Passalidae of the World. 63

come from Queensland. This specimen has hairy mesosternal scars and may belong to
a distinct species.

Passulus punctatissimus differs from P. toriferus chiefly in having all the cephalic
tubercles longer and more acute, and in not having the eighth groove of the elytra hairy
in front. The frontal area may be partially or not at all punctured. The anterior ends
of the marginal grooves of the pronotum are not expanded but may be somewhat curved.
The pronotum is sometimes wider in front than behind, with acutely produced anterior
angles! The outermost rib of the elytra is sometimes hairy throughout instead of only
in its anterior half? The last two variations may perhaps indicate a tendency for the
species to split up into various local races, but more material is needed to settle this.

Passalus unicornis, Saint-Fargeau and Serville.
Passalus unicornis, Saint-Fargeau and Serville, 1825, p. 20.

Six specimens from Guadaloupe, 38:5—42°0 mm. long.

The central tubercle is extremely long and slender, much more so than in the preceding
species, but the other cephalic tubercles are much less prominent and more obtuse than
in that species. The pronotum is unpunctured, except in the uniformly narrow marginal
grooves, and in and close to the scars. The epipleura, shoulders, tips, and eighth and tenth
(usually also the ninth to a less extent) ribs of the elytra are covered with hair-bearing
punctures.

Passalus opacus, n. sp.
Fig. VII, 22, p. 53.

One specimen from Farinas, Bolivia, 39:5 mm. long.

The whole surface of this insect is dull as in P. languidus (Kuwert, 1898, p. 275), from
which it differs in having all the grooves of the elytra much more strongly punctured.
Apart from its dulled surface P. opacus differs from P. unicornis in having the head more
rugose with a much shorter central tubercle, the sides of the pronotum more extensively
punctured, the marginal grooves of the pronotum very broad in front of the scars, the
mesosternal scars indistinct, no hair on the ribs of the elytra above or behind the extreme
anterior part of the tenth, and all the grooves of the elytra much more coarsely punctured,
the punctures in the lateral grooves being transverse.

Passalus interruptus (Linnaeus).

Lucanus interruptus, Linnaeus, 1767, p. 560.3

One or more (often numerous) specimens from each of the following localities :—Texas,
Mexico Guatemala (including one specimen from Escuintla), Honduras (San Pedro
Sula), Nicaragua, Panama (Chiriqui), Columbia, Venezuela (Caracas), Guiana (Demerara,

‚Surinam, Cayenne), Peruvian Amazon, Upper Amazon, Brazil (Amazonas, Pernambuco,

1 This is most marked in the series from the Upper Amazon.

? This is so in the two specimens from the Peruvian Amazon.

® This is the reference usually given, but Linnaeus himself described the species at greater length in 1764 (p. 33)
‘and refers there to yet earlier descriptions. I have been unable to consult thes: and cannot say in which or by
-whom the name interruptus was first introduced.

64 Memoirs of the Indian Museum. [Vor. VII,

Bahia, Petropolis, Rio de Janeiro, Santa Catharina, Blumenau) and Bolivia (Farinas and.
S. Antonia). Length 17-51 mm.

This appears to be a very common, widely distributed and variable species. Large
specimens may be as much as three times as long as small ones, are much more robust-
looking, have the sides of the pronotum unpunctured except in the scars and marginal:
grooves instead of extensively punctured, have the dorsal grooves of the elytra smooth
instead of distinctly punctured and have the lateral grooves moderately strongly instead of
very coarsely punctured.! The mesosternal scars are usually deep, narrow and smooth as
in P. wucorms, but may be larger and more irregular, with or without a few hair-bearing
punctures, or the whole of the sides of the mesosternum may be densely covered with
hair-bearing punctures. The hair on the elytra is usually distributed as in P. opacus, but
may be more extensive as in P. unicornis.

P. interruptus differs from P. opacus in having the surface of the body glossy and the
punctures in the grooves of the elytra much less coarse, the lateral ones not being
transverse. In these characters it resembles P. unicornis, trom which it differs in having a.
much shorter central tubercle.

Passalus glaber, n. sp.
Ho WAU 23}, jos D3},

One specimen without locality record. Length 37 mm.

This species is very like the last, but has only a few hair-bearing punctures on the
elytra, these being confined to the lower parts of the shoulders immediately in front of the
epipleura ; it has the mesosternal scars replaced by broad matt areas which are not sunk
below the general level of the plate ; and it has the metasternum somewhat less extensively
punctured.

Passalus binominatus, Percheron.
Passalus binominatus, Percheron, 1841, pp. 23-24.

One specimen from Santa Catharina, 32 mm. long.

The secondary tubercles on the anterior margin of the head are less widely separated,
and although the frontal ridges are lightly arched the frontal area is more nearly triangular
than semicircular. The central tubercle is obtuse and has no free forwardly directed apex..
There are no coarse punctures on the general surface of the pronotum. The mesosternal
scars are entirely absent. There are only a few coarse punctures on the inner side of the
posterior intermediate areas of the metasternum. In all other respects this species resembles.
P. glaber.

Passalus erosus, Truqui.
Fig. VII, 24, p. 53.

Passalus erosus, Truqui, 1857, p. 268.

One specimen doubtfully recorded from Brazil, and one said (doubtless incorrectly)
to have come from India. Length 35-36 mm.

1 The punctures in the lateral grooves are really of about the same size in all specimens ; consequently they are pro—
portionaliy larger and look much coarser in small than in big ones.

1918.] F. H. Gravety: Passalidae of the World. 65:

This species differs from the last only in the form of the frontal area, which is slightly
longer in proportion to its width and is more or less distinctly grooved in continuation of
the median notch on the anterior margin.

The species of Passalus known to me may be distinguished from one another thus :—

The punctures in the grooves of the elytra extremely coarse,
transverse laterally ; the anterior margin of the head straight
between the outer tubercles ; the lateral areas of the metas-
ternum hairless (small insects only)

IA The punctures in the grooves of the elytra less coarse, not
transverse laterally ; or, the anterior margin of the head
strongly notched in the middle ; or, the lateral areas of the

| metasternum hairy ur os 2. as OB ae 3.
The epipleura hairless de Le Be TE Io MOTI, 405 We
2} The epipleura hairy 3 a 56 2122721005153 992
The anterior margin of the head straight or simply notched
in the middle, without secondary tubercles; the frontal
area large, with the central tubercle about half as far from
the anterior margin as the outer tubercles are from each
other, and with the frontal ridges usually straight and
} diverging at about a right angle ; the central tubercle small,
2 without free apex; the outer tubercles prominent, the inner
tubercles usually situated very near them, always nearer
to them than to the central tubercle ; the lateral areas of
the metasternum, and the epipleura and shoulders of the

elytra, hairless 50 = 00 .. P. punclato-striatus, p. 52.
Not as above .. Mr Se of Ee a 4.
The anterior margin of the head straight between the outer
tubercles ; the clypeus entirely hidden; the central tubercle
smal’ with no trace of free apex ; the inner tubercles usually
much less widely separated than the outer, and when
4 distinct always situated a considerable distance behind them ;
the frontal ridges straight, not arched, meeting in a distinct
though sometimes more or less obtuse angle; the epipleura
usually hairless, when hairy the lateral areas of the
metasternum always hairy also SE on ae sis Se 5.
Not as above .. as Br ae of aXe ste 9.
( The inner and outer tubercles equally widely separated, the
| fomer situated at a considerable distance behind the latter ;
2 the epipleura densely hairy ue 30 .. P. rhodocanthopoides, p. 53.
Not asabove .. si es SE 55 45 56 6.

{ The anterior margin of the head usually somewhat thickened ;
the outer tubercles obsolete, obtuse; the frontal ridges
ending in the inner tubercles, which are very pronounced
: and are situated about half way between the central and
| outer tubercles aa on > P. morio, p. 54.

Not as above oe BG se ei Tex

66 Memoirs oj the Indian Museum.

( The anterior angles of the pronotum produced to form a pair
1) of small but distinctly acute forwardly-directed processes
The anterior angles of the pronotum not so produced

JE.

| Vou.

latijrons, p. 54.

The frontal area small, coarsely and closely punctured in front P. pertyi, p. 54.

| tm frontal area large, smooth or rugose with a few large

punctures in front

The elytra fused, their vertical anterior part lightly convex

The elytra separate, their vertical anterior part not convex

The anterior margin of the head straight between the outer
tubercles or very faintly notched in the middle ; the lateral

a areas of the metasternum hairy
| The anterior margin of the head strongly notched or provided
with secondary tubercles between the outer tubercles; or

the lateral areas of the metasternum hairless .

The frontal area about three times as wide as long, broadly
rounded and not angular behind ; the anterior margin of the
head straight or lightly convex between the outer tubercles

11l< The frontal area about twice as wide as long, or angular
behind ; the anterior margin of the head usually notched
or provided with secondary tubercles between the outer

tubercles

the frontal ridges strong

12 The central tubercle with long and ae free apex; the

frontal ridges weak

The epipleura hairy, the apex of the central tubercle more or
13 less free

The epipleura hairless ; the apex of the central tubercle not free

__( The apex of the central tubercle scarcely free, somewhat blunt
I The apex of the central tubercle distinctly free, sharper

The lateral areas of the metasternum hairy
The lateral areas of the metasternum hairless

- The anterior margin of the head convex between the outer
tubercles, but without median notch or secondary tubercles. .
Not as above

The clypeus to some extent exposed as a more or less vertical
(slightly overhanging) oe between the frons and the
labrum

The clypeus entirely hidden

The lateral areas of the metasternum hairless ; the epipleura
hairy throughout ; the central tubercle without free apex..

Not-as above...

| The central tubercle without very long and slender free apex ;

Ie.
IP,

12,

ae)

IP,

mi ht vy ©

converus, p. DD.

quitensis, p. 55.

affinis, p. 55.

euacadorens.s, p. 56.

catherinae, p. 55.

. recticlypeatus, p. 56.

. curtus, p. 56.
. gracilis, p. 56.

. prominens, p. 57.

. quatemalensis, p. 57.

incertus, P. IT.

WIDE,

10.

11.

16.

15.

14.

18.

1918.]

directed forwards above

The the

somewhat smaller outer tubercles, close behind the outer

inner tubercles large,

side of which they are situated; a pair of well developed
and somewhat widely separated secondary tubercles present
between the outer tubercles; the frontal ridges straight,
meeting in a right or slightly obtuse angle ; the lateral areas
of the
anterior parts of the sides of the elytra densely hairy

metasternum, and the epipleura, shoulders and

Not as above ke + :

“ The central tubercle small, without free apex, normal; the
anterior margin of the head not thickened ; the lateral areas
of the metasternum hairless
Not as above
The middle and hind tibiae armed with at most one small
spine Le :

The middle and hind tibiae more extensively armed

( The pronotum punctured only in the marginal grooves and

\ in and immediately above the scars 5° ;

The sides of the pronotum with an extensive band of

a OS ——— En Se

| punctures above the scars =
\ The grooves of the elytra somewhat finely punctured
(ms grooves of the elytra very coarsely punctured
inner and outer tubercles clearly separate ; the former
never more distant from one another than the latter,
sometimes more or less obsolete ae
The inner and outer tubercles situated close together, ate
together a pair of small oblique ridges on the anterior
margin of the head, with the former tubercles somewhat
more distant from one another than the latter
N The epipleura hairless, or the central tubercle with free apex
5+ The epipleura denselv hairy; the central tubercle without
un free apex © si
The central tubercle without free apex, normal .
The central tubercle with free apex, or more or less columnar
The lateral areas of the metasternum hairy ; a more or less
rudimentary fourth lamella often recognizable on the

a antennae : oc
( The lateral areas of the metasternum hairless Ae
The frontal area fully twice as broad as long; the central
| tubercle decumbent throughout, with long slender free apex :
the lateral areas of the metasternum hairless .. ive

| The frontal area less than twice as broad as long ; the lateral
areas of the metasternum more or less hairy .. BA
The central tubercle strongly elevated at base, massive, with
small decumbent forwardly directed free apex oe
( The central tubercle not having this form Si =

. interstitialis, p.

F. H. GRAVELY : Passalidae vf the World.

58,

. glaberrèsius, p. 58.

. Spinosus, p. 59.

. Spimipes, p. 59.

. spiniger, p. 59.

. polli, p. 62.

. abortivus, p. 60.
. jansoni, p. 60.

. mucronatus, p. 60.

. quadricollis, p. 61.

20.

bo
or

31.
26.

68 Memoirs of the Indian Museum. IMoc avi

The central tubercle erect, more or less columnar, broadly

0 rounded above : ue .. P. oceipitalis, p. 61.
The central tubercle de throughout, oath slender free
apex ys id eh Er TIrRsmasutuss 962%
\ The epipleura densely hairy ue ie re mM ve 32.
| The epipleura hairless un Le an ie 2% 2e 36.

nn general surface glossy ; the punctures in the lateral grooves

of the elytra coarse but scarcely transverse in 5 7. En 33.
The general surface quite dull; the punctures in the ne

grooves of the elytra very coarse, more or less transverse . P. opacus, p. 69.

tubercle without free apex eR se .. BD. toriferus, p. 62.
The frontal area more than twice as broad as long ; or the
central tubercle with free apex a Le We oe 4e 34.

an The central tubercle with long and slender free apex Se ae 2% 35.
The central tubercle not extensively free Fe .. P. interruptus, p. 63.
“The eighth ribs of the elytra hairless and unpunctured

throughout .. P. punctatissimus, p. 62.
The eighth ribs of the anne Sakura and He ec

in front a a ae ws =. Ba unicornis, p. 63:

The frontal area more than twice as broad as long on Ey glaber pas:
36

The frontal area less than twice as broad as long 38 Nie Ae 37.

| The frontal area less than twice as broad as long ; the central
\
i frontal area not grooved in continuation of the median
concavity between the secondary tubercles on the anterior
margin es 3% 56 ie .. P. binominatus, p. 64.
ST\ The frontal area grooved medially in front, in continuation
with the concavity between the secondary tubercles on the

anterior margin 3% de 5s 56) Jes GROSWS, {D> Ot.

Genus PTICHOPUS, Kaup, 1869, p. 27.
Type, Passalus angulatus, Percheron, 1835, pp. 84-86, pl. vi, fig. 5.

Ptichopus angulatus (Percheron).

Passalus angulatus, Percheron, 1835, pp. 84-86, pl. vi, fig. 5
Several specimens from Mexico, Guatemala, Honduras (San Pedro Sula), and Nicaragua.
Length 24-35 mm. ‘These specimens show the species to be a somewhat variable one,
even in the characters used by Kuwert to distinguish others from it; and the validity
of species thus distinguished seems very doubtful. The generic definition, therefore will
probably suffice for the identification of the present form, which is very unlike any other

Passalid known to me.

Subfamily SOLENOCYCLINAE.

The characteristics of this subfamily have been discussed and defined above (pp. 10-1 &)}e
Specimens from Madagascar are characterized by the presence of a pair of more or
less pronounced marginal tubercles immediately on the inner side of the fronto-vertical

1918.] F. H. GrAvELY : Passalidae of the World. 69

suture, no tubercles being developed in this position in specimens from Africa. Kuwert
has, it is true, described from Madagascar one species of each of his otherwise purely African
genera Hrionomus and Didimoides ; but it remains to be seen whether he was right. In
the case of the former genus the sentence “ Der ganze Clypeus in der Breite der ganzen
‘Oberlippe scharf vorgezogen ” suggests, for instance, that studi may belong in reality
to the new genus Malagasalus, established below to receive the only species of
Solenocyclinae known to me in which the clypeus is exposed, and the pair of marginal
tubercles most characteristic of the family as a whole—i.e., those immediately above the
lateral extremities of the clypeus—are entirely absent.

None of the Malagasy genera hitherto described contain very many species ; and the
differences between them may advantageously I think be regarded as specific rather than
generic. I propose, therefore, to unite all of them under the name Solenocyclus.

Some of the African genera appear to be decidedly larger ; but in spite of this I am
unable to find satisfactory characters by which to define more than two. I propose,
therefore, to unite under the name Pentalobus, Kaup, all species with hairy sides to the
metasternum and a more or less distinct tubercle or pair of tubercles or excavation in
the middle of the anterior margin of the head ; and to unite under the name Erionomus»
Kaup, all species in which the sides of the metasternum are hairless and the middle of the
anterior margin of the head is straight.

The genera of Solenocyclinae may now be defined as follows :—

| A more or less distinct pair of marginal tubercles present imme-
diately on the inner side of the fronto-vertical sutures (Mala-

| gasy forms) a TE
Tubercles not present in this position (African forms)

À ( The clypeus exposed aS de = .. Malagasalus, p. 69.
m The clypeus hidden 5 ve ar .. Solenocyclus, p. 70.

The sides of the metasternum hairless ; the anterior margin of
the head more or less distinctly notched in the middle line,
3 or with a median tubercle SE = .. Pentalobus, p. 72.
The sides of the metasternum hairy ; the anterior margin of
the head without any median notch or tubercle .. Erionomus, p. 74.

Genus MALAGASALUS, n. gen.

Type, Malagasalus clypeatus, n. sp.
The clypeus exposed ; a pair of marginal tubercles present immediately on the inner
side of the fronto-vertical sutures ; the sides of the metasternum hairless.

Malagasalus clypeatus, n. sp.
Fig. VIIL, 1, p. 70.

Two specimens from Fenerive, Madagascar. Length 34 mm.

The antennae each have three well developed lamellae. The labrum is concave in
front, convex at the sides, slightly narrower behind than in front. The central tubercle
is strongly elevated above the short parietal ridges, but 1s somewhat obtuse. The frontal
ridges diverge from it in an angle of about 90° ; they extend to the strongly developed

70 Memoirs of the Indian Museum. Vor. VI,

inner tubercles, which are directed obliquely upwards and are in contact anteriorly with
the still larger, but more forwardly directed outer tubercles. The outer tubercles are
separated by a distance which is less than the width of the labrum ; they are equidistant from
each other and from the tubercles on the inner side of the fronto-vertical suture, which are
almost equally large. The clypeus is exposed, but is directed almost vertically downwards.

The pronotum is slightly wider behind than in front, and its anterior angles
are obtuse. Its anterior margin is straight, and its posterior margin lightly convex.
The marginal groove is somewhat widely incomplete in front, and is strongly punctured.
The median groove is strong and complete. The scars are punctured, and there may be a
few punctures in the anterior angles. The surface of the prothorax is closely punctured
and hairy. but the hair is not very long. The mesosternum is smooth and glossy, with
large and deeply impressed roughened scars. The anterior intermediate areas of the
metasternum are somewhat coarsely and sparsely, and the lateral areas more finely
and dense.y punctured and hairy, the latter being, however, to some extent roughened
and hairless behind. A somewhat extensive patch of close, coarse, hairless punctures is
present beside the posterior margin of the posterior intermediate areas, and a band
of finer hair-bearing punctures extends from behind this patch outwards along the
posterior margin to the posterior angles. The elytra are hairless, with the lateral
grooves strongly and the dorsal somewhat more weakly punctured.

0°85 9290 TE
©. SUT Yor Py
BIN

40
os
e

Fie. VIM.
Solenocyclinæ ; specific characters in the upper surface of the head x 4.
1. Malagasalus elypeatus, Gravely. 4. Eriomomus trichostigmoides, Gravely.
2. Solenocyclus exaratus (Klug). 5. Erionomus planiceps (Eshscholtz).

3. Pentalobus punctupectis (Kaup).

Genus SOLENOCYCLUS, Kaup, 1868a, p. 10.
Incl. Oiceronius+Semicyclus, Kaup, 1871. Also Flaminus+ Vitellinus,
Kuwert.

Type, Passalus exaratus, Klug, 1832, p. 173.

Solenocyclus approximatus (Klug).

Passalus approximatus, Klug, 1832, p. 174.
Passalus approximatus, Percheron, 1841, pp. 16-17, pl. Ixxvii, fig. 5.
Six specimens from Madagascar, five of them being from Fenerive. Length 26°5-29:0-

mm.
The clypeus is hidden. The inner tubercles are smaller than in Malagasalus clypeatus,.

and are situated at a greater distance behind the outer tubercles, which are larger.

1918.] F. H. Gravety: Passalidae of the World. hl

Between the outer tubercles is a pair of small and more or less closely approximated
(sometimes fused) secondary marginal tubercles, and another such tubercle is situated
immediately on the outer side of each. The tubercles situated immediately on the inner
side of the fronto-vertical suture are somewhat larger than the two pairs of marginal
tubercles last referred to, and are widely separated from them, but they are less pronounced
than in Malagassalus clypeatus. Coarse punctures are scattered all along the sides of
the pronotum. The metasternum is practically hairless. In all other respects the present
species resembles the preceding one.

Solenocyclus morbillosus (Klug).
Passalus morbillosus, Klug, 1832, p. 175.
Passalus morbillosus, Percheron, 1841, pp. 18-19, pl. Ixxvii, fig. 6.

Eleven specimens from Madagascar, eight being from Fenerive and one (marked
“ Ciceronvus antanarivae, Kuw.’’) from ‘ Antanarivo ” (?=Antananarivo). Also two from
Andakana, belonging to M. Guy Babault. Length 20°5-240 mm. A smaller but more
robust insect than the last.

The pair of secondary marginal tubercles situated next to the outer side of the true
outer tubercles is distinctly larger than the pair situated close to the fronto-vertical suture,
and is situated much nearer to the latter than to the former, which are much larger than
either. The margin is often notched medially, and the notch may be bounded by
an additional pair of minute secondary tubercles. The frontal ridges diverge at right
angles from the low central tubercle and extend direct to the inner tubercles where they
bend more or less abruptly and extend parallel to one another, or slightly converging, to
the outer tubercles. The general surface of the head is smooth and glossy.

The pronotum is strongly punctured at the sides, and as a rule more or less all over
the dorsal surface also. The median groove is strong and complete ; the marginal groove
is somewhat widely incomplete in front. The mesosternum is more or less coarsely rugose,
especially near the lateral sutures ; the scars are ill-defined or absent. The metasternum
and abdominal sterna resemble those of S. approximatus. The grooves of the elytra are
almost uniformly punctured.

Solenocyclus exaratus (Klug).
Fig. VIII, 2.

Passalus exaratus, Klug, 1832, p. 173.
Passalus manouffi, Percheron, 1835, p. 62, pl. iv, fig. 7.

A number of specimens, mostly from Fenerive. Length 28-32 mm.

The outer tubercles, and the three pairs of secondary tubercles on the anterior margin
of the head, are all of about equal size and about equidistant from each other, except that
the secondary tubercles between the outer tubercles are sometimes weaker than the
rest. The central tubercle is very large, with a forwardly directed free apex; the whole
surface of the head in front of it is coarsely rugose ; the inner tubercles are minute, and
are situated close to the base of the central tubercle, and the frontal ridges are absent in
front of them.

The pronotum is without strong punctures, except in the lateral parts of the marginal
groove and in and beside the scars. Both the marginal and the median grooves are

72 Memoirs of the Indian Museum. Nora

complete. In other respects this species resembles the preceding one, except that the:
lateral grooves of the elytra are somewhat more coarsely punctured.

Solenocyclus grayi (Kaup).
Semicyclus grayi, Kaup, 1871, p. 28.

Five specimens from Madagascar, including two from Andakana sent for identification.
by M. Guy Babault. Length 305-360 mm.

The head is smooth and glossy except close behind the anterior margin, where there:
are a few large and more or less coalescent punctures. The outer tubercles, though obtuse,
are distinctly larger than in S. exa atus, and there are no secondary tubercles between
them. The free apex of the central tubercle is much larger than in S. exaratus, and there
is no trace of frontal ridges or inner tubercles.

There are a few punctures in the scars and marginal groove of the pronotum,,
the pronotum being otherwise unpunctured. The median groove is not very deeply
impressed, and neither it nor the marginal groove are complete in front. Along the
lateral margins of the mesosternum there extends a band of hair-bearing punctures, on
the inner side of which the scars are more or less clearly recognizable. The anterior
intermediate and lateral areas of the metasternum, and the posterior border of the:
posterior intermediate areas, are covered with somewhat fine hair-bearing punctures ;
there are no coarse punctures anywhere on the metasternum. The elytra are somewhat
hairy at the shoulders. The puncturing of their grooves is more or less obsolete, at least
dorsally.

Genus PENTALOBUS, Kaup, 1868a, p. 17.
Incl. Didimus, Kaup, 1871. Also Didimordes+ Eumelosomus, Kuwert, 1896.
Type, Passalus barbatus, Fabricius, 1801, p. 256.

Pentalobus klugi (Kaup).
Leptaulax klugii, Kaup, 1868a, p. 12.

Two specimens from Barombi, Cameroons Interior ; one from Abetefi, Ashanti ; one:
from Franceville, French Congo ; and several from Gaboon. Length 22°0-25°5 mm.

Each antenna bears three short lamellae. The anterior margin of the head bears five
more or less equidistant tubercles. The central and inner tubercles are moderately
elevated, and the frontal ridges distinct throughout. The surface of the head is punctured
sparsely behind the central tubercle, and somewhat more densely in front. The sides.
of the pronotum are very broadly and coarsely punctured; the median groove of the
pronotum is complete; the marginal groove is incomplete in front. The mesosternum
is usually more or less matt with somewhat obscure scars. The anterior intermediate:
areas of the metasternum are more or less distinctly punctured ; the lateral areas are
very sharply defined throughout and are enlarged behind ; the posterior intermediate
areas are smooth and glossy. The abdominal sterna are densely and extensively
punctured. The lateral grooves of the elytra are transversely punctured.

1918.] F. H. Gravety: Passalidae of the World. 73

Pentalobus sansibaricus (Harold).
Passalus sansibaricus, Harold, 1880, pp. 262-263.

Five specimens from Zanzibar, one from Abyssinia, one from Abetefi, one from
Rhombomp (Sierra Leone), a number from Dar-es-Salaam, and six said to come from
Bolivia. Length 16°5-27°5 mm.

P. sansibaricus differs from P. klugi only in having the antennal lamellae somewhat
longer, in having the upper surface of the head thickly punctured more or less all over, in
having the mesosternum smooth and polished, with well defined scars, and in having the
posterior intermediate areas of the metasternum coarsely punctured. The central plate of
the metasternum is lightly and sparsely punctured in small specimens.

Pentalobus punctipectus (Kaup).
ne, MID, 33, 10. 10
Leptaulax punciipectus. Kaup, 1868a, p. 11.

One specimen from Gaboon. Length 18 mm.

This species differs from small specimens of the last in having a median pair of secondary
marginal tubercles instead of a single median tubercle, and in having the central plate of
the metasternum less sparsely covered with stronger punctures.

Pentalobus barbatus (Fabricius).
Passalus barbatus, Fabricius, 1801, p. 256.

A large number of specimens from Abetefi, Ashanti; also a few from Amu, Ashanti ;
Gaboon ; Old Calabar ; Aquapim, Guinea; Barombi, Cameroons ; and Angola ; all 23-29
mm. long. Also one specimen from. East Africa, 20 mm. long, and two from Franceville,
French Congo, 17-19 mm. long, between which and the larger forms I am unable to find
any structural difference.

This species differs from P. punctipectis in having five antennal lamellae, all of them
very long and slender ; and in having the central area of the metasternum unpunctured.

Pentalobus parastictus (Imhoff).
Passalus parastictus, Imhoff, 1843, pp. 171-172.

One specimen from Aquapim, Guinea, and one from Barombi, Cameroons Interior.
Length 21-24 mm.

The punctures on the head are somewhat shallow, and the anterior margin of the head
is lightly exavate medially, but has no median tubercles. The punctures on the central
area of the metasternum are more or less concentrated beside the posterior margin. The
transverse punctures of the lateral grooves of the elytra are more or less obsolete in the two
outermost of these grooves, and all the grooves except the fifth and sixth tend to unite in
a matt depression behind. In other respects this species resembles P. punctipectis.

74 Memoirs of the Indian Museum. [Voz. VII,

Pentalobus fur (Kuwert).
Didimus duplicatus ab. fur, Kuwert, 1898, p. 307.

Three specimens from Abetefi, Ashanti. Length 21-23 mm.

It is possible that this form, which Kuwert regarded as an aberration of P. duplicatus
(Har.) may be no more than a variety of P. parastictus. It differs from the latter only in
being without punctures in the posterior angles of the pronotum, in having the punctured
area on the central area of the metasternum less strongly marked, in having the punctures
of the seventh and eighth grooves of the elytra weaker and less distinctly transverse, and
in having the posterior angles of the elytra polished instead of matt. These are much
the same characters as those by which Kuwert distinguishes this form from the form
which he regards as the typical dupiicatus, a species which I have not seen.

The above-mentioned species of Pentalobus may be distinguished as follows :—

( The anterior margin of the head with a strong median tubercle Le ae 2
"| The anterior margin of the head medially concave Le a Bic 3.
The posterior intermediate areas of the metasternun unpunc-
2) tured A ER ne er TENNIS Os Te
Ü the posterior intermediate areas strongly punctured .. P. sansibaricus, p. 73.
( The anterior margin of the head with a strong median concavity
bounded by a pair of strong secondary tubercles . de a de 4.
The anterior margin of the head with a faint median concavity
not bounded by definite tubercles .. : at a aM 5.
Antennae each with three moderately stout leslie a) ae punetypectis, pelos
Antennae each with five long and slender lamellae bo IRS Hao pate:
Elytral grooves 7-8 strongly marked with transverse punctures,
| the posterior angles of the elytra matt De .. BD. parastichus, pP. 13.
Elytral grooves 7-8 more faintly punctured, the posterior
angles of the elytra glossy a ae Bo Ea UT D (es

Genus, ERIONOMUS, Kaup, 1868a, pp. 16-17.
Incl. Calidas+ Epeus+ Eriosternus, Kuwert, 1896.
Type, Passalus planiceps, Eschscholtz, 1829, pp. 22-23.

Erionomus palini (Percheron).
Passalus palinii, Percheron 1844, pp. 8-9, pl. exxxv, fig. 1.
A number of specimens from Abetefi, Ashanti, and two from Gaboon. Length
34'5-38°0 mm.
This species superficially resembles Pentalobus barbatus. The antennae have, however,
only four lamellae ; the anterior margin of the head is approximately straight in the middle ;
the puncturing of the sides of the prothorax is somewhat less extensive ; the lateral areas

of the metasternum are hairy; and the punctures of the lateral grooves of the elytra,
though very strong, are not transverse.

1918.] F. H. GrAvELY : Passalidae of the World. 75

Erionomus trichostigmoides n. sp.
Fig. VIII, 4, p. 70.

Two specimens, one from Dar-es-Salaam, and one said to come from Santa Catharina
in $. E. Brazil. Length 27°5-29'5 mm.

This species bears a close superficial resemblance to species of the Oriental genus
Trichostigmus from which, however, it may readily be distinguished by the structure of
the mandib'es, and of the lateral areas of the metasternum, which resemble those of its
African allies.

The antennal lamellae are extremely short. The surface of the head is polished and
somewhat rugulose ; the frontal area is fully as broad as long, and the inner tubercles are
somewhat indistinct. The pronotum is without strong punctures ; its grooves resemble
those found in other members of the genus. The mesosternum is punctured and hairy
in front, and roughened and hairy behind, with a small smooth and hairless area in the
middle. The metasternum is punctured and hairy, except in the central and posterior
parts of the central area. The punctures in the grooves of the elytra are more or less
obsolete, but the eighth. to tenth ribs are covered with small hair-bearing punctures.

Erionomus alterego (Kuwert).
Eriosternus alterego. Kuwert, 1898, p. 138.

Two specimens from Abetefi, Ashanti. Length 29-31 mm.

The antennal lamellae are very short. The anterior margin of the head bears a more or
less distinct median pair of marginal tubercles, the outer tubercles are somewhat small,
and the secondary tubercles on the outer side of them are broadly truncate. The pronotum
resembles that of E. trichostigmordes, except that the marginal grooves are more strongly
punctured. The smooth and hairless central areas of the mesosternum and metasternum.
are much larger, though the punctures on the latter plate are stronger where they occur.
The elytra are hairless except at the shoulders ; their dorsal grooves are scarcely, their lateral
grooves distinctly but not transversely, punctured.

Erionomus planiceps (Eschscholtz).
Inne, WLU, 6, jos U;
Passalus planiceps, Eschscholtz, 1829, pp. 22-23.

Three specimens from Guinea and two from Abetefi, Ashanti. Length 38°0-41°5 mm.

In this species the outer tubercles are directed upwards instead of forwards, and appear
in consequence to be situated a little behind the anterior margin of the head, beneath.
which, however, dissection shows the true clypeus to be hidden as usual. The marginal
grooves of the pronotum are not strongly punctured. The posterior intermediate areas of
the metasternum bear a posterior marginal band of fine close punctures, the rest of these
areas being smooth much as in Æ. alterego, which the present species also resembles in all
other respects.

The above-mentioned species of Erionomus may be distinguished as follows :—

Each antenna with three short stout lamellae me ae a ar DAL
1b,

Each antenna with four long slender lamellae de a1) palin, pala.

76 Memoirs of the Indian Museum. [Voz. VII,

( The sides of the etytra covered throughout with hair-bearing
9 punctures... = ER or: .. E. trichostigmoides, p. 75.
The sides of the elytra hairless except at the shoulders 7 bie Se 3
( The outer tubercles situated as usual on the anterior margin of
the head and directed forwards "= ae .. Æ. alterego, p. 75.
3) The outer tubercles situated shehtly behind the anterior margin

\ ol the head and directed upwards .. me .. Æ. planiceps, p. 75.

Sub-family MACROLININAE.

As defined above (pp. 12-13) this sub-family includes the Macrolininae, Pleurariinae
Aceraiinae, Gnaphaloeneminae and Tarquintinae of my “ Account of the Oriental Passalidae.”
When that account was written only the Indian and Burmese genera and species were
adequately represented in the Indian Museum collection. Before it was published I was
able to revise to some extent, in the light of a hurried study of the collections in London,
Berlin and Hamburg, my ideas regarding the forms from further east; but the
arrangement of these forms there suggested is, I believe, capable of considerabe
improvement, as indicated in the present paper.

The symmetrical genus Macrolinus has here been placed next to the genus
Pleurarius, which seems to replace it in the Indian Peninsula, 7.e., before, instead of
after, all Oriental genera containing asymmetrical! species.

Kuwert’s Heferochilus wallace: has been removed from the genus Aceraius to the genus
Ophrygonius, where it has been put next to O. birmanicus and O. singapurae, which it
resembles much more closely than it does any species of Aceraius. To permit of this
change the genera Ophrygonius and Aceraius have been redefined, greater importance
being attached to the character of the mandibles than to that of the elytra ; with the result
that Aceraius minor and aequidens of my previous paper have also to be transferred to
Ophrygonius. The remaining species of Aceraius can then be arranged in a single series
leading up from forms allied to minor and aequidens to grandis and occulidens, which
appear to be the most highly specialized members of the genus.

Parapelopides, Trapezochilus, Gnaphalocnemis, Pelopides! and Plesthenus? are entirely
Oriental or Celebean. They resemble the Oriental forms dealt with above in that,

1 Kuwert placed two species, schraderi and gravidus, in this genus (1898, p. 322). Zang, who had not seen either of
them (1905 a, p. 316) pointed out the improbability of their being congeneric (1905 b, p. 227), and suggested that the former
should be regarded as the type of the genus, presumably on account of its probable relationship with the remaining genus
of Kuwert’s group Pelopinae. The material in the Van de Poll collection tends to confirm my opinion (1914 c, p. 201,
footnote 2) that schraderi actually belongs to the genus Protomocoelus ; if,therefore, this species is to be regarded as the
type of the genus Pelopides this name, having priority over Protomocoelus, will probably have to replace it: but there is
little hope of settling the identity of schraderi with certainty without reference to Kuwert’s type. Ido not think, however,
that Zang’s suggestion should be accepted ; for Kuwert, in his first definition of Pelopides (1896, p. 229), gives only
Mindanao as its locality, and this is the locality of gravidus, not ot schraderi. Moreover gravidus was known to him before
schraderi, as it alone is mentioned in his 1891 list, being placed there in the genus Pelops (= Protomocoelus). P. gravidus
and not schraderi should therefore, I think, be regarded as the type. It is represented in the Van de Poll collection by
specimens which appear to have been named by Kuwert himself, and there seems to be no longer any doubt as to its
identity.

2 Boisduval’s lottinii, and Kaup’s quadricornis are, it is true, recorded from “ New Holland.” But itis quite
‚uncertain whether lottinii is a Plesthenus at all (Kaup, 1868 a, p. 26, and 1871, p. 40; Blackburn, 1900, pp. 207-208) ;

1918.] F. H. GRAvELY : Passalidae of the World. 77

whenever asymmetry occurs in the mandibles, the dentition is reduced chiefly on the right

side.
The distinctive characters of the first four of these genera do not appear to me to have

more than specific significance, and I propose to unite them under the name Pelopides.
The fifth genus, Plesthenus, seems to be distinct. It is possible (see below, p. 121) that
these genera may have been derived from Tiberioides, but as this is by no means
certain I have not placed Tiberivides beside them, but have left it next to the symmetrical
species of Episphenus.

Pharochilus, Episphenoides, Mastochilus, Analaches and Cetejus may likewise be united,

the scars on the mentum, with the aid of which they have hitherto been defined, being
variable and not sharply distinctive. The somewhat large genus resulting from this union
is, however, composed of four more or less distinct groups of species, for which four of
the above names may be retained in a subgeneric capacity. Thus Pharochilus may be
-defined so as to include only large Australian species with extremely short antennal
lamellae and more or less extensive matt lateral borders to the mentum. And Æpisphenoides
may advantageously, I think, be re-united with the very few known species of Mastochilus,
and may then be defined so as to include the remaining large symmetrical species.
Analaches and Cetejus are very difficult to separate from Mastochilus on structural
grounds, but contain species of a much smaller size, many of which are distinctly
asymmetrical. Analaches contains somewhat larger and flatter species than Cetejus
with distinctly longer antennal lamellae. According to Heller, who has examined Zang's
‘material (1910, pp. 14 and 21), the upper edge of the left mandible is toothed near the base,
-and this is in agreement with my observations on the few species before me. This tooth
is sometimes, however, situated so near to the base as to be hidden beneath the anterior
-angles of the head. This is so, for instance, in Stoliczka’s australiensis, which Heller
places in the genus Cetejus, in spite of its flattened form and long antennal lamellae, but
which dissection shows to be an Analaches.

Mastochilus (s. lat.) appears to represent the simpler stock from which the remaining
genera, all more highly specialized, have been derived. All these other genera are found
mainly in the islands east of Celebes, and whenever their dentition is reduced this occurs
most markedly on the left side, instead of on the right as in Oriental groups (see pl. I).

Kaupnoloides, Hyperplesthenus, Aurelius, Labienus and Kaupiolus all have symmetrical
mandibles with complete dentition. In the most highly specialized species the elytra are
united, and in all of them the intermediate and lateral areas of the metasternum are fused,

-a fusion which is closely associated with the union of the elytra, tending to follow it in the

- other groups of Passalidae in which it occurs. Probably, therefore, these genera are losing,
if they have not already lost, the power (or at least the habit), of flight, the wings doubtless
becoming more efficient stridulatory structures at the same time. None of these genera
are very large, and they may advantageously be united under the name Labienus.

and in Kaup’s original description of quadricornis (loc. cit.) the only locality referred to is that of the type of Jottinii,

a locality which he has quoted in his monograph, perhaps inadvertently, as that of quadricornis. Even if the type of

quadricornis should prove, on re-examination, to be labelled ‘ New Holland ” I should still doubt the validity of the

record, in view of the extreme improbability of any such highly specialized species, with Oriental rather than Papuan
- affinities, occurring there.

78 Memoirs of the Indian Museum. Vom. WILL.

Protomocoelus resembles Labienus in having the lateral and intermediate areas of the
metasternum fused, though so far as I know the elytra are never united ; but it has the
dentition reduced, especially on the left side. It is probably allied to the asymmetrical
forms of Mastochilus through the more primitive species of Labienus (t.e., trigonophorus

and inaequalis) but does not appear to be allied to the higher forms of the latter
genus.

Gonatas, with which the imperfectly separated Omegarius (? and Tatius of which I have

not seen a specimen) may be united, also tends to have the dentition reduced, especially
on the left side

Pseudepisphenus and Tarquinius form the last line of descent from Mastochilus (s. lat.).
They are so unlike superficially that it seems best to retain both genera in the absence of
other forms throwing light upon them. Concerning their relationship see Gravely, 19140,
PP. 328-320.

The genera of Macrolininae may now be redefined thus :—

The supra-orbital and supra-oceipital ridges discontinuous ;

Insects always symmetrical; the mentum always with

primary but without secondary scars sf .. Macrolinus, p. 80.
1% The supra-orbital and supra-occipital ridges continuous, the
latter sometimes produced outwards behind the former in
asymmetrical species without scars of any kind on the
mentum ee ar ar 5 og ae ae 2.
The mentum without scars; the outer side of the mandibles
angulate at the base or not at all ae oe N: a He om
The mentum with scars; or, the outer side of the mandibles
angulate opposite the anterior lower tooth ne or ne ae ie
Only three lamellae recognizable when the antenna is furled ; the
| mentum strongly grooved in the middle line from front to back Pleurarius, p. 84.
34 More than three lamellae recognizable when the antenna is
furled ; the mentum at most with an incomplete median
groove anteriorly 7 ge 5 1; D Er 4..
{ The inner tubercles separated by a space 3-} as iene as that
ji separating the outer tubercles se 88 .. Tiberioides, p. 84.
. The inner tubercles separated by a space 2-1 times as long as
| that separating the outer tubereles a Ren PAT à À h Br

The left outer tubercle acute and little or no larger than the right ;
or, much larger and curved inwards, with its extremity rounded
rather than truncate and never angular on the outer side in
front. The dentition complete in symmetrical species; the
lowest terminal tooth always present on both sides; the
right anterior lower tooth smaller than the left in the more
EN highly asymmetrical species. The sides of the elytra hairless. Episphenus, p. 85.
The left outer tubercle larger than the right, directed more or less
inwards, truncate distally, the outer angle of truncation dis-
tinct, forming a more or less forwardly directed apex to the tu-
bercle ; or, the sides of the elytra hairy. The dentition complete,
or both the lowest terminal and anterior lower teeth reduced =” aon nO

1918.] F. H. GRAVELY : Passalidae of the World. 79

The dentition complete in unworn specimens on both mandibles ;
the mandibles in most species symmetrical ; the elytra often
hairless except at the shoulders ate Ophrygonius, p. 86.

6< The lowest terminal and anterior lower teeth of the right
mandible absent, or represented only by very minute
denticles ; the anterior lower tooth of the left mandible very

large ; the elytra more or less extensively hairy at the sides Aceraius, p. 89.

The dentition of the left mandible not more reduced than that
of the right and the right outer tubercle at least as large as
the left ; the mentum without primary scars; the secondary
scars very large, more or less completely cutting of the
median from the lateral pieces, usually more or less linear ;
the lateral and intermediate areas of the metasternum never

fused se =i ae 22 Ke an > 8.
Not as above ; the dentition of the right mandible never more

reduced than that of the left Ke ae Er x Er 9,

|
The anterior margin of the labrum with a (frequently indistinct)
tooth near the middle; all the elytral grooves deeply
impressed throughout, at least the lateral ones strongly
{ punctured ; the pronotal scars hairless; at least one of the
outer tubercles more or less complex, or truncate with an
additional tubercle between itself and the anterior angle of
8 the same side of the head (Oriental forms) oe .. Pelopides, p. 93.
The anterior margin of the labrum not toothed ; some at least
of the elytral grooves feebly impressed and more or less
obsolete behind, none of them at all strongly punctured ; the
| pronotal scars more or less hairy ; the outer tubercles acute
| or truncate, but never complex or associated with secondary

tubercles (Celebean forms) Plesthenus, p. 96.

The mentum with well developed primary scars; the lateral

and intermediate areas of the metasternum distinct .. Mastochilus, p. 97.
3 : À :
The mentum without definite primary scars ; or, the lateral and
| intermediate areas of the metasternum fused a a ea 10.
The lateral and intermediate areas of the metasternum fused .. AN En il;
wf The lateral and intermediate areas of the metasternum not
fused de ar ah 12.
The dentition of both mandibles complete, normal; mandibles
more or less symmetrical tie ys .. Labienus, p. 103.
The dentition of both mandibles reduced—especially that of

th left one ; the anterior lower tooth (when present) widely
separated from the middle lower tooth, and partially fused
with the lowest terminal tooth We a .. Protomococlus, p. 107.

1 The fusion is less obvious than in the American forms where it occurs, as the greater part at least of the posterior
-intermediate areas are hairless and either smooth or coarsely punctured, while the areas on the outer side and in front of

“them are very hairy and densely but somewhat finely punctured.

80 Memoirs of the Indian Museum. Wom, NL.

[ The mandibles angulate on the outer side close to the base or
not at all; at most a weak groove extending from this angle
® to the upper margin ae Ss = .. Gonatas, p. 108.
~\ The mandibles angulate on the outer side about opposite the
anterior lower tooth; a strong groove extending from this
angle to the dorsal margin se a bs si a 13.
- The inner tubercles situated behind the anterior margin of the
head ; the outer tubercles asymmetrical ie .. Pseudepisphenus, p. 111.
134 The inner tubercles situated on the anterior margin of the head,
giving the insect a Leptaulax-like appearance; the outer
tubercles symmetrical... 28 aS .. Larqunius, p. 111.

Genus MACROLINUS, Kaup, 1868a, p. 18.
Type, Passalus latipennis, Percheron, 1841, pp. 8-9, pl. Ixxin, fig. 3.

Macrolinus andamanensis (Stoliczka).

Basilianus andamanensis, Stoliczka, 1873. pp. 160-161.

Macrolinus andamanensis, Gravely 1914c, p. 242. pl. xin, figs. 41-41a.
Ten specimens from the Andamans, where Mr. Kemp recently collected two at Port Blair.
Length 32-8-36°5 mm.

Macrolinus sikkimensis (Stoliczka).

Basilianus sikkimensis, Stoliczka, 1873c, pp. 161-162.

Macrolinus sikkimensis, Gravely, 1914c, pp. 243-244, pl. xin, figs. 42-424.
Nine specimens from the Darjiling District (Tukvar, also specimens recently collected

by myself at Pashok, ca. 2,000 ft.), Assam (Margharita) and the Naga Hills, 2,000-5,000 ft.

>, 5
THE, IDG

Macrolinus spp. ; specific characters in the upper surface of the head x 4.
1. M. obesus, Gravely. 2. M. depressus, Gravely.

M. Vitalis de Salvaza has submitted for examination a specimen from Xieng Khaoung,
Tonkin, belonging to the sub-species tavoyanus, Gravely. Length 27'3-32:0 mm.

Macrolinus rotundifrons, Kaup.
Macrolinus rotundifrons, Kaup, 1874, pp. 44-45.
Macrolinus rotundifrons, Gravely, 1914¢, pp. 244-245, pl. xin, fig. 43.
Four specimens from Ceylon, one being from Belihul-Oya. Length 27:0-28-5 mm.

Macrolinus obesus, n. sp.
Fig. IX, 1. 4
Four specimens from Ceylon, three being from Belihul-Oya. Length 29:4-33:4 mm.
The antennal lamellae are very short and stout, even more so than in M. rotundifrons,
the first three being scarcely twice as long as thick, and scarcely more than half as long as
the last three. The ridges and tubercles of the head resemble those of M. rotundifrons,
but the general surface is less extensively punctured and rugose.

1918.] F, H. GraveLy : Passalidae of the World. 81

The pronotum is at most sparsely punctured in the anterior angles and in and near the
scars. The marginal grooves are widely discontinuous in front and somewhat less widely
behind; they are very narrow and are unpunctured except at their anterior ends, which
are slightly enlarged and directed a little backwards from the anterior margm. The median
groove is very fine, and is incomplete in front. The sides and angles are lightly rounded.

The scutellum is smooth and glossy. The mesothoracic epimera are smooth and glossy
antero-dorsally, punctured and glossy below and behind this, then unpunctured and matt,
and finally smooth and glossy along the oblique ventral margin. The mesosternum is
glossy throughout, but is sometimes indistinctly punctured in the more or less rudimentary
scars.

The intermediate and lateral areas of the metasternum are fused and are densely
punctured. They are hairy except on the greater part of the space corresponding to the
posterior intermediate areas, where the punctures are specially coarse. The abdominal
scars are strongly and extensively, but somewhat finely, punctured. The elytra are
united ; they are lightly convex between the shoulders, short in proportion to their length
and lightly convex at the sides, being distinctly broader behind than in front ; the grooves.
are strongly and uniformly punctured, about as strongly as are the lateral grooves of M.
rotundifrons or the dorsal ones of M. crenatipennis.

Macrolinus batesi, Kuwert.

Macrolinus batesi, Kuwert, 1898, p. 187.

Four specimens from Perak, Malay Peninsula, and large numbers from Mana-Riang,
Renau, Palembang, Sumatra, 3,000 ft. and from Bng. Proepoe, Pad. Bovenland (=interior
of Padang), Sumatra, ca. 1,600 ft.. Mr. Holman-Hunt has sent specimens for examination
from the Selangor-Pahang boundary, ca. 3,000 ft., Malay Peninsula ; and M. Guy Babault
from Medan, Sumatra. Length 25-30 mm.

In my “ Account of the Oriental Passalidae ” AZ. batesi was regarded as identical with
M. latipennis. The specimens which I now refer to the former are distinguished by the
relative shortness of the second antennal lamella, whose apex does not fall in line with the
apices of the first and third lamellae when the antennae are furled; by the small
unpunctured frontal area ; by the more or less distinctly shouldered outer tubercles ; by
the large inner tubercles with strongly concave instead of straight or convex connecting
ridge ; by the narrow marginal grooves of the pronotum, which are hairless except below
the scars; by the matt but entirely unpunctured mesosternal scars; and by the coarser
puncturing of the lateral grooves of the elytra.

Macrolinus depressus, n. sp.

Fig. IX, 2.
One specimen from Java, 33 mm. long. Closely allied to M. batesi, but larger and
proportionally flatter. The outer tubercles are strongly shouldered. The pronotum is un-

punctured except in the small round hairless scars, and between these and the posterior
M

89 Memoirs of the Indian Museum. . - |Vor. VII,

halves of the sides, where it is densely hairy. The posterior intermediate areas of the
metasternum and the dorsal grooves of the elytra are unpunctured.

Macrolinus latipennis (Percheron).

Passalus latipennis, Perche on, 1841, pp. 8-9, pl. Ixxin, fig. 3.
Macrolinus latipennis, Gravely, 1914c, pp. 245-246, pl. xin, figs. 45-46.

One specimen from P.Oelak Tanding ; two from Hili Madjedja, N. Nias ; one each from
Bedagei Interior, ca. 600 ft. and Bng. Proepoe, Padang Interior, 1,600 ft., Sumatra ; one
from Tengger Mountain and several from Buitenzorg, Java; two from Mt. Marapok,
Dent Province, Borneo ; and one from Mt. Kina-Balu, Borneo, as well as a few without
definite locality records. Length 22-5-25:2 mm.

Macrolinus sulciperfectus, Kuwert.
Macrolinus sulciperfectus, Kuwert, 1898, p. 184.

One specimen from Toli-Toli, N. Celebes, 26-7 mm. long.

Macrolinus duivenbodei, Kaup.
Macrolinus dwivenbodei, Kaup, 1868 a, p. 19: 1871, p. 43, pl. iv, fie. 6.
Four specimens from Celebes, three being from Menado and one from Loewoe. Length
26-28 mm.
The first two antennal lamellae are more or less distinctly shorter than the remaining
four; but all six lamellae are slenderer than in the Ceylonese species waterhouser &
rotundifrons associated by Kaup (1871) in this respect with the present species.

Macrolinus urus, Heller.
Macrolinus urus, Heller, 1898, pp. 23-24. pl. i, fig. 26.

Numerous specimens from Bua-Kraeng, 5,000 ft., S. Celebes. Length 35°4-41°5 mm.

The first three antennal lamellae are much shorter than the last three.

Although the elytra are united in most specimens’ the lateral and intermediate areas
of the metasternum are distinct. This is also the case in Pleurarius brachyphyllus, from
the Indian Peninsula,” a species in which the elytra are concave between the shoulders and
the wings show little or no trace of reduction. The elytra of the present species are
convex between the shoulders and the wings or always reduced, being intermediate in
form between those of Pleurarius brachyphyllus and those of Macrolinus obesus.

The key given to the identification of the different species of Macrolinus, on pp.
323-24 of my ° Account oi the Oriental Passalidae ” requires considerable modification for
the species which I have now seen for the first time to be included in their proper places,
It may be emended as follows :—

The frontal ridges complete and well developed, the mner

tubercles more or less distinct
The inner tubercles, and the anterior part or whole of the
frontal ridges, obsolete ; species confined to Celebes “8 BE mA 113,

1 In several they are separate, and show no signs of ever having been united.
* See Gravely, 1915, p. 496.

1918.] F. H. GRAVELY : Passalidae of the World.

The six antennal lamellae stout and as a rule not very long,
the first three usually. very short ; species from the Indian
Empire, Indo-China, (?) Siam and Ceylon "

The six antennal lamellae long and slender ; species from the
Malay Peninsula, Sunda Islands, Philippines and Celebes

margin of the head throughout its whole length by a more or
less concave surface; species from the Indian Empire,
Indo-China and (?) Siam .. Bs Pr
The ridge joining the inner tubercles closely approximated
to the anterior margin of the head either in the middle or
throughout ; species from Ceylon .. = ‘ie
“ The lateral grooves of the elytra. narrow, their punctures
normal an : fs tes ae:

4
The lateral grooves of: mi elytra. broad, their punctures

| The ridge joining the inner tubercles separated from the anterior
| transversely linear : ie ie
The outer tubercles slender in profile, truncate ; the ridge
| joining the inner tubercles concave
| The outer tubercles stouter and distinctly bifid in ae she
ridge joiming the inner tubercles straight ;
The ridge between the inner tubercles straight throughout
almost its whole lensth ; the anterior angles of the pronotum
unpunctured ; the antennal lamellae somewhat long
The ridge between the inner tubercles convex, evenly curved
throughout ; the anterior angles of the pronotum strongly
| punctured ; the antennal lamellae short
\

The lateral grooves of the elytra almost as broad as the
intervening ridges, their punctures very coarse indeed
The lateral grooves of the elytra much narrower than the
intervening ridges : 53 = Sr
The elytra separate, parallel-sided, with finely punctured
grooves xg a
The elytra united, more ovate, with coarsely punctured grooves
The frontal area small and unpunctured ; the inner tubercles
large and connected by a strongly concave ridge ; the marginal
grooves of the pronotum hairless in the anterior angles
The frontal area large and punctured all over ; the inner
tubercles small, and connected by a ridge which is straight as
a whole and situated immediately above the anterior margin
of the head, and may have a more or less angular median
convexity ; the marginäl grooves of the pronotum hairless
only in their anterior terminal enlargements
( The anterior angles of the pronotum strongly and ince
punctured ; the pronotal scars with a few small hairs
The anterior angles of the pronotum with not more than one
or two punctures ; the pronotal scars thickly hairy

M. sikkimensis, p. 80.

M. nicobaricus, Gravely.

M. andamanensis, p. 80.

M. waterhousei, Kaup.

M. crenatipennis, Kuwert.

M. rotundifrons, p. 80.

M. obesus, p. 80.

M. batesi, p. 81.

M. depressus, p. 81.

108

84 Memoirs of the Indian Museum. [Vor. VII,

The third lamella of the antennae not distinctly shorter than
the fourth ; the median groove of the pronotum obsolete .. he Fie 12.
114 The third lamella of the antennae distinctly shorter than the
fourth; the median groove of the pronotum distinct,
complete ore ar a se . M. sulciperfectus, p. 82.

The tip of the second lamella not reaching the line joining the
tips of the first and third lamellae when the antenna is
12 turled Bis ne se ues .. M. weberi, Kaup.
The tips of all six lamellae arranged in a straight line when the
antenna 1s furled oP ae oe .. WM. latipennis, p. 82.

/ The apex of the central tubercle acute, directed forwards,
somewhat overhanging .. if ne .. M. duivenbodei, p. 82.
The apex of the central tubercle approximately rectangular in
profile, directed upwards, not overhanging ae M urus, p. 82,

Genus PLEURARIUS, Kaup, i868), p. 1.
Type, Pleurarius pilipes, Kaup, 18685, pp. 1-2.

Pleurarius brachyphyllus, Stoliczka.
Pleurarius brachyphyllus, Stoliczka, 1873, pp. 152-153.

This species is not represented in the Van de Poll collection.

Since compiling the list of localities in my “ Account of the Oriental Passalidae ” I have
collected specimens in Cochin at Kavalai, ca. 1,300-3,000 ft., and between miles 10 and 14
on the State Forest Tramway, 0-300 ft. M. Guy Babault has presented specimens from near
Mahé on the Malabar Coast, and from Kodaikanal in the Palni Hills ; he has also sent for
examination specimens from Wallardi in Travancore and from the Coorg region.

It is doubtful whether P. pilipes, the only other species of the genus described, is really
distinct from P. brachyphyllus, although the former is supposed to come from Sumatra and
not from India (see Gravely, 1914¢, p. 320).

Genus TIBERIOIDES, Gravely, 1913, p. 405.
Type, Tiherius kuwerti, Arrow, 1906, p. 446.

Tiberioides kuwerti (Arrow).
lel IE

Tiberius kuwerti, Arrow, 1906, p. 446.
Tiberioides kuwerti, Gravely, 1914c, pp. 215-216, pl. xi, fig. 14.

Two specimens from Darjiling and one from N. Manipur, 3,000-9,000 ft. Length
38°5-39°5 mm. RE

1918, F. H. Gravety: Passalidae of the World. 85

Tiberioides borealis (Arrow).
Fig. X.
Chilomazus borealis, Arrow, 1906, pp. 467-468.

One specimen from N. Manipur, 3,000-9,000 ft. Length 38 mm.

Fie. X.
Tiberioides borealis, Arrow, mentum X 8.

The species of Tiberioides may be identified thus :—

“The lateral grooves of the elytra broad, their punctures trans-
versely linear Ke = a .. I. kuwerti, p. 84.
The lateral grooves of the elytra narrow, their punctures
normal on N na aie te .. . 2.

The median part of the mentum without depressions or ridges T. austeni, Gravely.
The median part of the mentum with a low convexity flanked

by broad shallow depressions close to the middle of the

anterior margin, with a strong transverse ridge behind it .. T. borealis, p. 85.

Genus EPISPHENUS, Kaup, 1871, p. 45.
Type, Episphenus moorei, Kaup, 1871, p. 45.

Episphenus moorei, Kaup.
Pls af,
Episphenus moorei, Kaup, 1871, p. 45.
Episphenus moore: + pearson, Gravely, 1914c, pp. 217-218, pl. xi, figs. 16-17.
Eight specimens from Ceylon (Belihul-Oya and Colombo) and, one said to be from the
Himalayas. Length 30:7-34°2 mm.

Episphenus comptoni (Kaup).
Pie
Aceraius Comptom, Kaup, 18682, p. 28.
Episphenus compton + var. flachı, Gravely, 1914c, pp. 218-219, pl. xi, figs. 18-19a.
Eleven specimens from Ceylon, including four from Belihul-Oya and one recently
collected by Mr. Kemp on Horton Plains. Length 29-3-42°0 mm.

86 Memoirs of the Indian Museum. [Voz. VII;

Episphenus neelgherriensis (Percheron).
IP; I.

Passalus neelgherriensis, Percheron, 1841, p. 4, pl. Ixxvii, fig. 1.
Episphenus neelgherriensis, Gravely, 1914c, pp. 222-223, pl. xi, figs. 21-21a.

Five specimens from India, one being from Cochin and one from the Madras Presidency.
Also specimens presented by Mr. T. Bainbrigge Fletcher from the Nilgiris (Ootacamund),
Mysore (Bababudin Hills, 4,500 ft.) and Coorg (Mercara, Santi Koppa and Pollibeta) ; by
M. Guy Babault from Wallardi in Travancore and from the neighbourhood of Trichinopoli
and Mahé ; and by myself from Cochin (Kavalai, ca. 1,300-3,000). Length 23:3-28:5 mm..

Epishenus indicus (Stoliczka), 1873, pp. 159-160.
Pie
Basilianus indicus, Stoliczka, 1873, pp. 159-160.
Episphenus indicus, Gravely, 1914c, pp. 220-222, pl. xi, figs. 20-200.

Hight specimens, one from the Madras Presidency, one from Mercara, Coorg, and six
said to be from Assam. Also specimens presented by M. Guy Babault from Wallardi in
Travancore, and from near Mahé on the Malabar Coast; by Mr. T. Bainbrigge Fletcher
from Santi Koppa in Coorg ; and by myself from IEavelaı, 1,300-3,000 ft., in Cochin..
Length 27-2-36-2 mm. |

The species of Episphenus may be identified thus :—

The anterior margin of the head symmetrical, the nor
margin of the mentum not depressed or grooved .. E. moorei, p. 85.
14 The anterior margin of the head more or less asymmetrical ;
the anterior margin of the mentum more or less depressed or
grooved ac a 30 ne ae oa 59 2.
The anterior margin of the head not very strongly asymmetrical
| as a rule; the anterior margin of the mentum strongly
grooved on either side of a strong median tubercle (occasion-
ally paired) .. 5 = Le .. E. comptont, p. 85.
[m anterior margin of the head strongly asymmetrical ; the
mentum without any strongly marked tuberele ie a 5 D.
The anterior angles of the head not prominent .. .. E. neelgherriensis, p. 86.
3 The anterior angles of the head more or less prominent .. E. indicus, p. 86.

Genus OPHRYGONIUS, Zang, 19044, pp. 697-700.
Incl. Heterochilus, Kuwert, 1896 (preoccupied)—Rhipsaspis, Zang, 1905.

Type, Ophrygonius quadrifer, Zang, loc. cit.=Passalus inaequalus, Burmeister, 1847,
p. 468. |
Ophrygonius cantori (Percheron).
Passalus cantori, Percheron, 1844, pp. 3-4, pl. exxxiv, fig. 2
Ophrygonius cantori, +subspp. convexifrons and dunsiriensis, Gravely, 1914c, pp. 224-295, pl.
xi, figs. 22-220.
Numerous Himalayan specimens, including three from Tukvar ; one specimen from.
N. Manipur, 3,000-9,000 ft. ; and one from the Ruby Mines District of Upper Burma,
5,000-7,000 ft. Specimens have been presented by M. Guy Babault from Kulu (Kandi.

1918.] F. H. GRAVELY : Passalidae of the World. 87

and Mandi); by Mr. C. Beeson from Kumaon (Ramgarh, 6,000 ft., under bark of dead
oak); by Dr. B. L. Chaudhuri and myself from the Darjiling District (Senchal, ca. 8,000 ft.,
and Pashok 5,500 ft.) ; by Mr. 8S. W. Kemp from the Khasi Hills (Shillong 6,400 ft., and
Maflong, 5,900 ft.) ; and by M. Vitalis de Salvaza from Tonkin (Lao Kay).

One specimen labelled “ Darjeeling ” in the Van de Poll collection is only 28-5 mm.
long. All the remaining Himalayan specimens are 30 mm. long or over. The Manipur
specimen is 30 mm. long; all the remaining specimens found east of the Darjiling District
are 29 mm. long or under. Although, therefore, occasional specimens may transgress the
normal limits of the race characteristic of their locality, the validity of difference in size as
a distinction between O. cantori, s. str., and its sub-species convexifrons is confirmed. The
form of mentum, on the other hand, described as distinctive of the sub-species dunsiriensis,
does not appear to be a good character, and although specimens from the Dunsiri valley
range from 28 to as much as 31 mm. in length I think that they can best be regarded as
belonging to the sub-species convexifrons.

Ophrygonius birmanicus, Gravely.
Ophrygonius birmanicus, Gravely, 1914e, p. 226. text-fig. 3A.

One specimen from the Ruby Mines District of Upper Burma. Another has been
presented by Mr. J. Coggin Brown from Man Lom, Homang, N. Tawngpeng, N. Shan States,
Upper Burma, 4,500-5,500 ft. ; and others by M. Vitalis de Salvaza from Chapa, Ht. Tonkin,
Length 29-33°5 mm.

Ophrygonius singapurae, Gravely.
Ophrygonius singapurae, Gravely, 1914c, pp. 226-227, text-fig. 3B.
One specimen from Laos, and two from Mt. Marapok, Dent Province, British North
Borneo. Length 30°5-32°2 mm.
In fresh specimens the anterior lower teeth each have their apices produced into a
sharp and slender point. The left outer tubercle is distinctly thicker at the base than in
the preceding species, but may point more definitely inwards than in the type specimen.

Ophrygonius wallacei (Kuwert).

Heterochilus Wallacei + crinitus + oculitesselatus, Kuwert, 1898, p. 334.
Aceraius wallacei, Gravely, 1914c, pp. 228-229, pl. xu, figs. 26-266.

One specimen from Singapore and a number from Borneo (Mts. Kinabalu and Dispo
Length 33:5-38:0 mm.

The anterior lower tooth on each side is much broader than the lowest terminal. Both
are small on the right mandible, but are perfectly distinct in fresh specimens.

Ophrygonius inaequalis (Burmeister).

Passalus inaequalis, Burmeister, 1847, p. 468.
Ophrygonrus inaequalis, Gravely, 1914c, pp. 227-228, pl. xu, figs. 24-24a.

A number of specimens from Mt..Marapok, Dent Province, British North Borneo :
and one from Mt. Kinabalu. Length 23-27 mm.

The punctures in the lateral grooves of the elytra are variable, and 1 no longer think it
at all likely that oroleius, Smith, will have to be recognized as a distinct variety.

88 Memoirs of the Indian Museum. [Vor. VII,

Ophrygonius aequalis, n. sp.
Is RÜE, IL,

A number of specimens from Chapa, Tonkin, have been sent by M. Vitalis de Salvaza.
They vary from 27:3-31°6 mm. in length, and differ from O. minor only in their larger size,
in having only four antennal lamellae pubescent, in having the upper tooth of both
mandibles well developed, in having the left outer tubercle still more nearly identical in

eo
a RR

Fic. XI.
1. Ophrygonius aequalis, Gravely, part of 3. Ophrygonius javanus, Gravely, left mandible
upper surface of head x 4. from without, highly magnified.
2. Ophrygonius javanus, Gravely, part of 4. Aceraius lamellidens, Gravely, left mandible
upper surface of head x 4. from without, highly magnified.

size and form with the right, in having the frontal area shorter and distinctly transverse,
and in having the punctures on the sides of the pronotum densely clustered in and
confined to the scars, instead of very thinly scattered over a slightly more extended area.
Ophrygonius javensis, n. sp.
Fig. XI, 2-3.

One specimen from Boeloe Lawang, Pasoeroean, Java. Length 26 mm. Closely allied
to O. minor, from which it differs only in having somewhat better developed upper teeth,
and in having the punctures and hair on the elytra confined to the ninth and extreme
hinder end of the seventh ribs, instead of over the whole of these two ribs.

Ophrygonius aequidens (Gravely).
Aceraius aequidens, Gravely, 1914c, p. 240, text-fig. 4E., p. 234.
Six specimens from Mt. Kinabalu. Length 27-31 mm.

The species of Ophrygonius as re-defined in the present paper may be recognized as.
follows :—
The antennal Jamellae short and stout ; the anterior marginal
depressions of the mentum quite small; the sides of the
elytra hairy .. NR ue as .. OÖ. cantori, p. 86.

The antennal lamellae long and slender; or, the anterior

marginal depressions of the mentum very large ; or, the sides

of the elytra hairy a sig 0 56 a5 50 ae
The antennal lamellae long and slender a: 56 ac 00 3.
© { The antennal lamellae short and stout as ae Go 5.

The tips of the first two lamellae not falling in line with those
of the last four when the antenna is furled ; the anterior
marginal depressions of the mentum quite small ; the sides

3 ot the elytra hairless ER an 2 .. O.birmanicus, p. 87.
The tips of all six lamellae forming a straight line when the
antenna is furled ; or the sides of the elytra hairless. The

anterior marginal depressions of the mentum usually large a oc 4.

1918.] F. H. GRAVELY : Passalidae of the World. 89

The tips of all six lamellae forming a straight line when the
antenna is furled ; the anterior marginal depressions of the
mentum always large and separate ; the sides of the elytra
| hairless oh crs = ae .. O. singapurae, p. 87.
*\ The tips of the first two lamellae not falling in line with those
of the other four when the antenna is furled; the anterior
| marginal depressions of the mentum very variable, often fused
or rudimentary ; the sides of the elytra densely hairy in front O. wallacei, p. 87.
The anterior marginal depressions of the mentum very large :
the sides of the elytra hairless we PE .. O. inaequalis, p. 87.
“ } The anterior marginal depressions of the mentum more or less
rudimentary ; the sides of the elytra hairy ae SE De ce 6.
The upper tooth of both mandibles well developed, more or

less rectangular ; the frontal area markedly transverse .. O. aequalis, p. 88.

more obtuse ; the frontal area about as long as broad
The anterior lower tooth of the left mandible very large, much

larger than that of the right mandible
1 The anterior lower tooth of the left mandible scarcely larger
than that of the right a 5. .. O. aequidens, p. 88.
The upper tooth of the left mandible obtuse but moderately
large ; the seventh ribs of the elytra almost unpunctured .. QO. javensis, p. 88.

: The upper tooth of the left mandible minute ; the seventh ribs of

(
|
| The upper tooth of both mandibles rudimentary or at least
f
| the elytra sparsely punctured and hairy throughout .. O. minor (Gravely).

Genus ACERAIUS, Kaup, 1868a, pp. 26-27.
Type, Passalus grandis, Burmeister, 1847, p. 463.

Aceraius lamellatus n. sp.

Mr. Holman-Hunt has submitted two specimens from the Malay Peninsula (Ulu
Gombak), and Mr. Bryant two from Penang, one of each of which has been presented to the
Indian Museum collection. Length 23 mm.

This species appears to be allied to various species of the genus Ophrygonius on the oue
hand, and to the genus Acerarus on the other. The antenna! lamellae and mentum resemble
those of Ophrygomus singapurae, the former being exceptionally large for so small an
insect. The mandibles are of the Aceraius type, and resemble those of A. helferi except that
the convexity of the posterior part of the upper margin is pressed further back from the small
upper tooth. In other respects this species resembles Ophrygonius minor, except that the
eighth ribs of the elytra are punctured instead of (or as well as) the seventh.

Aceraius helferi, Kuwert.

JPL, IE,

Aceraius helferi, Kuwert, 1898, pp. 346-347.
Acermus helferi + tavoyanus + assamensis + himalayensis, Gravely, 1914c, pp. 236-238, pl. xii,
figs. 36-39a.
One specimen from Burma and four from Siam, three of the latter being from Renong.
M. Vitalis de Salvaza has presented a fine series of this species from Tonkin (Chapa
and Xieng Khouang), and has sent for examination specimens from Ban Tink and Lao
N

90 Memoirs of the Indian Museum. [Voz. VIT,

Kay in the same country, and from Cambodia. Mr. J. Coggin Brown has presented
specimens from Loi Tawng Kyaw 5,500-7,000 ft., Man Lom, Hamang 4,500-5,500 ft., and
between Man Lom and Man Hpat, 4,500-5,500 ft., all in Tawnpeng in the Northern Shan
States, Upper Burma. Mr. Holman-Hunt has sent for examination a specimen said to
come from the Malay Peninsula (Ulu Gombak) and M. Guy Babault one said to come from
Sumatra (Médan). Length 29:3-37:2 mm.

I have already pointed out (19146, p. 292 footnote) that À. tavoyanus, from Southern
Tenasserim, is not really distinct from A. helferi from further north. The series trom
Tonkin shows in addition that A. assamensis, and A. himalayensis are no more than
imperfectly separated local races of the same form. Specimens from northerly localities
(see pl. 1, ‘“ Aceraxus, other spp.) ordinarily have much squarer and less slender left outer
tubercles than have specimens from further south. But the shape of this tubercle is not
altogether constant in specimens from a single locality ; and the gradation of the southern
form into the northern seems to be so complete as to render impossible the separation of the
species even into two definite races.

Aceraius alutaceosternus, Kuwert.
Aceraius alutaceosternus, Kuwert, 1898, pp. 347-348.
Aceraius alutaceosternus, Gravely, 1914c, p. 236, text-fig. 4E, pl. xi, figs. 34-34a.

One specimen from Perak and one from Borneo. Mr. C. Holman-Hunt has presented
a specimen from Bukit Kutu in the Malay Peninsula. Length 35-4—39:5 mm.

The shape of the left outer tubercle of this species is the same as that of the most
extreme form found among southern specimens of the preceding, of which it is little more
than a local race. The size of the posterior convexity of the upper margin of the left
mandible, and the faint median groove on the anterior part of the mentum, appear, however,
to afford constant distinctive characters by which it may be recognized.

Aceraius borneanus, Kaup.
Aceraius borneanus, Kaup, 1871, p. 52.
Aceraius borneanus, Gravely, 1914c, pp. 238-239, pl. xi, figs. 25-25b.

One specimen from the Malay Peninsula (Pahang), four from Sumatra (three being
from the interior of Bedagei on the east coast, ca. 600 ft.), many from North Nias (Hili
Madjedja and G. Madjeja), five from Middle Nias (Dyma and Kalim Bungo), one from Java,
and many from Borneo (Doesonlanden, Brunei, and Mts. Marapok and Kinabalu). Mr.
C. Holman-Hunt has presented a specimen from Ulu Gombak, Malay Peninsula. Length
23°0-29°2 mm.

This species appears to be somewhat rare in the Malay Peninsula and Java, but to be
abundant in Sumatra and Borneo.

Aceraius pilifer (Percheron).

Passalus pilifer, Percheron, 1835, pp. 23-24, pl. 11, fig. 2.
Aceraius pilifer, Gravely, 1914c, pp. 235-236, pl. xii, fig. 35.

Numerous specimens from the following places in Java: Tjı Bodas, ca. 4,000 ft. ;
Pengalengan, 4,000 ft. ; Mt. Tjikorai, 4,000 ft.; Mt. Gede, 4,coo ft.: Telaga Bodas

1918. | F. H. GRAVELY : Passalidae of the World. OI

Garoet Preanger, 4,000-5,000 ft. ;: G. Tjı Salimar, W. Preanger, ca. 3,000 ft. ; Boeloe
Lawang Res. Pasoeroean. Length 27-32 mm.

This species appears to be very common in Java, where it probably replaces the
preceding one. It seems to be rare in Sumatra and Borneo. It has not been recorded
from the Malay Peninsula.

Aceraius perakensis, Kuwert.

Aceraius perakensis, Kuwert, 1898, p. 308.
Aceraius perakensis + laevimargo, Gravely, 1914c, pp. 229 and 235.

A number of specimens from Mt. Kinabalu. M. Guy Babault has presented a
specimen from near Dolok-Baros, Médan, Sumatra. Length 34-40 mm.

A. laevimargo, Zang, appears to be identical with the species which I previously
separated as A. perakensis.

Aceraius tricornis, Zang.

Aceraius tricornis + kuwerti, Zang, 1903a, p. 339.
Aceraius tricornis + kuwerti, Gravely, 1914c, p. 235, text-tig. 4 B, pl. xi, fig. 31.

Two specimens from Mt. Maropok, and a number from Mt. Kinabalu. Length 46°0-
525 mm.

The characters by which A. tricornis is separated from A. kuwerti prove to be variable,
and the second name must fall.

Aceraius laniger, Zang.
Aceraius laniger, Zang, 1905a, pp. 191-192.
Aceraius laniger, Gravely, 1914c, p. 234.
Three specimens from Borneo, two being from Mt. Kinabalu. Length 54°0-56°5 mm.
The characters distinguishing this species from the last, small though they are, do not
appear to vary.
Aceraius moschleri, Kuwert.

Aceraius möschleri, Kuwert, 1898, p. 344.
Aceraius möschleri, Gravely, 1914c, pp. 229-230, pl. xu, fig. 33.

Four specimens from Mt. Kinabalu in Borneo. Length 35-37 mm.

Aceraius illegalis, Kuwert.

Acerarus illegalis, Kuwert, 1898, p. 345.
Aceraius illegalis, Gravely, 1914c, p. 230, pl. xii, figs. 32-32a.

Nine specimens from Borneo (Mts. Kinabalu and Marapok). Length 40-43 mm.

Aceraius laevicollis (Illiger).
Passalus laevicoliis, Uliger, 1800, p. 103.
Aceraius laevicollis, Gravely, 1914c, pp. 230-231, pl. xii, figs. 27-27a.
Many specimens from the Malay Peninsula (Singapore, Larut, Penang, Perak) ; Sumatra
(Bedagei Interior, ca. 600 ft. ; Tandjong-Djati, Ranau, Palembang, ca. 2,000 ft.; 8. E.
Serdang, E. Coast, ca. 1,000 ft.); N. Nias (Hili Madjedja and G. Madjeja) ; Middle Nias

N 2

92 Memoirs of the Indian Museum. [Vor. VII,

(Dyma) ; Java; Borneo (Mts. Marapok and Kinabalu, British N. Borneo, Riam Kanan and
Pengaron, Martapoera) ; Bali; and S. Celebes (Tjamba). Length 30'7-37'3 mm.

Aceraius grandis (Burmeister).
ell, Jt,
Passalus grandis, Burmeister, 1847, p. 463.
Aceraius grandis + var. rectidens + subsp. hirsutus, Gravely, 1914c, pp. 231-233, pl. xu, figs. 28-30.

Specimens of the typical form from Sumatra (Mana-Riang, Renau, Palembang. 2.000-
3,000 ft.; Bng. Proepoe, interior of Padung, ca. 1,600 ft. ; also specimens presented by
M. Guy Babault from Médan) ; North Nias (Hil Madjedja and G. Madjeja) ; Middle Nias
(Kalim Bungo and Dyma); Java (G. Tji Salimar, W. Preanger, +3,000 ft. ; Kawie
Mountains, Pasoeroean ; Tji Bodas, ca. 4,000 ft.) and Borneo (Mt. Kinabalu and Brunei).
Length 40-55 mm.

Specimens of subspecies hirsutus, Kuwert, from Nepal; Darjiling District (Tukvar);
Assam (Margherita ; Chandkhira, Sylhet) ; Upper Burma (Cachin Cauri) ; Laos ; Tonkin
(Xieng Khouang, Hoabink, Chapa and Napé—all presented by M. Vitalis de Salvaza, who
has also submitted one from Ban Tink for examination) and 8. Palawan. Length 36°8-
49°0 mm.

The examination of the above recorded specimens has shown that southern specimens
must all be regarded as belonging to a single race. Neither of the names rec'idens and
addendus (see Gravely, 1914¢, p. 322 footnote) need, therefore, be retained.

Aceraius occulidens, Zang.
Aceraius occulidens, Zang, 1905a, pp. 190-191.
Aceraius occrhdens, Gravely, 1914c, p. 234, text-fig. 4A.
Specimens from the Malay Peninsula have been presented by Mr. Holman-Hunt, who
has also submitted one for examination from Gap, 2,700-3,000 ft., on the Selangor-Pahang
boundary. Length 45°5 mm.

The species of Aceraius, as re-defined in this paper, may be identified as follows :—

The upper tooth of the left mandible simple, set in a hollow in
front of a convexity of the upper margin, from which it is
distinctly separated at base ar ae ae aS 36 2.

14 The upper margin without a convexity behind and distinct

from the upper tooth ; this margin concave or straight, or

else uniformly convex the whole way from the tip of the

upper tooth backwards ; or, the upper tooth bifid a 55
The six antennal lamellae exceptionally long and slender . A. lamellatus, p. 89.
|

24 The antennal lamellae short and stout, the first two more or
less rudimentary he 4e 50 30

The posterior pars oi the tenth and the whole of the eighth ribs

j of the elytra unpunctured 3e an Je 56 . Le

? The seventh to tenth ribs of the elytra (inclusive) punctured

throughout .. 52 ER Br . A. borneanus, p. 90.

1918.]

“The convexity of the upper
moderately high; the mentum with no trace of a median
groove

margin of the left mandible

The convexity ae the upper margin a the left: tr very
pronounced ; the mentum with a fine groove in front

“ The anterior angles of the head more or less obtuse, never
prominent 36 =

The anterior angles of the nee sharper, more or je as
prominent -

en most 32 mm. long ; ile shorter

in least 34 mm. long, usually much longer En

At most 40 mm. long; the right outer tubercle distinct and
more or less sharply pointed in unworn specimens, the
antennal lamellae and other characters very variable

At least 43 mm. long; the oe outer tubercle more obtuse or
absent ae a:

The right outer tubercle more or less leer the tenth ribs
of the elytra hairless and unpunctured throughout 5

The right outer tubercle distinct ; the tenth ribs of the elytra

punctured and hairy anteriorly

The right outer tubercle usually shorter, truncate or rounded ;
sometimes, however, divided into two separate tubercles of
which the outer one or both may be more or less long and
acute

‘ The anterior à of the ed eaveoly, ones the tip
the right outer tubercle not bent outwards de

10 The anterior angles of the head distinctly prominent ; the tip
of the right outer tubercle bent more or less abruptly
outwards se se 5c

At most 38 mm. long ; the anterior angles of the head scarcely
more prominent chan’ in A. illegalis, somewhat variable

Specimens no larger than A. laevicollis with the left anterior

angle of the head produced into a long slender inwardly curved
process ; larger ones (which may be as much as 55 mm. in
length)

largest closely resembling A. laevicollis in form

with the angles more moderately prominent, the

The canthus without any upwardly directed tubercle
A small and stout erect tubercle arising trom the dorsal surface

|
The right outer tubercle simple, acute and eh long
|

of the canthus immediately in front of the eye

Ale

4
Jal

Ar.

F. H. Gravety: Passalidae of the World.

. helferi, p. 89.

alutaceosternus, p. 90.

pilifer, p. 90.

perakensis, p. 91.

tricornis, p. 91.

laniger, p. 91.

moschleri, p. 91.

illegalis, p. 91.

laevicollis, p. 91.

grandis, p. 92.

occulidens, p. 92.

Genus PELOPIDES, Kuwert. 1896, p. 220.

Incl. @naphalocnemis, Heller, 1900 (=

Also Trapezockilus, Zang, 1995 (=
Zang, 19044.

Type, Pelopides gravidus,! Kuwert, 1398, p. 322.

1 See above, p. 76, footnote 1.

Eriocnemis, Kaup, 1868, preoccupied).
Phraortes, Kuwert, 1896, preoccupied) + Parapelopides ,

10.

Il.

94 Memoirs of the indian Museum. [Voz. VII,

Pelopides symmetricus (Zang).
PPT
Parapelopides symmetricus, Zang, 1904a, pp. 695-697, figs. 1-2.
Parapelopides symmetricus, Gravely, 1914c, pp. 246-247, text-figs. 6A-B.
Two specimens from Mt. Marapok, and a number from Mt. Kinabalu. Length 37'0-

42°7 mm.
Pelopides gravidus Kuwert.

Fig. XII.
Pelopides gravidus, Kuwert, 1898, p. 322.
Five specimens, apparently cotypes, from Davao, Mindanao. Length 43-46 mm.
P. gravidus is to some extent transitional between P. symmetricus and P. simplex.
The anterior lower tooth of the right mandible is distinct when unworn, but is smaller than
in P symmetricus. The head has both the outer tubercles broad as in P. symmetricus,

Fig. XII.

Pelopides gravidus, Kuwert ; head from above x 4.
but the inner portion of the right one is distinctiy more prominent than that of the
left, while the left one is flanked by a very deep and the right by a shallower concavity.
The elytra are very like those of P. monticulosus. In all other respects P. gravidus
resembles P. symmetricus.
Pelopides dorsalis (Kaup).

IPM, IL.
Eriocnemis dorsalis, Kaup, 1871, p. 41.
Trapezochilus nobilis,’ + respectabilis, Gravely, 1914c, pp. 247-248, text-fig. 5 C-E., pl. xii, fig. 48.
Six specimens from Perak and one from Sumatra ; also four from Médan (nr. Dolok-
Baros , Sumatra, presented by M. Guy Babault. Length 3275-4000 mm. The smaller
specimens have the grooves of the elytra more strongly punctured than the larger ones.

Pelopides burmeisteri (Kaup).
Eriocnemis Burmeister’, Kaup, 1868a, p. 22.
Gnaphalocnemis burmeisteri, Gravely, 1914c, p. 249, pl. xi, fig. 49.

A number of specimens from Sumatra (Mana-Riang and Tandjong-Djati, Renau,
Palembang; Bandar, Palembang; Bng. Proepoe, interior of Padang; Kandg. Ampat,
Padang Benedenl; also two from Médan, presented by M. Guy Babault) ; two from Java
(one being from Tji Solak, Wynkoop’s Bay) and six from Borneo (one being from Sintang
and three from Sarawak). Length 38-49 mm. ‘

1 Sec also Gravely, 1914c, p. 297 (T. dorsalis).

1918. | F. H. GRAVELY : Passalidae of the World. 95

Pelopides monticulosus (Smith).
Passalus monticulsus, Smith, 1852, p. 6, pl. 1, fig. 1.
Gnaphalocnemis monticulosus, Gravely, 1914c, pp. 249-250, pl. xiii, fie. 49a.

Three specimens from Sumatra (two being from Bedagei Interior, ca. 600 ft.), and a
number from Borneo (Mts. Kinabalu and Marapok). Length 30-45 mm.

Pelopides tridens (Wiedemann).

Biel.
Passalus tridens, Wiedemann, 1823, pp. 109-110.
Gnaphalocnemis tridens, Gravely, 1914c, p. 250, pl. xii, figs. 50-50a.

Nine specimens from Sumatra (Bedagei Interior, ca. 600 ft.; Gng. Talang, Padang
Interior ; Mt. Singalang) ; many from Java (Telaga Bodas, Garoet, 4,000-5,000 ft. ; G. Tji
Salimar, ca. 3,000 ft. ; Tji Bodas, Gng. Gede, ca. 4,000 ft., all in Preanger ; also Buitenzorg
and Tjandiroto) ; and one said to come from the Key Islands (near New Guinea). Length
46-56 mm.

The species of Pelopides may be identified as follows :—

The anterior lower tooth of the right mandible small hut distinct

1 | in unworn specimens
| The anterior lower tooth of the right mandible absent BE 32 st 3.

{ The outer tubercles at least approximately symmetrical ; the
anterior lower tooth of the right mandible larger .. P. symmetricus, p. 94.

| The inner angle of the right outer tubercle produced forwards
and inwards beyond that of the left ; the former acute, the
latter obtuse ; the anterior lower tooth of the right mandible
smaller en dt Be ag .. P. gravidus, p. 94.

The left outer tubercle consisting of a single, somewhat slender,
obliquely truncate process ; the right one, of a similar but
broader and slightly bifid mner process together with smaller
pointed outer and middle processes .. 55 .. P. simplex, (Gravely).
Both tubereles consisting of two or three denticles more or less
fused together, the tubercle as a whole being very broad
on the right or on both sides > a 3% er ase 4.
(The right and left outer tubercles of equal size, though not

always of identical form .. Ss oo Pons als Apt

The right outer tubercle much broader than the left

Grooves 5-7 of the elytra more or less broad ; each containing a
polished flattened band, which is marked by a single row of
punctures, and defined on either side by a more or less dis-
tinct roughened line with which the punctures may be to

some extent confluent .. Ets ed 5% de ie 6.

| The lateral grooves of the elytra all narrow, simply punctured P. burmeisteri, p. 94.

96 Memoirs of the Indian Museum. (Vor. VII,

Grooves 5-7 of elytra somewhat variable in width ; the posterior

part of groove 8 rarely wider than the anterior part, never

as wide as groove 7 ; insects at most about 45 mm. long .. P. monticulosus, p. 95.
Grooves 5-7 of elytra always very broad ; the posterior part (and

sometimes the whole) of groove 8 like them ; insects at least

about 45 mm. long as oH Tee imidenss pn 92.

Genus PLESTHENUS, Kaup, 1871, p. 40.

Type, Eriocnemis quadricornis, Kaup, 1868a, p. 26.

Of the four species referred by Kuwert (1898, pp. 324-5) to the genus Plesthenus one,
lottini, Boisduval, probably belongs to an Australian genus,’ the other three being
Celebean (see above, p. 76, footnote 2). Of these three 2nuitus, Kuwert, is the most distinct.
This species is represented in the Van de Poll collection by perhaps three specimens.
Each of them is, however, distinguished by some definite character from the others ; and at
least two localities are represented by the three specimens, one of which bears no locality
record. It is impossible, therefore, to be certain at present whether the three specimens
belong to three separate species or not ; and it will probably be best to describe all under
the one specific name, referring at the same time to the individual differences.

The other two species, quadricorms, Kaup, and gelon, Schaufuss, are together
represented by eleven specimens from five localities, two in the north of the island and
three in the south. The southern specimens are distinctly larger than the northern, and
they have an acuminate or very obliquely truncate right outer tubercle, the left outer
tubercle being more or less obsolete ; in the northern specimens, on the other hand, the right
outer tubercle tends to be more abruptly truncate, and the left outer tubercle to be more
strongly developed, in some instances much more so. Here again it seems impossible to
determine with certainty how many species are represented in the material before me, the
differences between different forms being in this case undoubtedly correlated with locality.
I propose, therefore, to treat all as a single species quadricornis, recognizing ge'on as a more
or less distinct southern race. The races seem to be distinguished more definitely by size
than by structure.

I am unable to identify any of the specimens before me with either of the species
described since Kuwert’s work was published (see above p. 9), but it is possible, I think,
that these may prove to be no more than varietal forms of P. quadricornis.

Plesthenus invitus, Kuwert.
PIME
Plesthenus invitus, Kuwert, 1898, p. 325.

Three specimens, one of which is from Tondano, Minahassa and another from Menado.
Both the specimens whose localities are recorded are 45°5 mm. long ; the other specimen is
much bigger, being 53 mm. long.

This species differs from Pelopides burmeisteri, monticulosus and tridens only in the.
generic character smentioned above (p. 79), in having the left anterior lower tooth wholly
distinct from the lowest terminal tooth instead of partially fused with it, and in the weaker

1 Presumably Mustochilus.

1918.] F. H. GRAVELY : Passalidae of the World. 97

and more crescentic scars on the mentum. The outer tubercles of the head are practically
symmetrical, the right one being slightly broader than the left, at least in the Tondano-
specimen. In this specimen the space between the inner and outer tubercles is surrounded
by fine but distinct ridges. of which the lateral and posterior are straight, and the anterior
procurved. In the large specimen the lateral ridges are absent, the anterior one is
procurved and the posterior one recurved ; and in the Menado specimen even the anterior
and posterior ridges are indistinct.
Plesthenus quadricornis, Kaup.
eae AL.

Eriocnemis quadricornis, Kaup, 1868a, p. 26.
Plesthenus quadricornis, 1871, p. 40, pl. iv, fig. 4.

Four specimens of the northern race, quadricornis, s. str. {(see above, p. 96), from
Toli-Toli and three from Menado, both in Northern Celebes. One specimen of the southern
race, gelon (Schaufuss, 1885, pp. 187-188), from Samanga, one from Patunuang, and two-
from Macassar, al in Southern Celebes. Length of the northern race 49-53 mm. Length
of the southern race 55-60 mm.

P. quadricornis differs from P. wmvitus only in having the right outer tubercle
somewhat longer, and sometimes more obliquely truncate or even acuminate, and in having
the left outer tubercle smaller and acuminate or obsolete. In the Toli-Toli specimens
the left outer tubercle is an acutely pointed process, scarcely shorter than the inner side of
the somewhat obliquely truncate right outer tubercle. In the Menado specimens it 1s much
shorter, the apex being rectangular or even obtuse. This is the case in the specimens from
Samanga and Patunuang also ; but in these the right outer tubercle is somewhat slenderer
and more obliquely truncate ; in size, moreover, these specimens resemble those from
Macassar and not the preceding. In the Macassar specimens the left outer tubercle is.
scarcely recognizable, and the right outer tubercle is acuminate.

The two above described species of Plesthenus may be identified thus :—

The outer tubercles approximately symmetrical, both of them

truncate ern oe a6 Ge -. P.ınvius, p. 96.
The outer tubercles asymmetrical, the right one much larger

than the left and truncate, except when the latter is obsolete,

in which case it may be acuminate; the left one never

truncate 50 So 56 | .. P. quadricornis, p. 97.

Genus MASTOCHILUS, Kaup, 18684, pp. 19-20, corrected 1868b, p. 31.

Type, Passalus polyphyllus, MacLeay, 1826, p. 430.
This genus, as pointed out above (p. 77), may be divided into four more or less distinct
sub-genera :—

The mentum laterally with a narrow depressed matt border, or
more extensively matt ; large Australian forms with very

1 short antennal lamellae ; always symmetrical .. .. Pharochilus,! p. 98.
The mentum uniformly glossy in fresh specimens ; the antennal
lamellae much longer ; or not Australian = ae na sis 2.

1 Kaup, 1868a, pp. 20-21. Type, Passalus dilatatus, Dalman.

98 Memovrs of the Indian Museum. [Vor. VII,

Robust. symmetrical insects at least 34 mm. long ; almost all

Australian! .. ee se oe .. Mastochilus, s. str., p. 100.
Smaller insects—never more than 33 mm. long; almost all

from New Guinea and the neighbouring islands ;? often

asymmetrical .. + se sy ae a ne 3.
The upper margin of the left mandible strongly angulate ;
the antennal lamellae longer and the body more flattened .. Analaches,? p. 101.
34 The upper margin of the left mandible, behind the upper tooth,
not or scarcely angulate ; the antennal lamellae shorter and
the body more robust of sa at .. Cetejus,* p. 102.

Mastochilus (Pharochilus) dilatatus (Schönherr).
Passalus dilatatus, Schoherr, 1817, p. 334, appendix, p. 144.

A few specimens from Queensland (Cardwell) and New South Wales (Richmond River
and Sidney). Length 31-36 mm.

The inner tubercles are large, somewhat as in M. quaestionis, but they are widely
separated, the frontal area being always strongly transverse, and not elevated. The general
texture of the upper surface of the head resembles that of M. australasicus. The more or
less confluent and transverse grooves on the mentum are deeply crescentic, being almost
equidistant from the anterior and posterior margins near the middle line; and the
punctured lateral parts of the mentum have a narrow matt border on the outer side.
The elytra resemble those of M. quaestionis and M. australasicus, except that the punctures
are somewhat transverse, especially towards the posterior end, where the ridges between
them are apt to be more or less obsolete.

Mastochilus (Pharochilus) nitidulus, Macleay.
IE

Mastochilus nitidulus, MacLeay, 1871, p. 175.
Pharochilus nitidulus, Kuwert, 1898, p. 333.

Two specimens from New England, New South Wales, presented by Mr. H. Schrader.

Length 41 mm.
The lateral lobes of the mentum resemble those of M. dilatatus. The median part of the

mentum resembles that of M. politus, which species this one resembles in all other

respects.

1 The only exception known is M. pectinigera (Heller) from New Guinea (length 39:5 mm). Concerning the position
of obliquus, Kirsch, see Heller, 1910, pp. 17-18.

? The only exception known is Analaches australiensis, which is apparently found both in Australia and New Guinea
(length 25:5-30:0 mm). ;

® Kuwert, 1896, p. 230 Type, Epilaches puberilis, Kuwert, 1898, p. 337 (see Zang, 1905a, p. 238, footnote ; also
p. 24). ;

4 Kaup, 1871, p. 53. Type, Aceraius virginalis, Kaup, 1868b, p. 5.

1918.] F. H. Gravety: Passahdae of the World. 99

Mastochilus (Pharochilus) politus (Burmeister).
Fig. XIIL 1.
Passalus politus, Burmeister, 1847, pp. 465-466.

A number of specimens from Queensland and Victoria, and two from Tasmania
(Brighton). Also three presented by Mr. H. Schrader, two being from Tingha and one from
the Clarence River, both in New South Wales. Length 305-370 mm.

The upper surface of the head resembles that of M. dilatatus, except that there is a
secondary tubercle developed on the upper side of the base of each of the outer tubercles,
which consequently appear very high and obliquely truncate in side view. The ridge

Fie. XIII.
Mastochilus spp., specific characters in the upper surface of the head x 4.
1. M. (Pharochilus) politus (Burmeister). 4. M. (Analaches) australiensis (Stoliczka).
2. M. (Pharochilus) punctiger, Gravely. 5. M. (Cetejus) grabowskii (Kuwert).
3. M. (s. str.) quaestionis (Kuwert). 6. M. (Cetejus) peltostictus (Kaup).

bounding posteriorly the crescentic groove on the mentum is usually more pronounced,

and the lateral matt border is very broad and extends round in front on to the inner

side of the punctured area. The punctures in the lateral grooves of the elytra, though
sometimes irregular, and inclined to be more or less transverse behind, as a rule resemble
those ot M. quaestionis and australasicus more closely than they do those of M. dilatatus.

Mastochilus (Pharochilus) punctiger, n. sp.
Fig. XIII, 2.

Two specimens from New South Wales. Length 33 mm.

The small triangular frontal area is bounded laterally by high frontal ridges and inner
tubercles, somewhat as is M. quaestionis, but is not itself elevated. The ridges joining the
inner tubercles to each other and to the outer tubercles are absent, a broad transverse
flattened rugose band extending between the supraorbital ridges from the inner tubercles
to the anterior margin of the head ; behind this band the head is smooth and glossy. The
mentum is somewhat coarsely and sparsely punctured between the scars, with a more or
less triangular, extensive (but sometimes indistinct) depression in front. Its lateral parts
are more closely and finely punctured on the inner side, obliquely striate with a more or
less smooth matt border on the outer.

The median groove of the pronotum is deeply impressed, but may be incomplete in
front ; the marginal grooves are coarsely punctured in their anterior extremities, which
bend slightly inwards; the remainder of them is very finely punctured, as are also the

pronotal scars. The general surface of the pronotum is unpunctured. The scutellum
02

100 Memoirs of the Indian Museum. [Vor. VII,

‘bears a number of scattered punctures, especially in front. The mesosternum is matt,
with a coarsely punctured A-shaped area in front. It is without scars. The posterior
‘intermediate areas of the metasternum are closely and coarsely punctured, with the
exception of a band bordering the somewhat narrow and rugose lateral areas. The elytra
resemble those of M. australasicus.

Mastochilus (s. str.) quaestionis (Kuwert).
Je UNE, 8 [Ds SL
Episphenoides quaestionis, Kuwert, 1898, p. 327.

A number of specimens from New South Wales, several being from the Richmond
River. Length 44-51 mm.

Easily distinguished from all other known species by its large inner tubercles, which
‚are situated very close to one another and to the central tubercle, the whole of the small
triangular concave frontal area being strongly elevated. The small central part of the
mentum is usually marked with a pair of more or less confluent shallow transverse grooves
a little behind, and a small depression in the middle of, the anterior margin ; but either or
both of these may be faint or absent ; or one may be so strongly developed as to obliterate
the other ; strong ridges, however, are never present. The lateral parts of the mentum, in
front of the primary scars, are glossy, punctured and hairy throughout. The pronotum
has no median groove ; its scars are punctured. he elytra are more coarsely punctured
at the sides than above.

Mastochilus (s. str.) australasicus (Percheron).
PISE
Passalus australasicus, Percheron, 1841, pp. 67, pl. Ixxvu, fig. 2.

A number of specimens from Queensland, New South Wales (Richmond River) and
Victoria (Melbourne) ; also a series said to come from Dodinga in Halmaheira ;! also
‘specimens presented by Mr. H. Schrader from Tingha and New England, New South Wales.
Length 37-48 mm.

The small inner tubercles are situated nearly half as far from the outer tubercles as
the latter are from each other, and almost directly behind them. The ridge uniting the inner
tubercles is often absent. When present it may lie close in front of the frontal ridges, or
some distance in front of them. The frontal area, which may thus be almost non-existent
-or of considerable size, is smooth and glossy ; the area between it and the anterior margin of
the head is very rough. The pronotum, mentum and elytra resemble those of M. quaestionis.

Mastochilus (s. str.) polyphyllus (MacLeay).
Passalus polyphyllus, MacLeay, 1826, p. 439.
Several specimens from Queensland and New South Wales (Sidney) ; also one said to
come from Dodinga in Halmaheira. Length 34-40 mm.
The central and inner tubercles are more widely separated than in the preceding species ;
and the frontal ridges, instead of running direct between the two, diverge either from an

—

1 Probably part of a series from Australia which has been wrongly labelled.

1918.] F. H. GRAvVELY : Passalidae of the World. 101

anterior prolongation of the central tubercle or at very acute angle, and curve first outwards
and then a little forwards, with the result that the frontal area appears markedly
transverse. The ridge joining the inner tubercles together is present, but there are no
ridges in front of these tubercles. The mentum is glossy throughout, and bears a small but
deeply impressed V- or U-shaped groove in the middle in front ; only the lateral parts are
punctured. The pronotum resembles that of P. punctiger, the median groove being very
strongly developed. The dorsal grooves of the elytra are imperfectly, the lateral ones
strongly but not very coarsely, punctured.

Mastochilus (Analaches) australiensis (Stoliczka).
Ihe, MODULE 44, jo, OY),
Cetejus australiensis, Stoliczka, 1873, pp. 157-158 (continuation of footnote to p. 156).

A number of specimens from Queensland, one from New South Wales and one from
New Guinea (Stephansort, Astrolabe Bay). Length 25:5-30:°0 mm. The smallest and
flattest species known from Australia, also the only one with asymmetrical outer tubercles

All six antennal lamellae are extremely long and slender. The upper tooth of both
mandibles is obsolete, and the denticle behind it, though well developed, is more or less
hidden beneath the ends of the supra-orbital ridges; from this denticle a slight ridge
extends downwards to the outer angle of the mandible, which is produced into an acute
and outwardly (almost forwardly) directed tooth. The outer tubercles are very large,
the left one slightly more so than the right; the gap between them is semi-circular.
The inner tubercles are situated at their base ; the frontal area is about twice as broad
as long. The general surface of the upper: side of the head is more or less strongly
punctured. The secondary scars on the mentum are transverse and matt; with the
middle of the anterior margin they enclose more or less completely a small triangular
glossy area.

The pronotum is more or less sparsely covered at the sides with large punctures, which
tend to concentrate in and around the scars. Its marginal grooves are punctured throughout
and are scarcely bent wards at their anterior ends. The median groove is more or
less obsolete. The dorsal grooves of the elytra are finely punctured ; the lateral ones are
much broader than the ridges between them, and the transverse ridges between their
enlarged matt punctures are more or less obsolete.

Mastochilus (Analaches) puberilis (Kuwert).
Epilaches pubarilis, Kuwert, 1898, p. 337.1

One specimen from Milne Bay, British New Guinea, and one presented by the British
Museum, also from New Guinea. Length 31-33 mm.

The mandibles are not angulate externally, and the denticle near the base of the upper
margin is not covered by the anterior angles of the head. The outer tubercles resemble
those of M. australiensis, but the inner ones are situated a considerable distance behind
them. The frontal area is smaller and more triangular. The pronotum is somewhat
indistinctly punctured in the scars and marginal grooves ; except for this it is unpunctured ;
the median groove may be somewhat stronger. The dorsal grooves of the elytra are

1 First mentioned in Kuwert’s 1891 list, where the name is spelt puberilis as above.

102 Memoirs of the Indian Museum. [Vor. VII,

unpunctured ; the lateral ones are strongly punctured. In other respects this species.
resembles M. australiensis.

Mastochilus (Cetejus) grabowskii (Kuwert).
Fig. XIII, 5, p. 99.
Cetejus grabowski, Kuwert, 1898, p. 330.

Three specimens from Humboldt Bay, New Guinea.t Length 28-5-29:0 mm. A stouter
insect than M. australiensis or M. puberilis, with shorter antennal lamellae.

The mandibles are not angulate externally, and there is no denticle exposed on the
upper margin behind the rudimentary upper tooth. The upper surface of the head resembles
that of M. puberilis, except that the outer tubercles are more outwardly directed, that the.
central and inner tubercles are more widely separated, and that the frontal ridges are more.
or less incomplete between them. The secondary scars on the mentum are represented by
a more or less broken transverse groove a little behind the anterior margin. The scars,
anterior angles, and marginal grooves of the pronotum are coarsely punctured ; the median
groove is very strong. All the grooves of the elytra are somewhat coarsely punctured.

Mastochilus (Cetejus) sodalis (Kaup).
Aceraius sodalis, Kaup. 1868a, pp. 29-30 : and 18685, p. 5.
A number of specimens from Ternate and Batjan, one of the latter being from Labuan ;
also one irom Taruna, Gt. Sangir, 2,000 ft. Length 24-0-25-8 mm.
This species closely resembles the last, but is smaller and therefore less perceptibly
asymmetrical ; it has a somewhat shorter frontal area, and has the groove formed by the-
secondary scars on the mentum deep and unbroken.

Mastochilus (Cetejus) peltostictus (Kaup).

Fig. XIII, 6, p- 995 also pl. I.
Aceraius peltostictus, Kaup, 18685. pp. 5-6.

Two specimens from Ceram and many from New Guinea, one of the latter being from.
Stephansort. Length 22-5-26-3.

The outer tubercles are strongly asymmetrical, the left one being slightly longer than
the right, broad and truncate instead of slender and pointed, and more inwardly directed.
The groove representing the secondary scars on the mentum is more strongly arched away
from the anterior margin than in M. sodalis, and the grooves of the elytra are more finely
punctured, especially the dorsal ones. In other respects M. peltostictus resembles M.
sodalrs.

1 The two specimens which I recorded from Stephansort (1914c, p. 334), though compared with named specimens in.
Berlin, do not agree with Kuwert’s description of the present species, and are apparently M. peltostictus, Kaup.

1918.] F. H. Gravety: Passalidae of the World. 103
The above-mentioned species of Mastochilus may be grouped into four sub-genera as
indicated above (pp. 97-98), and may be distinguished from one another thus :—

I. Sub-genus PHAROCHILUS, Kaup.
The median part of the mentum with at most two or three

punctures .. : : : - = + 2.
The median part of the mentum ray een eee
out ae à er .. M. punctiger, p. 99.
lateral lobes of the mentum En a more or less narrow
= matt border on the outer side af er = = = 3.
The lateral lobes of the mentum fonts matt on the outer
| side Be 55 D <i . M. politus, p. 99.
The ridges of the elytra obsolete in the extreme posterior ales M, dilatatus, p. 98.
sf The ridges of the elytra normal a: = .. WM. mtidulus, p. 98.

II. Sub-genus MASTOCHILUS, s. str.

The frontal area small, almost equilaterally triangular, raised
above the general surface of the head “6 .. M. quaestionis, p. 100.

The frontal area not raised above the general surface of the head,

usually much broader than long sis :
The secondary scars very variable in development ; when

present always extending inwards from the ends of the

anterior margin of the median part of the mentum .. WM. australasicus, p. 100.
The secondary scars always developed as a pair of fine grooves

defining a smal] and more or less equilaterally triangular

shield in the middle of the anterior margin ae .. M. polyphyllus, p. 100.

|
III. Sub-genus ANALACHES, Kuwert.
The lateral grooves of the elytra much broader than the ridges
between them ; the transverse ridges between their enlarged
matt punctures more or less obsolete .. M. australiensis, p. 101.
The lateral grooves of the elytra narrow and strongly punc-
tured 2: de + sts .. M. puberilis, p. 101.

IV. Sub-genus CETEJUS, Kaup.

The left outer tubercle pointed, directed forwards or a little
outwards

The left outer tubercle truncate, decried inwards .. .. WM. peltostictus, p. 102.
The secondary scars on the mentum represented by a more or
less broken transverse groove a little behind the anterior

2 margin Me à & . M. grabowskii, p. 102.
The groove formed by the dede scars on the ein dep
and unbroken = AR ss . M. sodalis, p. 102.

Genus LABIENUS, Kaup, 1871, p. 39.
Incl. Aurelius, Kuwert, 1896; Hyperplesthenus, Kuwert, 1808 ; Kaupioloides, Gravely,
1913 ; Kaupiolus, Zang, 1903b (=Vellejus, Kaup, 1871, preoccupied).
Type, Eriocnemis ptox, Kaup, 18682, p. 25.

104 Memoirs of the Indian Museum. - (Vor. VDE

Labienus inaequalis, n. sp.
Figs. XIV, 1 and XV. 1.

Two specimens from Hattam, Arfak, Dutch New Guinea. Length 37°7-38:7 mm.

This species is closely allied to 2. trigonophorus, from which it differs in the following
particulars only. The anterior margin of the labrum is more strongly concave. The
primary scars on the mentum are much smaller, and the secondary ones much larger, the
latter almost meeting in the middle line immediately in front of the posterior margin. The
general surface of the head is less rugose than in L. trigonophorus, and the left outer tubercle
is much larger and more definitely directed inwards ; the frontal ridges are obsolete in front.

SUR

Fic. XIV.
Labienus spp. ; specific characters in the upper surface of the head x 4.

1. L. inaequalis, Gravely. 3. L. ptoxoides, Gravely.

2. L. dohrni (Kuwert). 4. L. compergus (Boisduval).
of the inner tubercles. The anterior angles of the prothorax are somewhat more obtuse ;
the median groove is complete. The posterior intermediate areas of the metasternum are
almost unpunctured as well as being hairless ; the lateral areas are thickly punctured and.
hairy as in L. trigonophorus, their surface being on the same level as that of the intermediate
areas, an abrupt change of level occurring only where they touch the central area.
Neither in L. trigonophorus nor in L. inaequalis are the elytra united.

Labienus dohrni (Kuwert).
Fig. XIV, 2; also pl. 1.
Aurelius dohrni, Kuwert, 1898, p. 326.

Four specimens from Dutch New Guinea, three being from Hattam, Arfak, and one
from Kapaur. Length 42-47 mm.

This species is easily recognized by the small free forwardly directed apex of the central
tubercle and the absence of inner tubercles. The frontal and parietal ridges are obsolete
or absent. The outer tubercles may be simple and symmetrical, or the left one may be
more or less distinctly double ; the anterior margin of the head is usually more or less rough
between them, this rough area being separated from the rest of the head by a fine ridge
or groove.

The pronotum is much broader behind than in front ; its median groove is indistinct ;
it is without punctures except in the posterior angles (including the true scars, though
other unpunctured depressions may be present near them), which are densely hairy
and punctured. The scutellum bears two longitudinal lines of fine punctures. The
mesothoracic episterna are punctured except in the posterior angles. The mesosternum

‘is smooth and polished ; its scars are distinct, but very small and less close to the margin
than usual. The metasternum resembles that of L. trigonophorus. The grooves of the
elytra are deep but unpunctured.

1918.] F. H. GRAVELY : Passalidae of the World. 105

Labienus ptoxoides, n. sp.
Figs. XIV, 3 and XV, 2.
Two specimens, one from Andai and the other from Mt. Arfak, both in New Guinea.
Length 44-46 mm.
This species is allied to L. gracilis and L. ptox. The antennae resemble those of L. ptox,
the first two lamellae being quite short and the remaining four very long. The secondary
scars of the mentum are straight, and extend to the posterior margin. The head closely

INE, DOVE

Labienus spp. ; mentum x 8.
1. L. inaequalis, Gravely. 2. L. ptoxoides, Gravely.
3. L. ptox, Kaup.
resembles that of both L. gracilis and L. ptox, especially the latter. The pronotum resembles
that of L. ptox, the scars being more densely hairy and punctured than in L. gracilis. In
all other respects the present insect resembles both species.

Labienus ptox, Kaup.

Fig. XV, 3.
Eriocnemis ptox, Kaup, 1868a, p. 25.
Labienus ptox, Kaup, 1871, p. 39.

Numerous specimens from New Guinea (Stephansort, Bongu, Milne Bay and Roon
Islands), Aru (Ureiuning and Wamma Dobbo) and Waigeu. Also one said to come from
Borneo and three from Sumatra. Length 48-58 mm. _ |

This species differs from the last only in its larger size and in having the scars on the
mentum strongly arched instead of straight.

Labienus compergus (Boisduval).
Fig. XIV, 4; also pl. 1.
Passalus compergus, Boisduval, 1835, pp. 244-246.
Vellejus compergus, Kaup, 1871, p. 36.

Several specimens from New Guinea (Milne Bay ; Stephansort, Astrolabe Bay ; Anda;
Kapaur ; and Hattam, Arfak) and one from Waigeu. Length 32-36 mm.

The antennae resemble those of the majority of species belonging to the genus. The
right outer tubercle is more or less distinctly double. The left outer tubercle may be
similar, or may be composed of three more or less distinct processes. The pronotal scars
are unpunctured and hairless. The mesosternal scars are variable. The posterior

intermediate areas of the metasternum are unpunctured. All the grooves of the elytra»
P

106 Memoirs of the Indian Museum. [Vor. VII,

-especially the lateral ones, are distinctly punctured. In other respects this species resembles

L. ptox.
Labienus moluccanus (Percheron).

Passalus mollucanus, Percheron, 1835, pp. 31-33, pl. u, fig. 7.

Numerous specimens from Ceram (Roemasosa]-Pasama in the central part, Wahaai
in the north, Elpapoeti Bay in the south, Honitetoe in the West), Limtoe in Nusa Laut,
Saparua, Hitu and Leitimor in Amboina, Batjan. Length 42-52 mm.

This species differs from the last only in having the frontal ridges more or less obsolete
behind the inner tubercles, in having the two processes of each of the outer tubercles less
distinct (much as in L. ptox and ptoxoides) and in having the elytra united in the middle

line.
Labienus gigas (Kaup).

12h, Il,
Eriocnemis gigas, Kaup, 1868a, p. 23.

Seven specimens from Batjan, including five from Laboean, one from Halmaheira
one from Ternate, and two without locality records. Length 56-64 mm. I am unable to
distinguish between L. gigas from Ternate and L. crassus from Batjan.

The elytra are united as in L. moluccanus, which the present species resembles in all
respects, except in having each of the outer tubercles composed of three more or less distinct
blunt processes, and in having the posterior intermediate areas of the metasternum, and as
a rule the pronotal scars, punctured, though not hairy.

The species of Labienus known to me may be identified thus :—
( The secondary scars on the mentum convergent behind, curved
or straight, not very widely separated as a whole ss ae oie 2,
) The secondary scars on the mentum strongly arched outwards,
| very widely separated asa whole .. oc Se 5% si 8.
_ The outer tubercles simple, conical ; the grooves of the elytra
2 punctured
Not as above

/ The left outer tubercle scarcely longer than the right ; the

: mentum with primary scars on u .. L. trigonophorus (Zang)*.
‘ | The left outer tubercle much longer than the right ; the mentum
without primary scars .. os .. L. imaequalis, p. 104.
( The central tubercle free apically ae i .. L. dohrni, p. 104.
4 ( The central tubercle not free ai an a Re Bee 5,
_ C The outer tubercles long and simply truncate te .. L. glaber (Gravely).
> The outer tubercles much shorter, not simply truncate 5, IN ae 6.

Both outer tubercles composed of two blunt denticles ; the scars
of the pronotum without punctures or hair De .. LL. impar (Kuwert).!
6< The inner denticle of the right outer tubercle more or less
obsolete ; the scars of the pronotum with hair-bearing punc-
| tures X: I fs 2%, de Se Er To

1 See Gravely, 1913, for figures of these species,

1918. | F. H. Gravety: Passalidae of the World. 107

The antennal lamellae normal en or .. L. gracilis, Heller.!
7 The first two antennal lamellae very short, the remaining four
exceptionally long + 7 x: .. L. ptoxoides, p.105.
The first two antennal lamellae very short, the remaining four
8 exceptionally long a 55 ig .. L. ptox, p. 105.
The antennal lamellae normal a ae ae ae aS oh
( The elytra separate ; insects of moderate size Ne .. L. compergus, p. 105.
( The elytra united ; insects of large size a % = = 10.

The outer tubercles much narrower than the space between
them, each bemg composed of two more or less distinct
blunt processes only; the posterior intermediate areas
of the metasternum smooth ae Ss .. L. moluccanus, p. 106.
10% The outer tubercles at least as broad as the space between
them, each including an additional blunt process on the inner
side of, and some distance from, the two found in Z. moluc-
censis ; the posterior intermediate areas of the metasternum
coarsely punctured BA 9.0 Re .. L. gigas, p. 106.

Genus PROTOMOCOELUS, Zang, 1905), p. 154.
=Pelops, Kaup, 1871, preoccupied.
Type, Passalus australis, Boisduval, 1835, pp. 246-247, pl. vi, fig. 21.

Protomocoelus australis (Boisduval).
TA, Il:
Passalus australis, Boisduval, 1835, pp. 246-247, pl. vi, fig. 21.
Pelops australis, Kaup, 1871, p. 38.

Five specimens from the Solomon Islands (including one from San Cristoval and one
from Bougainville), three from New Brittain, many from New Guinea (Milne Bay)
Stephansort and Isola Yule), and several from Waigeu, Aru (Wamma Dobbo and Ureiuning)
and Ceram. Also one specimen said to come from Australia. Length 30-47 mm.
The Solomon Islands specimens (except the one from Bougainville, which is only 35 mm.
long), and the specimen labelled Australia, are much the largest, none of the others exceeding
37 mm. in length. Apart from the Bougainville specimen the smallest of the Solomon
Islands specimens is 41 mm. long. I am unable, however, to find any constant structural
difference between the Solomon Islands specimens and the others, and am consequently
unable to recognize P. solomonis (Kaup) as distinct. P. australis is somewhat variable,
and the validity of Kuwert’s species may be doubted.”

Protomocoelus australis is probably allied more closely to Labienus inaequalis than to any
other species yet described. It differs from it, however, in the structure of the mandibles (see
above, pp. 78 and 79) ; in the broader and often more widely separated, but very variable.

1 See Gravely, 1913, for figures of this species,

2 Passalus impressicollis, Boheman 1858, p. 40, cannot belong, as supposed by Kuwert, to the present genus, for its
outer tubercles are equal and obtuse instead of unequal and acute. It comes from Sydney, and not from Menado as stated
by Kuwert ; it is said by Boheman to be allied to Wastochilus polyphyllus, and doubtless belongs to the same genus.

108 Memoirs of the Indian Museum. - [Vor. VII,

scars on the mentum ; in having the frontal ridges more or less obsolete behind instead of in
front of the inner tubercles ; in the deep concavity between the outer tubercle and the
anterior end of the supraorbital ridge of the left or of both sides of the head ; in having
the right outer tubercle about as long as the left although more slender ; in having no
distinct median groove on the pronotum ; in usually having punctures on the posterior
intermediate areas of the metasternum ; and in having the dorsal grooves of the elytra
as distinctly punctured as the lateral ones. The elytra are not united in either species.

Genus GONATAS, Kaup, 1871, p. 50. ‘
Incl. Omegarius+ ? Tatius, Kuwert, 1896, p. 229.
Type, Passalus naviculator, Percheron, 1844, pp. 1-2, pl. exxxiv, fig. 1.
Gonatas minimus (Kuwert).
Tell, I,

Omegarius minimus, Kuwert, 1898, p. 313.
Omegarius minimus, Gravely, 1913, pp. 110-111, text-fig. 3A.

Three specimens from New Brittain, of which two are from Herbertshöhe ; one said
to be from Australia and one without locality record. Length 20-25 mm.

Gonatas pumilio, Kaup.

Aceraius pumilio, Kaup, 18685, p. 6.
Gonatas pumilio, Kaup, 1871, p. 50.
Omegarius pumilio, Gravely, 1913, p. 112, text-fig. 3B.
Several specimens from New Guinea (Torres Straits, Fly River, Kapaur), Waigeu
Amboina (Leitimor), and Ceram (Honitetoe in the western, 'Wahaai in the northern, and

Fie. XVI.
Gonatas spp. ; mentum x 8.
1. G. cetioides, Zang. 3. G. carolinensis, Gravely.
2. G. tenimbrensis, Gravely. 4. G. minor, Gravely.

5. G. naviculator (Percheron).

Roemasosal-Pasania in the central parts of the island). Also one specimen said to come
from the Sulu Islands. Length 18°5-23'0 mm.

Gonatas cetioides, Zang.
Fig. XVI, 1.
Gonatas cetioides, Zang, 1905a, p. 316.

One specimen from Herbertshöhe, New Brittain. Length 25 mm.
The antennae resemble those of G. minimus, the mandibles those of @. pumilio. The
posterior margin of the mentum is very lightly curved as in both species. but the lateral

1918.] F. H. Gravety: Passalidae of the World. 109

forwardly directed parts of the scars are much more, and the other parts less, deeply impressed
than in either. The lateral areas of the metasternum are somewhat smoother and the
prenotum and elytra somewhat more convex than in either.

Gonatas schellongi, Kuwert.
le
Gonatas schellongi, Kuwert, 1898, p. 314.

Numerous specimens from New Guinea (Stephansort, Milne Bay, Humboldt Bay,
Torres Straits), Kei Islands, and New Brittain (Herbertshöhe). Length 28-32 mm.

The antennal lamellae are longer than in the two preceding species, and the posterior
margin of the mentum is more strongly curved, the scars consequently forming a W- rather
than a w-shaped figure. The left mandible is as broad as the right and scarcely if at all
longer ; its anterior lower tooth is less distinct than in G. minimus, but more distinct than
in other species of the genus. The form of the outer tubercles varies slightly, and
G. tridentatus, Kuwert, is unlikely, I think, to prove distinct ; G. differens, albertisi, major
and novaebrittanniae will perhaps also prove to be identical with the present species. The
lateral areas of the metasternum are punctured and hairy. The dorsal grooves of the elytra
are less distinctly punctured than are the lateral ones.

Gonatas germari, Kaup.
Aceraius germari, Kaup, 1868a, pp. 30-31.
Gonatas germari, Kaup, 1871, p. 51.
Gonatas germari, Gravely, 1914c, pp. 250-251, pl. xin, fies. 47-47a.

Numerous specimens from Ternate and Batjan (Labuan), five from Halmaheira
(Dodinga) and Great Banda, one from Morty Island near Halmaheira, one from the Kei
Islands, and one said to be from Australia. Also one from New Guinea (Dorey) and four
from Buru (Wakollo in the central part of the island, and [lat on the east coast) all of much
larger size. Length, excluding the Dorey and Buru specimens, 23:5-26:5 mm. ; length of
Dorey specimens 29:0 mm. ; length of Buru specimens 31-0-32°5 mm.

The antennae and mentum resemble those of @. schellongi. The left mandible is
distinctly longer than the right, and its anterior lower tooth is more or less obsolete. The
lateral areas of the metasternum are punctured and more or less hairy. The difference
between the distinctness of the punctures in the dorsal and lateral grooves of the elytra is
less great than in G. schellongi.

Gonatas tenimbrensis, n. sp.
Fig. XVI, 2.

Five specimens from Tenimber (or Timor Laut), four being from Jandema. Length
25°5-27°0.

Closely allied to the preceding species, from which it differs only in having still less
or no trace of the left anterior lower tooth, in having antennae with shorter lamellae like those
of G. minimus and G. cetioides, and in having the lateral areas of the metasternum
unpunctured and hairless. The scars on the mentum are deeply impressed throughout.

|
I}

110 Memoirs of the Indian Museum. [Vor. VII.

Gonatas carolinensis, n. sp.
Fig. XVI, 3, p. 108.
Three specimens from the Caroline Islands. Length 22°4-23:2 mm.
This species differs from the last only in its smaller size, slightly longer antennal lamellae,
and more even mentum, the scars being less deeply impressed, especially medially.

Gonatas minor, n. sp.
Fig. XVI, 4, p. 108.

Four specimens from Mefor (“‘ Mafor ”) and one from Run (“ Roon ”) Islands. Length
21°3-22°2.

The antennae are very long as in G. germari. The difference in length between the
right and left mandibles is somewhat greater in G. minor and in the next species than in
any other. The mentum resembles that of G. germari ; its scars are less deeply impressed
than in G. tenimbrensis, but more deeply impressed than in G. carolinensis. The lateral
areas of the metasternum are smooth and hairless. In other respects the present species
resembles G. germari and G. naviculator.

Gonatas naviculator (Percheron).
Fig. XVI, 5, p. 108; pl. T.
Passalus naviculator, Percheron, 1844, pp. 1-2, pl. exxxiv, fig. 1.
Gonatas naviculator, Kaup, 1871, pp. 50-51.

Numerous specimens from Saparua Island, several from Ceram (Kairatoe in the
Western and Roemasosal-Pasama in the central part of the island) and Buano, one from
Nusa-Laut, and one said to come from New Guinea. Length 23-0-28:5 mm.

G. naviculator can be distinguished from all other species of the genus known to me
by the strongly and as a rule abruptly curved posterior margin of the mentum, though this
character is not always so clearly marked in specimens from the mainland of Ceram as in
those from the neighbouring islands. The antennal lamellae are comparatively short and
stout as in G. tenimbrensis, etc. The mandibles resemble those of G. minor. The outer
tubercles are somewhat slenderer than in @. germari, which the present species resembles

in other respects.

The above-mentioned species of Gonatas may be identified thus :—

The posterior margin of the mentum very lightly curved :
the scars more or less w-shaped 1 a x = ve 2.
The posterior margin of the mentum more strongly curved ;
the scars more or less W-shaped 7 Sn un oe SR 4.
The dentition of both mandibles complete and well developed G. minimus, p. 108.
24 The anterior lower tooth of the left mandible rudimentary or
absent
‘Che antennal lamellae very slender ; the scars on the mentum
distinctly w-shaped en 3
34 The antennal lamellae stouter ; the scars on the mentum
intermediate between the » and W form, very deeply

impressed laterally

G. pumilio, p. 108.

G. cetioides, p. 108.

1918.] F. H. Gravety: Passalidae of the World. 111

dible of about the same size as the right + .. G. schellongi, p. 109.
The left anterior lower tooth more or less rudimentary ; the
left mandible more or less distinctly longer and slenderer

The left anterior lower tooth small but distinct; the left man-
4
than the right we i i 66 52 5 5.

The posterior margin of the mentum moderately strongly

arched Ir a oe > x 6.
The posterior margin of ine mentum very strongly ea

the antennal lamellae somewhat short and stout ; the Talea

areas of the metasternum somewhat rough and hairy .. G. naviculator, p. 110.

The lateral areas of the metasternum more or less rough and
hairy ; the antennal lamellae very long and slender .. G. germari, p. 109.
The lateral areas of the metasternum smooth and hairless a ae a Ue

The antennal lamellae somewhat short and stout ; the scars
on the mentum very deeply impressed a .. G. tenmbrensis, p. 109.
The antennal lamellae longer and slenderer ; the scars on the
mentum less deeply impressed ds Me 7 ei a 8.

The left mandible and the antennal lameilae moderately long

and slender; the scars on the mentum very lightly im-

pressed, especially near the middle line à G. carolinensis, p. 110.
The left mandible and the antennal lamellae net Tone

and slenderer ; the scars on the mentum normal 56 Cie WOT 19, IND,

Genus PSEUDEPISPHENUS, Gravely, 10140, p. 327.
Type, Pseudepisphenus perplexus, Gravely, 1914¢, pp. 327-328, text-fig. 8, A-B.

Pseudepisphenus perplexus, Gravely.
Pil, lls
Pseudepisphenus perplexus, Gravely, 1914c, pp. 327-328, text-fig. 8, A-B.
One specimen from Snow Mts., 4,000-6,000 ft., Dutch New Guinea, presented by the
British Museum. Length 29:5 mm.

Genus TARQUINIUS, Kuwert, 1896, p. 227.
Type, Tarquinius paradoxus, Kuwert, 1898, p. 279 ; Gravely, 1914c, pp. 178 & 327, text-
dig. 8, ©.-D. (see pl. I).

Subfamily LEPTAULACINAE.
The genera of Leptaulacinae may be separated thus :—

The sides of the elytra hairy ap oe .. Trichostigmus, p. 112.
The sides of the elytra hairless 35 > .. Leptaulaz, p. 112.

112 Memoirs of the Indian Museum. [Vor. VII,
Genus TRICHOSTIGMUS, Kaup, 1871, p. 31.

Trichostigmus ursulus, (Schautuss).
Leptaulax ursulus, Schaufuss, 1885, p. 187.

A number of specimens from 8. Celebes (Lompa-Battau, 3,000 ft., and Tjamba).
Length 16-0-19°5 mm.

Trichostigmus ursulus resembles Leptaulax bicolor, except in the generic character and
in having the sides of the pronotum more sparsely punctured in the neighbourhood of the
scars and not at all in the anterior angles.

Trichostigmus thoreyi, Kaup.
Trichostigmus thoreyi, Kaup, 1868a, pp. 13-14.
A single specimen presented by Mr. C. F. Baker from Imugin, N. Viscaya, Phillipines.
Length 16-7 mm.
T. thoreyi differs from the preceding and only other known species of the genus only
in the structure of the pronotum.

The species of Trichostigmus may be distinguished thus :—
( The pronotum with a few punctures in the anterior angles, its
marginal grooves broad and deep and coarsely punctured T. thoreyi, p. 112.
The pronotum unpunctured except near the scars, its
| marginal grooves very fine ae oe so 1 Sons, ap NID,

Genus LEPTAULAX, Kaup, 18682, p. 11.

Incl. Leptaulacides, Zang, 19054, p. 106, footnote 1.

Type, Passalus dentatus, Fabricius, 1792, p. 241.

The account of this genus which I published in 1914 was based ete on the
examination of specimens from Continental Asia. I had, it istrue, received a few
specimens from the Archipelago; and I was able to make a hurried examination of the
named collection in Berlin. But I had had no opportunity of examining a large and
representative collection at leisure, an opportunity which has now been afforded by the
obtaining of the Van de Poll collection for the Indian Museum.

The careful examination of this additional material convinces me that the drastic
reduction in the number of species, advocated in my previous paper, was fully justified
except in the case of L. barbicauda, Zang ; and, indeed, that a few further reductions must
be made. Thus L. obtusidens proves to be a synonym of L. bicolor ; and L. novaeguineae,
together with the names regarded as synonyms with it, are almost certainly synonyms of
the same species, or partly of the same species and partly of L. dentatus.

I have seen nothing in the collection that can be distinguished as L. macassariensis ;
but a specimen labelled with this name, and associated with specimens both of L. bicolor
and L. dentatus, proves to belong to the former species. I am still inclined to think, judging
from Schautuss’s description, that the type may prove to be of a distinct species, the
puncturing of the head being apparently much coarser than in L. bicolor, and the convexity

1918.] F. H. GRAvVELY : Passalidae of the World. 113

of the body much greater than in L. cyclotaenius, the only other species known to me from
Celebes with which it can possibly be identified. But for the present it seems best to drop
the name macassariensis, raising the Bornean anibarbis to specific rank.

The variation of L. cyclotaenius in size, form, and head-puncturing proves to be much
greater than I previously supposed, especially in Malaysian specimens ; and the distinction
between the Malaysian and continental races breaks down. The name himalayae therefore
becomes a synonym only. L. anipunctus is very near L. cyclotaenius and may prove to be
identical with it. For the present it seems best to regard it as a variety of that species.

I am no longer able to regard the varieties vicinus and glabriventris, of L. bicolor

and dentatus respectively, as distinct.

The three species L. bicolor, L. cyclotaenius and L. dentatus are so variable as to require
very special care in their discrimination. The first and third can always be told apart by
the structure of their parietal ridges, which extend outwards to the supraorbital ridges in
the former, and end abruptly not far from the central tubercle in the latter. In
L. cyclotaenius these ridges are variable ; but the puncturing of the lateral grooves of the
elytra is much more distinctly scalariform than is ever the case in L. bicolor (this is usually,
but not always, so in L. dentatus also); and the central area of the metasternum is
almost invariably punctured either irregularly or over a more or less V-shaped area,
punctures being absent or confined to a single symmetrically placed pair in L. dentatus.
In the rare cases where the general appearance of the specimen resembles that of
L. cyclotaenius, and the central area of the metasternum is entirely without punctures—
I have only seen one such, and very few in which these punctures were not at least
moderately numerous, all of these being from Sumatra or the Malay Peninsula—one can
only base one’s identification on the somewhat indefinite and variable characters afforded
by the shape of the frontal ridges and the puncturing of the head.

Leptaulax planus (Illiger).
Passalus planus, Uliger, 1800, p. 104.
Leptaulax planus, Gravely, 1914c, pp. 260-261 and 310, pl. xiii, fig. 58.

One specimen from Siam, nine from the Malay Peninsula (four of them from Perak,
and one from Larut), many from Sumatra (Bedagei Interior, ca. 600 ft. ; Tandjong Morawa;
Serdang ; S. E. Serdang, ca. 1,000 ft.; Png. Pandjang, Padung Interior, ca. 2,000 ft.,
Tandjong-Djati, Ranau, Palembang, ca. 2,000 ft.) and Borneo (Sarawak; Brunei ;
Doesonlanden ; Martapura ; Mt. Marapok, Dent Province; Mt. Kina-Balu) and one from
Celebes (Tondano, Minahassa). M. Guy Babault has sent specimens for examination from
Medan, Sumatra. Length 12-3-14-0 mm.

Leptaulax glaber (Kirsch).

Trichostigmus glaber, Kirsch, 18775, pp. 139-149.
Leptaulax glaber, Gravely, 1905c, p. 307.

One specimen from Batjan and four from New Guinea (Humboldt Bay, Mt. Arfak and
Takar). Length 14:0-15°8 mm.

But for the reddish-brown colouration commonly found on the anterior parts of the

elytra, this species might easily be confused, with small and much flattened specimens of the-

114 Memoirs of the Indian Museum. [Vor. VII,

Polynesian form of L. bicolor. The specimens before me, however, show the extent of this
colouration to be extremely variable ; for in one of the Humboldt Bay species it covers
about a half and in the other about a third of the whole area, while in a third specimen from
New Guinea it is confined to a somewhat indistinct patch between the shoulders and in the
fourth it is entirely absent. The chief characteristics of the species, apart from colour,
are its extreme flatness, the fineness of the marginal grooves of the pronotum, and the
almost entire absence of punctures from the pronotum and metasternum. The frontal
ridges extend outwards and slightly forwards, to end somewhat abruptly at a considerable
distance behind the outer marginal tubercles.

Leptaulax sambawae, n. sp.

Four specimens from B. Aroe Hassa, Sambawa, 2,000-5,000 ft., and two from Poera,
Allor Islands, 3,000-4,000 ft. Length 24-27 mm.

This species differs from L. bicolor only in having the pronotum somewhat less
distinctly rectangular in shape, and entirely unpunctured except somewhat indistinctly in
the scars and still more indistinctly in the marginal grooves ; in having the punctures on
the posterior intermediate areas of the metasternum more or less obsolete ; and in having
the elytra distinctly wider behind than in front with their lateral grooves much less strongly
punctured. The abdominal sterna are polished and are entirely unpunctured in two
specimens, the terminal segment being marked in others with hair-bearing punctures.

Leptaulax barbicauda (Zang).
Leptaulacides barbicauda, Zang, 1905a, pp. 164-165.
Several specimens from the Malay Peninsula, (Gap, ca. 3,000 ft., Selangor-Pahang
boundary) submitted by Mr. C. Holman Hunt. Length 27-30 mm.
This species is transitional between L. sambawae and L. bicolor, and 1s so near the latter
most variable species that I have some hesitation in regarding it as distinct. It is, however,

distinctly bigger, with large and strongly rectangular pronotum, the general appearance

of the insect consequently resembling that of L. dentatus. The puncturing of the pronotum
and metasternum is weaker than is usually the case in L. bicolor, tending to resemble rather
that found in L. sambawae ; there are always, however, a few punctures in the anterior
angles of the pronotum, and the punctures in the pronotal scars and on the posterior
intermediate areas of the metasternum are somewhat stronger.

Leptaulax bicolor (Fabricius).
Passalus bicolor, Fabricius, 1801, p. 256.
Leptaulax bicolor + var. vicinus, Gravely, 1905c, pp. 257-259 and 307-309.

Numerous specimens, including one or more from each of the following localities :—
Ceylon (Belihul-Oya) ; Parambikulam, Cochin State, 1,700-3,200 ft. (collected by myself) ;
Santi Koppa, N. Coorg (presented by Mr. T. Bainbrigge Fletcher); Tukvar, Darjeeling
District ; Pashok, Darjeeling District, 2,000 ft. (collected by myself) ; Margherita, Assam ;
Port Blair, Andamans (collected by Mr. 8. W. Kemp); Tonkin (Nape, Thadua, Chapa,
Hoabink, and Xieng Khouang, submitted by M. Vitalis de Salvaza ; and Cape Fouquet,
submitted by M. Guy Babault); Siam; Perak, Malay Peninsula; Gap, 3,000 ft.,

1918.] F. H. Gravety: Passalidae of the World. 115

Selangor-Pahang boundary, Malay Peninsula (one specimen sent for examination by
Mr. C. Holman-Hunt) ; Hili Madjedja and G. Madjedja, North Nias ; Kalim Bungo, Middle
Nias , Sumatra (Bedagei Interior, ca 600 ft. ; Beloe Lawang, Pasoeroean ; Mana Riang,
Palembang, 2,000-3,000 ft.; Tandjong-Djati, Ranau, Palembang, ca. 2,000 ft.; Kandg.
Ampat, Lower Padang; Bng. Proepoe, Pad. Bovenland, ca. 1,600 ft.; Engano Island,
Benkoelen : also specimens from Medan, submitted by M. Guy Babault) ; Java (Bogor—
Buitenzorg; Tji Bodas, Gng. Gede, ca. 4,000 ft. ; Telega Bodas, Garoet, Preanger, 4,000-
5,000 ft.; Mt. Tjikorai, 4,000 ft., Sukabumi, 2,000 ft., and Pengalengan, 4,000 ft., W.
Java; G. Tji Salimar, ca. 3,000 ft., W. Preanger ; Tji Solak, Wunkoops Bay; Mt.
Tengger, 4,000 ft., E. Java; Malang); Borneo (Mt. Marapok ; Mt. Kinabalu ; Sarawak ;
Pontianak ; Doesonlanden ; 1° S., 115° E.; Banguey Island) ; Philippine Isiands (Davao:
also specimens presented by Mr. C. F. Baker from Imugin, N. Viscaya ; Mt. Makiling,
Luzon ; Zamboanga, Mindanao ; Mt. Bonatao and Los Banos) ; Talaut Islands (Salıbabu) ;
Celebes (Lompa-Battau, 3,000 ft., Tjamba and Bantimoeroeng in the south ; Menado ; Loka,
Bonthain) ; Halmaheira (Gilo); Morty; Ternate ; Batjan (Labuan); Wakollo, Central
Buru ; Ilat, Buru East Coast ; Mysol ; Kei Islands ; New Guinea (Humboldt Bay ; Kapaur ;
Dore ; Run). Length 12-24 mm.

In the large and representative collection now before me I find it impossible to subdivide
the species satisfactorily into groups distinguished hy the amount of puncturing on the
abdominal sterna. There is, however, a marked though imperfect correlation of the extent
of this puncturing with the localities from which the specimens come, specimens with smooth
sterna being characteristic of Ceylon, the Andamans and Nicobars, the Philippines, and the
Archipelago east of the Sunda Islands. In specimens from Java the abdominal sterna are
as a rule less extensively punctured than in specimens from Sumatra and Borneo ; but
specimens with absolutely unpunctured abdominal sterna do occur in Borneo and in
small islands near Sumatra, if not actually on the mainland.

The form of the mesosternal scars is also variable. Normally they are rounded on the
inner side, and are not very large ; but in specimens from the archipelago east of Borneo
the inner side is usually straight, extending much further backwards. Such forms also
occur further west, though more rarely. Celebes specimens appear to occupy a somewhat
intermediate position.

In very small specimens. from the Archipelago east of Celebes, which are usually
extremely flat ike L. planus, the frontal ridges resemble more or less closely those of
L. glaber, ending behind the anterior margin of the head and usually between its inner and
outer tubercles ; and the marginal tubercles of often closely approximated. When a series
of specimens is examined, however, this character also proves to be somewhat indefinite,
and I am no longer able to regard L. obtusidens, Kuwert, as distinct.

The size of the punctures in the lateral grooves of the elytra 1s very variable both in
L. bicolor and in L. dentatus. As a rule it is much smaller in the former than in the latter,
but the difference in the case of extreme specimens is very small. There is never any
difficulty, however, in distinguishing the two species from each other, by the structure of
the parietal ridges, which extend to the supra-orbital ridges in L. bicolor, but end abruptly
about half way between the central tubercle and the supra-orbital ridges in Z. dentatus.

Q 2

116 Memoirs of the Indian Museum. (Vor. VII,

Leptaulax anibarbis, Kuwert.
Leptaulax anibarbis, Kuwert, p. 293.
Leptaulax macassariensis, subsp. anibarbis, Gravely, 1914c, pp. 256 and 305-306, pl. xiii, fig. 54.

One specimen from Mt. Kinabalu. Length 22-2 mm,

Leptaulax cyclotaenius, Kuwert.

Leptaulax angustifrons + cyclotaenius + himalayae, Kuwert, 1898, pp. 285-286,
Leptaulax anıpunctus, Zang, 1905a, pp. 234-235.
Leptaulax cyclotaenius + anipunctus, Gravely, 1914c, pp. 255-257, pl. xi, figs. 53 and 55.

A number of specimens of the typical form from the following localities :—Margherita,
Assam; Xieng Khouang, Tenkin and Cambodia (presented by M. Vitalis de Salvaza) ;
Perak, Malay Peninsula (also specimens from Gap, 2,700-3,000 ft., Selangor-Pahang
Boundary, and foothills of Gunong Hitam, Selangor, Malay Peninsula, submitted by
Mr. C. Holman-Hunt); Sumatra (Kandg. Ampat, Lower Padang; Gunung-Agung,
Palembang, 5,000 ft. ; S. E. Serdang, ca. 1,000 ft. ; Engano Island, Benkulen Residency ;
also specimens from Medan, Sumatra, submitted by W. Guy Babault) ; Borneo (Mts.
Kinabalu and Marapok) ; and North Celebes (Tondano, Minahassa ; Toli-Toli).

Also several specimens of the variety anipunctus, Zang, from Chapa and Lao Kay,
‘Tonkin, and from Cambodia, presented by Mr. Vitalis de Salvaza.

Length 11-7-20°5 mm.

The varietal form anvpunctus differs from the typical form only in having the pronotum
somewhat sparsely, though extensively, punctured in the anterior angles and round about
the scars, instead of densely punctured at the sides from end to end ; and in having the
posterior intermediate areas of the metasternum somewhat weakly punctured on the inner
side only.

The structure of the head is very variable. The parietal ridges are usually long as in
L. bicolor in small specimens, and short as L. dentatus in larger ones. The latter usually
have the surface of the head densely punctured and the frontal area longer than broad ;
the former usually have the surface of the head more or less unpunctured and the frontal
area broader than long. Very small specimens are usually extremely flat like L. planus,
larger ones being somewhat stouter ; this is the case in L. bicolor also.

Leptaulax dentatus (Fabricius).
Passalus dentatus, Fabricius, 1792, p. 241.
Leptaulax dentatus + var. glabriventris, Gravely, 1914c, pp. 252-255, pl. xiii, figs. 52-52d,
Numerous specimens from the following localities :—Madras; Nepal; Darjiling
District (Tukvar, Van de Poll collection ; Singla, presented by H. E. Lord Carmichael ; and
Kalimpong, presented by myself); Tonkin (Lao Kay, Vientiane, Hoabink, Napé), Laos
(Kham-Keut) and Cambodia (Kompong Kedey) submitted by M. R. Vitalis de Salvaza ;
Renong, Siam; Karen Hills, Burma, 4,000 ft.; Andamans (Port Blair, presented by
Mr. 8. W. Kemp); Penang; Perak, Malay Peninsula; Carey Island (presented by
Mr. C. Holman-Hunt); Hili Madjedja, N. Nias; Kalim Bungo, Middle Nias ; Sumatra
(Médan, submitted by M. Guy Babault ; Tanjond-Djati, ca. 2,000 ft. and Mana-Riang,
2,000-3,000 ft., Renau, Palembang; 8S. EH. Serdang, E. Coast, ca. 1,000 ft.; Bedagei

1918.] F. H. GRAVELY : Passalidae of the World, 117

Interior, East Coast, ca. 600 ft.; Kandg. Ampat, Lower Padang) ; Java (Malang ; Tengger
Mt., 4,000 ft. ; Tjicopo; Boeloe Lawang, Pasoeroean ; Senggoro, southern Pasoeroean ;
Central Java, 1,500 ft.); Bali; Borneo (Mts. Kinabalu and Marapok ; Doesonlanden ;
Martapura, S. E. Borneo) ; Philippines (Mindoro ; 8. Palawan; Balabac: also specimens
presented by Mr. C. F. Baker from Imugin, N. Vissale; Mt. Makiling and Limay, Luzon ;
Thgan, Mindanao ; Mt. Banalao ; and Los Banos) ; Taruna, Great Sangir ; Celebes (Tondano ;
and Tangari, Minahassa, Menado and Toli-Toli in the north ; Bonthain, Bua-Kraeng 5,000
ft., Tjamba and Bantimurang in the south) ; Sapit, Lombok, 2,000 ft. ; Buru (North Coast ;
Kajeli; That, East Coast; Wae Kibo; Tifu Bay); Hitu, Amboina; Ceram (Wahaai ;
Rumasosal-Pasania ; Kairatoe) Buano; Nus Laut. Also a specimen said to come from
British Honduras, and others from the following localities which I have been unable to
trace :—Sula Besi (Doherty); Labunarang, Andonara (Doherty) ; Pach. (Mouhot) ; Mat.
(Wallace). Length 17-5-32:7 mm.

The puncturing of the abdominal sterna is very variable and proves, as in L. bicolor,
to be of no use for the distinction of definite varieties—hence the name glabriventis becomes
a synonym. The central area of the metasternum bears at most a pair of symmetrically
placed punctures. It never bears irregular punctures such as are ordinarily characteristic
of L. cyclotaenwus.

Leptaulax timoriensis (Percheron).

Passalus timoriensis, Percheron, 1841, pp. 19-21, pl. Ixxviüi, fig. 1.
Leptaulax timoriensis, Zang, 1905c, p. 223.

Three specimens from Gng. Leo, Dutch Timor, 2,000-4,000 ft.; five from Dilli, Port
Timor, 2,500 ft. ; one from Ilwaki, Wetter ; and two from the Alor Islands. Length 24-35
mm.

This species is very near L. dentatus, being distinguished only by the structure of the
pronotum, which is less distinctly rectangular, and is unpunctured in the anterior angles,
except in very small specimens in which one or two punctures may be present in this
position. In small specimens the puncturing in and around the pronotal scars and marginal
grooves is more extensive than in large ones.

Leptaulax anna, Zang.
Leptaulax anna, Zang, 1905a, p. 316.

Four specimens from B. Aru Hassa, Sambawa, 2,000-5,000 ft. Length 30-31 mm.

L. anna is very like L. timoriensis, but has the pronotal scars less densely punctured,
has the elytra more distinctly broadened behind with their lateral grooves matt and
marked with somewhat worn-looking punctures, and has the metasternum hairy laterally
and in front. In the Van de Poll specimens (30-31 mm. long) the lateral and intermediate
areas of the metasternum are united ; but in a smaller specimen in our collection (26 mm.
long and a co-type) they are distinct, though the ridge between them is somewhat weak
behind. The elytra are separate.

il
|

118 Memoirs of the Indian Museum. . [Voz.. VII,

Leptaulax humerosus, Kuwert.

Leptaulax humerosus, Kuwert, 1898, p. 289.
Leptaulax humerosus, Gravely, 1914c, pp. 251-252, pl. xin, fig. 51.

Numerous specimens from the following localities :—Perak, Malay Peninsula ; Sumatra
(Mana-Riang, 2,000-3,000 ft., and Tandjong-Djati, ca. 2,000 ft., Renau, Palembang ;
Bng. Proepoe, Padang Interior, ca. 6,000 ft.; 8. E. Serdang, ca. 1,000 ft. and Bedagei
Interior, ca. 600 ft., Hast Coast ; Beloe Lawang, Pasoeroean) ; Java (Malang ; Pengalengan,
S. Preanger, 4,000-5,000 ft. ; G. Gedeh, N. W. Preanger, 4,000 ft. ; Telaga Bodas, Garoet,
Preanger, 4,000-5,000 ft.; G. Tji Salimar, W. Preanger, 3.000 ft.; Tengger Mountain,
E. Java, 4,000 ft.) ; Borneo (Martapura and Kinabalu). Length 15-8-22°5 mm.

Easily distinguishable from L. anna, which it resembles as regards the sculpturing of
the elytra, by its smaller size, by its more strongly rectangular pronotum with thickly
punctured sides and more or less prominent anterior angles, and by its slenderer elytra.

The species of Leptaulax which I have been able to recognize may be distinguished
from one another thus :—

The elytra polished throughout

1 4 The depressed surface of the two or three outermost grooves
of the elytra dull, the punctures somewhat worn-looking 6 se IN

The puncturing of the lateral grooves of the elytra not strongly
transverse ; the parietal ridges united with the supraorbital
ridges = Be LE OY ae je on 3.
transverse ; or, the parietal ridges ending more or less

2\ The puncturing of the lateral grooves of the elytra strongly
abruptly about half way to the supraorbital ridges a 7 8.

The abdominal sterna covered eveniy all over with somewhat
obscure, broad, shallow punctures .. . Z. planus, p. 113.

34 The puncturing of the abdominal sterna arabe in estent,

sometimes absent, when present always finer, and when

extensive deeper and less uniform .. 54 5% = oo 4,

puncturing more or less obsolete ; the sides of the pronotum
and the posterior intermediate areas of the metasternum

at most weakly punctured

The marginal grooves of the pronotum coarser, strongly
punctured ; the sides of the pronotum and the posterior
intermediate areas of the metasternum as a rule strongly

The marginal grooves of the pronotum extremely fine, their
4

and extensively punctured 3 oS 55 3. BK Us

The pronotum strongly rectangular, the elytra more or less
parallel-sided ; small insects, not more than 18 mm. long .. 46 6% 6.
52 The sides of the pronotum somewhat rounded, the elytra more
or less dilated behind ; large insects, not less than 24 mm.
long .. 30 = 50 .. L. sambawae, p. 114.

1918.]

various species of Passalidae.
together in my “ Account of the Oriental Passalidae ?

The frontal ridges ending in the inner marginal tubercles ;
insects always unicolorous above ?
The frontal ridges extending parallel to the anterior margin

[
‘ of the head to a point between the inner and outer marginal
tubercles, where they end somewhat abruptly ; the anterior
parts of the elytra commonly reddish brown in otherwise
black insects .. Ne bas He :
[ Large insects (over 28 mm. long); the puncturing of the
pronotum and metasternum very scanty :
Smaller insects (not more than 25 mm. long) ; the puncturing
of the pronotum and metasternum much denser on
The parietal ridges united with the supraorbital ridges ; the
| central area of the metasternum unpunctured
The parietal ridges ending more or less abruptly about half
way to the supraorbital ridges ; or, the central area of the
metasternum punctured . co
” The central area of the metasternum almost invariably ah
at least a few more or less irregularly placed punctures ; the
parietal ridges variable a
The central area of the metasternum with at most one pair of
symmetrically placed punctures ; the parietal ridges always
ending more or less abruptly about half way to the
supra-orbital ridges eS
( The pronotum strongly rectangular ; its anterior angles more or
| less extensively punctured of si
The pronotum more rounded ; its anterior angles unpunctured,
except in small specimens. where a small group may be
present ‘€ =:
‘ The grooves of the elytra not tuberculate
114 A more or less distinct polished tubercle formed out of each
of the transverse ridges in the lateral grooves of the elytra. .
The pronotum convex, punctured only in the scars and
marginal grooves and usually in the anterior angles ; the
elytra short, dilated behind; the metasternum hairy
laterally and in front, its lateral and intermediate areas
124 often united
The pronotum somewhat flattened, densely punctured laterally,
its sides practically straight ; the elytra slender, more or less
parallel sided ; the metasternum hairless, the intermediate
and lateral areas always distinct

ZOOGEOGRAPHICAL RESULTS.

F. H. GravELy : Passalidae of the World.

119
L. roepstorfi, Kuwert,
L. glaber, p. 113.
L. barbicauda, p. 114.
L. bicolor, p. 114.
L. anibarbis, p. 116.

9.
L. cyclotaenius, p. 116.

10.
L. dentatus, p. 116.
L. timoriensis, p. 117.

12.

L. beccarii, Kuwert.

L. anna, p. 117.

L. humerosus, p. 118.

It would be useless to attempt to give here a detailed account of the distribution of the

For in the case of Oriental genera the information gathered
can readily be supplemented by the

120 , - Memoirs of the Indian Museum. . [Vor. VII,

additional records contained in the present paper; and in the case of other genera no
compilation is possible without.a much more detailed revision of synonymy than I am at
present able to achieve. But the general distribution of the family requires some further
consideration in the light of certain facts set forth in the present paper.

It will be convenient to deal with the Indo-Australian area first.

This area is inhabited by three subfamilies of Passalidae, namely the Aulacocyclinae,
Macrolininae and Leptaulacinae.

The Aulacocyclinae, though not a very large subfamily, appears to be a somewhat
highly specialized one. In none of its species are there frontal and parietal ridges or inner
and outer tubercles, such as are found in the more primitive species of all other subfamilies ;
and in the three largest genera, Comacupes, Taeniocerus and Aulacocyclus the basal piece
and lateral lobes of the male genital tube form one piece, either by consolidation or by the
suppression of the basal-piece, instead of being separate as in other subfamilies (see Sharp
and Muir, 1912, p. 580; also above, p. 5), while the middle lower tooth on each mandible
is immovable. In all other Passalidae, even in such primitive forms as O\leoides
subrecticornis, this tooth is jointed. Jointing does not occur, so far as I know, in any
beetles other than Passalids, and is clearly an indication of specialization ; but its absence
in Comacupes, Taeniocerus and Aulacocyclus is probably secondary and not primitive,
especially as it is correlated with specialization of the male genital tube. In the two
remaining genera, Ceracupes and Cylindrocaulus, the structure, both of the tooth in question
and of the male genital tube, resemble those found in other subfamilies.

The largest genus, Aulacocyclus, is centred in the Australian Region, but extends into
the Sunda Islands and Indian Peninsula. This discontinuous distribution suggests that
ground is being lost in the Oriental Region, where the smaller genera Comacupes and
Taeniocerus predominate. These genera are confined to the Oriental Region, except for a
single species of Comacupes (C. foveicollis) which has established itself in Celebes. Only
one species, Taeniocerus bicuspis, is found north of the Malay Peninsula ; this extends.
northwards to the Himalayas.

The genera Ceracupes and Cylindrocaulus, in which the male genital tube and middle
lower tooth resemble those of other subfamilies, only occur towards and beyond the
northern confines of the Oriental Region. With these presumably primitive characters
they combine cephalic excrescences which give them a most unusual appearance. Such
excrescences frequently indicate the senility of a group, and it seems probable that
Ceracupes and Cylindrocaulus are senile survivors of a transitional group through which
the more typical Aulacocyclinae of the present time have been derived. Ceracupes is less.
abnormal than Cylindrocaulus and occurs in Burma, the Himalayas, Tonkin and, Formosa.
The latter only occurs still further north, namely in China and Japan. Its species are the.
only Aulacocyclinae known to have fused elytra.

The Macrolininae fall into two series of genera, whose distribution must be considered.
separately. The first of these comprises the genera Macrolinus and Pleurarius, whose
combined range covers the Oriental Region and Celebes, but does not extend into the
Papuan Sub-Region. Ceylon is occupied by species of Macrolinus which are closely allied:
to one another but differ in certain characters, common to all of them, from the remaining

1918.] F. H. GRAvELY : Passalidue of the World. 121

species of the genus.! The genus Pleurarius appears entirely to replace Macrolinus in the

Indian Peninsula. This genus has otherwise been recorded only from Sumatra, whence-

it was originally described. In the absence of further records from that island I am
inclined to doubt the validity of the record and to believe the genus to be confined to the
Indian Peninsula, If this is so the genus probably contains one species only, a species
whose elytra are united. Other groups of Macrolinus occupy respectively (1) the
Indo-Chinese Sub-Region, (2) the Malayan Sub-Region and (3) Celebes, except that one
rare Cebelean species belongs to the Malayan group. Species of Macrolinus with fused
elytra are known only in the Ceylonese and Celebean groups.

The second series of genera of Macrolininae (Pl. I) is found throughout the Indo-Aus-
tralian area and is remarkable for the pronounced asymmetry which is developed in most of
its more highly specialized members. It comprises the Aceraiinae and Gnaphalocneminae of
my previous papers, one of which was devoted to a special study of its distribution (19140).

The study of more extensive material fully confirms the geographical separation, in
Ceylon and Australia respectively, of the primitive and closely related symmetrical forms
by the more highly specialized and less closely related descendants of each ; but shows that
I was mistaken in confusing the Celebean Passalid fauna with the Papuan, and that my
suggestions regarding phylogeny can be improved upon.

Concerning the distribution of the genera Episphenus, Ophrygonius and Aceraius there
is nothing fresh to add. The first named is confined to the Indian Peninsula and Cevlon,
the two last to the rest of the Oriental Region. The species inhabiting Ceylon are less
highly specialized than those inhabiting the Indian Peninsula, which in their turn are less
highly specialized than those found on the other side of the Ganges, taking these as a whole.
And in each of these areas the most asymmetrical (7.e., the most highly specialized) is also
the most abundant, the most variable, and among the largest. It also has gregarious
habits (Gravely, 19145, pp. 202-204 ; 1914¢, pp. 311-313).

Similarly, in the genus Pelopides, the most abundant species in the Sunda Islands are
large and highly asymmetrical (P. tridens, etc.) ; but in the Malay Peninsula the most
abundant species (P. dorsalis) is smaller and more nearly symmetrical. The most symme-
trical species of all appears to be confined to Borneo, the island where a primitive form
would be least expected ; but 1t does not seem to be common there.

The genus Pelopides is found all over the Malayan Sub-Region, and extends beyond
it into the extreme south of Burma, but no further. Its connection with simpler genera
is obscure, but it would be quite in keeping with the general relationship between the
evolution and distribution of asymmetrical Passalids for some ancestral form to be found in
Continental Asia. It seems to me possible that such may be represented in the genus
Tiberioides, a symmetrical genus whose presence in the area occupied by Ophrygonius and
Aceraius does not accord well with any direct relationship with them. If this is the case,
the grooves on the mentum of T. borealis no doubt represent an early stage in the
development of the large secondary scars found in all species of Pelopides. Closely allied
to Pelopides is the genus Plesthenus, which is confined to Celebes (see above, p. 96).

1 For the distinctive characters of the several local groups of species of Macrolinus see sections 1-3 of the table on
pp. 82.84 above.

122 Memoirs of the Indian Museum. [Vor. VI,

The most primitive species of the Australian Region belong to the genus Mastochilus,
a genus which, like the Oriental Episphenus, contains both symmetrical and more or less
strongly asymmetrical species. The subgenera Pharochilus and Mastochilus, which with one
exception (M. pectinigera, Heller, from New Guinea) are confined to Australia, contain large
and robust symmetrical insects. The subgenera Analaches and Cetejus, which with one
exception (M. australiensis from Australia) are found in the islands north of Australia,
contain smaller and often slighter insects which are almost always more or less asymmetrical.
Of the two species of Æpisphenus inhabiting Ceylon the dominant one is slightly
asymmetrical, the other, which is symmetrical, being closely allied to it, but much less
abundant and of smaller size. In Australia, on the contrary, the various symmetrical
species are dominant, the asymmetrical Mastochilus (Analaches) australiensis being
comparatively rare; which suggests that M. ausiraliensis is a comparatively recent
importation and has not been derived directly from its symmetrical compatriots. This
suggestion is supported by the fact that M. australiensis is much more closely related to
Papuan than to Australian species, being indeed one of the most highly asymmetrical
members of its genus, and by the fact that it has been recorded from New Guinea as well
as from Australia.

The genus Mastochilus probably represents the primitive stock from which the genera
Labienus (with Protomocoelus), Gonatas and Pseudepisphenus (with Tarquinius) have been
derived.

In Labienus specialization affects mainly the metasternum, apparently in association
with the wings, which tend to lose their normal function and doubtless to become more
efficient stridulating organs at the same time. Inthe most highly specialized members of
the genus, which appear to be confined to the Moluccas, the elytra are united in the middle
line, species with separate elytra being apparently to be confined to New Guinea, the
Aru Islands, ete.

Protomocoelus appears to have been derived from the simpler forms of Labienus. Its
dentition is reduced, in which respect it is the most highly specialized of all the species
with a modified metasternum. But the elytra are not united nor do they show any tendency
to become ovate. The genus occurs in the Solomon Islands and has been recorded from the
Moluccas, as well as from the Islands inhabited by the simpler forms of Labienus.

The genus Gonatas constitutes a second line of descent from Mastochilus. The
metasternum, wings and elytra are always normal; but the mandibles become very
strongly asymmetrical, and the posterior margin of the mentum very strongly arched, in
highly specialized forms. The progressive stages of this development are still preserved in
the less highly specialized species. The most primitive species of all, G. minimus, appears
to be confined to New Guinea and its neighbouring islands; but @. pumilio, the species
most closely allied to it, although occurring there appears to be centred in the Moluccas.
Much larger and more abundant than either are G. scheliongi, G. germari and G. naviculator,
which must be regarded as the dominant species of the genus. G. schellongi is somewhat
more primitive than either of the others, and is confined to New Guinea, the other two
being centred in the Moluccas, though recorded from New Guinea and from Java and the
Philippines also.

1918. | F. H. Gravety: Pussulidae of the World. 123

The third and last line of descent from Mastochilus is found in the genera
Pseudepisphenus and Tarquinius, two extremely rare forms known only from New Guinea.
Their affinities have already been fully discussed elsewhere (Gravely, 1924c, 328-329).

The Leptaulacinae are centred in the Malayan Sub-Region, whence several have spread
westwards and eastwards to the Indo-Chinese Sub-Region and Celebes respectively. The two
dominant species, L. bicolor and L. dentatus, have spread beyond these limits into the Indian
Peninsula and Ceylon in the west, and into the Moluccas, New Guinea, and possibly even
Australia in the east. Isolated species have arisen in several of the islands or island groups
of both the eastern and western parts of the archipelago. The number of distinct species
appears, however, to be small and the more widely distributed species especially are
extremely variable and often difficult to distinguish from one another. They are also
extremely abundant. This probably indicates that the subfamily is of relatively recent
origin and that it has not yet reached a condition of equilibrium.

The importance of Palk Strait, the Gangetic Plain, the China Sea and Isthmus of Kra
(together), the Straits of Macassar and Torres Strait in the distribution of the Macrolininae
has already been pointed out (Gravely, 1914¢, p.338). The further study of the Passalidae
cf the Australian Region shows that the Molucca and Gilolo Straits are of no less
importance and, indeed, that to the former belongs the special importance which I
previously attached to the Straits of Macassar, the fauna of Celebes being even more
unlike that of the Australian Region than it is unlike that of the Oriental Region.

The Passalids hitherto recorded from Celebes are as follows 1—

ea Genus otherwise purely Oriental; ©. joveicollis,
Comacupes fovercollis subsp. minor

s. str., confined to Borneo.
Aulacocyclus celebensis re is Endemic. Genus Indo-Australian.
(Endemic. Belongs to the group of Maerolinus other-

( wise known only from the Malayan Subregion.

a duivenboder bis Eye )

DEEE ‚These two constitute a group which is endemic.
22 I Que O° oe

Macrolinus suleiperfectus

A Malayan species. The genus is so definitely
Oriental that I am inclined to doubt this record.
Endemic. Allied tothe Oriental Pelopides.

Aceravus laevicollis {
ee The only other known species of the

Plesthenus spp.

Trichostigmus ursulus ; É
genus is Oriental.

Leptaulax planus ..

: aan Oriental species which appear to be extending their

range. One at least has reached New Guinea
= cyclotaenius

fie and possibly Australia.

Although a large proportion of these species are endemic, and it is doubtful whether
either of the two most characteristic asymmetrical Oriental genera, Aceraius and Pelopides,
occur in Celebes at all, it will be seen that every species known from Celebes is related to
‘species which are essentially Oriental, although some have established themselves in the
Australian Region also.

1 Concerning Kuwert’s record of ‘‘ Pelops ” impressicollis see above, p. 107, footnote 2.

R 2

124 Memoirs of the Indian Museum. MOVE

The Passalid fauna of the Moluccas is closely allied to that of New Guinea, and several
species have been recorded as common to both. In the genus Gonatas, however, it is
noteworthy that of the two species with most primitive mentum the one with complete
dentition is only known from New Guinea ; while of the three common species with more
highly specialized mentum the one with the most primitive dentition seems to be confined
to New Guinea and the other two to the Moluccas. Similarly, in the genus Labienus,
species with normal elytra appear to be confined to New Guinea and those with fused elytra
to the Moluccas. The allied Protomocoelus, in which the mandibles are modified instead
of the elytra, belongs however to New Guinea, and, although it is undoubtedly more
widely distributed than any of its allies, the single record of its occurrence in the Moluccas
should be confirmed before it is finally accepted. Pseudepisphenus and Tarquinius are only
known from New Guinea.

The information at present available regarding the distribution of American and
Ethiopian Passalidae is much less satisfactory than that regarding the Indo-Australian
subfamilies. The probable distinctness of the American and Ethiopian Passalid faunas, in
spite of several records to the contrary, has already been dealt with (above, pp. 10-11). It
is perhaps worthy of note here that no Ethiopian Passalidae are known to have the elytra
united, and that in America, although species with fused elytra attain the largest size, the
commonest and most widely distributed species have separate elytra. Among the
Pseudacanthinae Popilius cornutus is the largest and most highly specialized of the species
with separate elytra and is the commonest and most widely distributed species in the
subfamily. Among the Proculinae no species appears to be exceptionally abundant. Among
the Passalinae Paxillus leachii, Passalus interstitialis and Passalus interruptus are particularly
abundant and widely distributed. The last named is probably the most abundant and
widely distributed of all, and is also extremely variable. The group of species to which it
belongs appears to me to be the culminating point of the general trend of evolution through-

-out its genus, a genus whose wealth of closely interrelated species suggests that it bears the
‚same kind of relation to the rest of the American Passalid fauna as Leptaulax does to the rest
of the Indo-Australian.

SUMMARY.

1. External Morphology.

The clypeus 1s exposed and separated by a suture from the frons only in the subfamily
Pseudacanthinae. In a few other genera, mostly American, it is exposed but united
to the frons. In the majority of Passalids the whole of the upper surface of the anterior
part of the head, between the supra-orbital ridges and in front of the frontal ridges, is frons,
the whole of the clypeus being doubled beneath this out of sight (pp. 1-3, fig. 1, 1-4).

It is uncertain whether the inner and outer marginal tubercles of the Leptaulacinae
are homologous with the inner and outer tubercles respectively of other Passalidae
(pp. 3-4).

The dentition is reduced only in somewhat highly specialized forms. In American
subfamilies it seems to be associated with the loss of the habit of flight, and to come about

1918.] F. H. Gravety: Passalidae of the World. 125-

through the fusion of the two lowest terminal teeth. In Indo-Australian subfamilies it is
always associated with cephalic asymmetry and never with the loss of the habit of flight,
and comes about through the fusion of the anterior lower and lowest terminal teeth
(pp. 9-10, fig. 11).

The loss of the habit of flight appears to allow of greater specialization of the wings as
stridulating organs. It produces definite structural modifications in the insect (pp. 4-5).
The following genera contain, so far as is known, only flightless forms :—Cylindrocaulus,
Platyverres, Pleurarius, Proculejoides, Proculejus, Procululus, Proculus, Pseudacanthus
and Publius. The following species are also flightless :—Labienus moluccanus and gigas,
Macrolinus obesus and ursus, Passalus quitensis and Vindex synelytris.

The structure of the male genital tube is almost uniform throughout the family.
The genera of Aulacocyclinae other than Ceracupes and Cylindrocaulus differ, however, from
the rest of the family in that the basal piece and lateral lobes are represented by one
undivided plate (p. 5).

The central tubercle is usually larger in females than in males in species in which
it varies greatly in size (p. 5).

2. Classification.

Seven subfamilies have been recognized, of which one, the Aulacocyclinae, confined to
the Indo-Australian area with China and Japan, is somewhat widely removed from all
the others (p. 9). Two others are confined to the Indo-Australian area. These are
distinguished from American and Ethiopian subfamilies by the structure of the mandibles
(p.9). The Ethiopian subfamily is distinguished from the four American ones by the
structure of the anterior margin of the head (pp. 10-11).

The number of genera has been greatly reduced. Specific synonymy has not been dealt
with, but there can be little doubt that a similar reduction is required in the number of
species.

3. Geographical Distribution.

Passalidae appear to flourish only under more or less moist tropical conditions.

American, Ethiopian and Indo-Australian forms belong respectively to different
subfamilies, probably without exception (pp. 9-12).

The group of Macrolininae with strong asymmetrical tendencies is of special zoogeo-
graphical interest. Its most primitive species inhabit Ceylon and Australia. These
are closely allied to one another but give rise to divergent lines of descent, confined
respectively to the Oriental Region with Celebes, and to the Australian Region. Both these
Tegions are composed of a series of smaller areas, each characterized by a distinct Passalid
fauna, which is more highly specialized in those nearer to Celebes than in those further
away. These areas are: in the Oriental Region—Ceylon, the Indian Peninsula, the
Indo-Chinese Subregion and the Malayan Subregion; and in the Australian Region—
Australia, New Guinea and the Moluccas (pp. 120-124). These facts bear out the
suggestion (Gravely, 1913, p. 204) that conditions towards the centre of the Archipelago are
peculiarly favourable for evolution, and that as more and more highly specialized forms
have arisen there, they have migrated outwards, driving before them the less highly

126 : Memoirs of the Indian. Museum. : [Vor. VII,

specialized, which have rarely survived except where they have been able to establish.
themselves behind some obstacle to migration.

The fauna of Celebes, though related to the Oriental fauna, is very distinct from it, and
contains a large proportion of endemic species and one endemic genus (p. 123).

The genera Macrolinus and Pleurarius are Oriental. The former genus has produced
local races in Celebes and in each of the areas into which the Oriental Region proper has been :
divided above, with the single exception of the Indian Peninsula where it is replaced by the
latter genus (pp. 120-121).

The Leptaulacinae appear to be centred in the Malayan Subregion and to be undergoing |
rapid development and expansion (p. 123).

The Aulacocyclinae appear to be a very highly specialized subfamily, now on the
decline. The genera Ceracupes and Cylindrocaulus appear to be the senile representatives :
of an old group, in some respects more primitive than the forms at present dominant in the
subfamily. They are only found towards and beyond the limits of distribution of the rest
of the subfamily (p. 120).

The Solenocyclinae appear to be peculiar to the Ethiopian Region, and the Pseuda-
canthinae, Proculinae and Passalinae to America. The information at present available
as to their distribution is much less complete than is that available concerning Indo-
Australian forms (pp. 5 & 124).

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Stoliczka, F. ‘ A contribution towards a Monograph of the Indian Passal-
idae.” J. A. S. B. XLII (IN), 1873, pp. 149-162:

Kirsch, T. (a) “ Neue Käfer aus Malacca.” Mit. K. Zool. Mus. Dresden,

I, Was io. re.

(b) “ Beitrage zur Kenntniss der Coleopteren-Fauna von Neu
(Gnome Unten JX, Zool, Visa Dresden Wl, Use, D.
135-161.

Harold, E. von. Matth. Munch. Ent. Ver., p. 101.

Kirsch, T. “ Zwei neue Coleopteren-Arten aus Neu Guinea.” Ann. Mus.
Cw. Genova, XIV, pp. 18-20.

Fairmaire, L., in Le Naturaliste, 1880, p. 164.

Harold, F. von. ‘ Beschreibungen neuer, auf seiner, von der Akademie
unterstützten Reise in Ostafrika, vorzüglich in den Districten von Taita.
und Ukamba auf einer Tour von Mombassa nach dem Kenia, von Hrn.
J. M. Hildebrandt gesammelter Coleopteren.” Monatsber. R. Preuss. Ak.
Wiss. Berlin, 1880 (1880-1881), pp. 260-270.

1918.]

1883.

1884.

1885.

1886.

1887.

1889.

1890.

1891.

1892.

1896.

1897.

1898,

1898.

1900,

F. H. Gravety : Passalidae of the World. 129

Lewis, G. “ Lucanidae of Japan.” Trans. Ent. Soc., London, 1883, pp.
333-342, pl. xiv.

Wystmann, P. “Catalogue Systematique des Passalides.” Ann. Civ. Mus.
Genova, (2a) I (XXI), 1884, pp. 326-348.

Kirsch, T. ‘‘ Neue südamerikanische Käfer, III.” Berlin Ent. Zeitschr.,
XXIX, 1885, pp. 207-224, pl. ix c (Passalidae, pp. 208-210).

Schaufuss, L. W. “ Beitrage zur Fauna der Niederlandischen Besitzungen
auf den Sunda-Inseln.” Horae Soc. Ent. Ross., XIX, 1885, pp. 183-209.
Aurivillius, C. “ Insekter insamlade pa Kamerunberget af G. Waldau och

K. Knutson. I. Coleoptera: Cetoniidae et Lucanidae.” Bih. K. Sv.
Vet.-Ak. Handl., XII (4) 1, 18 pp., 3 text-figs.
Bates, H. W. Biologia Centrali-Americana. Insecta. Coleoptera IT (2)
Pectinicornia and Lamellicornia, 1886-1890 (Pectinicornia, 1886).
Fairmaire, L. “ Coleopteres de | ’Interieure de la Chine.” Ann. Soc. Ent.
Belg., XXXI, 1887, pp. 87-136.
Bates, H. W. Biologia Centrali-Americana. Insecta. Coleoptera II (2)
Pectimicornia and Lamellicornia, 1886-1890 (Supplement 1889).
Kuwert. (a) “ Einige neue Passaliden.” Deutsche Ent. Zeitschr. 1890,
pp- 97-104.
* (b) “ Die Passaliden und ihr Monographie von Kaup.” Soc. Ent.
V, 1890, pp. 1, 9 and 17.
Kuwert, A. “ Systematische Uebersicht der Passaliden Arten und Gat-
tungen.” Deutsche Ent. Zeitschr., 1891, pp. 161-192.

(?)Tryon, H. “Coleoptera collected by Mr. A. C. English in the St. Joseph
river district of British New Guinea, under the auspices of his Honour the
Administrator.” Rep. Administrator Brit. N. Guinea, Il, app. v, pp.
109-112.

Blackburn, T. “ New Genera and species of Australian Coleoptera, XX.”
Trans. R. Soc., S. Australia, XX, pp. 233-259.

Heller, K. M. “Neue Kafer von Celebes.’ Abh. u. Ber. d. K. Zool. u.
Anthr.-Ethn. Mus. zu Dresden, VI, 1896-7 (3), 24 pp., 1 pl.

Kuwert, A. ‘Die Passaliden dichotomisch bearbeitet ; 1.” Novit. Zool.
III, 1896, pp. 209-230, pl. v-vii.

Casey, T. L. “ Ooleopterological Notices, VII.’ Ann. N. York Ac. Sci.
IX, 1896-7, pp. 285-684.

Kuwert, A. “ Die Passaliden dichotomisch bearbeitet ; IL.” Novwit. Zool.
IV, 1897, pp. 274-306.

Heller, K.M. ‘ Neue Kafer von Celebes.”” Abh. u. Ber. K. Zool. u. Anthr.-
Ethn. Mus. Dresden, VII, 1898-9 (3), 42 pp., 1 pl.

Kuwert, A. ‘“ Die Passaliden dichotomisch bearbeitet ; LIT and IV.” Nonit.
Zool. V, 1898, pp. 137-205 and 259-349.

Blackburn, T. “ On some new Genera and Species of Australian Coleoptera.’”
Proc. R. Soc. Victoria XII, pp. 206-233 (Passalidae, pp. 207-211).

130

1900.

21902
1902.

1903.

1904.

1905.

1906.

1907.

1910.

Memovrs of the Indian Museum. | [Vor.. VII,

Heller, K. M. ‘ Neue Käfer von Celebes.”” Abh. u. Ber. K. Zool. u. Anthr.-
Ethn. Mus., Dresden, IX, 1900 (5), 46 pp.

Fruhstorfer. ““ Tagebuchblättern.” Insekten-Börse XIX, 1902, p- 28.

Rosmini, O. “* Viaggio del Dr. Enrico Fresta nella Republica dell’ Eucadore
regioni vieine—Passalidi.”” Boll. Mus. Torino XVII, 1902, no. 428, 10 pp.

Fauvel, A. “ Faune Analytique des Coléoptéres de la Nouvelle Calédonie.”
Rev. Ent. franc. XXII, 1903, pp. 203-378.

Zang, R. (a) “ Vorlaufige Diagnosen neuer Indo-Australischer Passaliden ”

Insekten-Börse, 20 Jahrg., No. 43, pp. 338-9.
(b) “ Bemerkungen zur Alteren Passaliden-Litteratur.”” Deutsche
Ent. Zeitschr., 1903, pp. 417-420.
Zang, R. (a) “ Parapelopides und Ophrygonius, zwei neue Gattungen der
Passaliden (Coleoptera).” Zool. Anz. XXVII, 1904, pp.
694-701, 3 text-figs.
(b) “ Ueber einige Passaliden,’ Tigjdscht. v. Ent. XLVII,
pp. 181-5.
Pangella, G. (a) ‘ Passalidi di Costa Rica.” Boll. Mus. Torino XX, 1905,
No. 498, 12 pp., 1 text-fig.
(b) “ Viggio del Dr. Alfredo Borelli nel Paraguay e nella Re-
publica Argentina—Passalidi.” Boll. Mus. Torino XX,
1905, No. 508, 16 pp.

Zang, R. (a) Numerous papers in Deutsche Ent. Zeitschr. for 1905.

(b) “ Anderung in der Nomenclatur der Passaliden (Coleoptera).”’
Zool. Anz. XXIX, 1905, pp. 154-5.

(c) Passalidarum Synonymia. Kritische Revision der von Kuwert
und anderen Autoren aufgestellten Gattungen und Arten.”
Notes Leyden Mus. XXV, 1905, pp. 221-232.

(d) ** Zwei neue Passaliden aus den Gattungen Comacupes, Kp.,
und Aceraius, Kp.” Notes Leyden Mus. XXV, 1905, pp.
233-8.

Pangella, G. ““ Spedizione al Ruwenzori di 8. A. R. Luigi Amedeo di Savoia
Duca degli Abruzzi. Nuovo specie di Passalidi (Diagnosi preventiva).”’
Boll. Mus. Torina, XXI, 1906, No. 540, 1 p.

Zang, R. (a) “ Uebersicht der Basilianus-Arten.” Deutsche Ent. Zeitschr.,

1906, pp. 177-183.

*(b) “ Passalini.” Nova Guinea, Résultats de (Expedition scien-
tifique Néerlandaise à la Nouvelle-Guinée en 1903, sous les
auspices de Arthur Wichmann, Chef de V Expedition. vd
pp. 23-26.

Arrow, G. J. “A contribution to the classification of the Coleopterous
Family Passalidae.” Trans. Ent. Soc., London, 1906 (1906-7), pp. 441-469.

Heller, K. M. “‘ Fünfter Beitrage zur Papuanischen Käferfauna.” Abh.
u. Ber. K. Zool. u. Anthr-Ethn. Mus. Dresden, XIII, 1910 (3), 42 pp., 1 pl.

1918.]

LOTS

1912.

1913.

1914.

1915.

1916.

F. H. GRAVELY : Passalidae of the World. 131

Heller, K. M. “ Eine neue Ceracupes-Art aus Formosa.” Ann. Soc. Ent.
Belg. LV, 1911, pp. 256-7, 1 text-fig.

Sharp, D. and Muir, F. “ The Comparative Anatomy of the Male Genital
Tube in Coleoptera.” Trans. Ent. Soc., London, 1912, pp. 477-642, pl.
xli-Ixxvii (Passalidae, pp. 579-580, pl. xliv, figs. 11-13a).

Gravely, F. H. “Three Genera of Papuan Passalid Coleoptera.” Mitt.
Mus. Hamburg, XXX, 1913, pp. 103-112, 6 text-figs.

Casey, T. L. “ Miscellaneous Notes and New Species.” Memoirs on the
Coleoptera V (Lancaster Pa., 1914), pp. 355-378.

Gravely, F.H. (a) “ H. Sauter’s Formosa-Ausbeute—Passalidae.” Sup-

plementa Entomologica III, 1914, pp. 30-32.

(6) The Evolution and Distribution of certain Indo-Austra-
lian Passalidae, J. A. S. B., X, 1914, pp. 201-209.

(c) An Account of the Oriental Passalidae based primarily
on the collection in the Indian Museum. Mem. Ind.
Mus. III, No. 4, 1914, pp. 177-353, 3 pl.

Grieve, 8. ‘The Occurrence and Distribution of the Beetle Passalus uni-
cornis, Serv., in the Antilles and the Northern portion of South America.”
Proc. R. Phys. Soc. XIX, pp. 159-160.

Gravely, F. H. “Notes on the Habits of Indian Insects, Myriapods and
Arachnids.” Rec. Ind. Mus., XI, pp. 483-593, pl. xxii-xxv (Passalidae,
pp. 495-497.

Gravely, F. H. ‘Some Lignicolous Beetle-Larvae from India and Borneo.”
Rec. Ind. Mus. XII, pp. 137-175, pl. xx-xxii.

Heller, R. M. “ Die Käfer von Neu-Caledonien und den benachbarten
Inselgruppen.” Sarasin and Roux’s Nova Caledonia, A, Zool. II (3) pp.
229-364, pl. x-xi, 22 text-figs. (Passalidae, pp. 352-353).

Schultze, W. “A Catalogue of Philippine Coleoptera.” Philippine J. Sci.
XI (D), pp. 1-194 (Passalidae, pp. 154-156).

1918. | F. H. GRAVELY : Passalidae of the World. 133

INDEX.

Synonyms are printed in italics : page numbers referring to keys, descriptions, locality
or synonymic records, and figures are printed in ordinary type ; other numbers are in bold
face.

The summary of the paper on pp. 124-126 has been designed partly as a guide to the
whereabouts of the principal facts recorded. References to it are not included in this index.
References to the list of recently described genera and species on pp. 7-9 have similarly been

omitted.

Page.
abortivus (Passalus) 53, 60, 67.
Aceraiinae 67121
Aceraius 76, 79, 89 ete., 121, 128.

Aceraius group

12°

aequalis (Ophrygonius) 88, 89.

aequidens (Aceraius, Ophrygonius) 76, 88, 89.

affınıs (Passalus) 2, 55, 66.

agassizi (Arrox, Sertorvus) 33, 34, 35.
agnoscendus (Vindex) 10, 11, 43, 46, 47.
albertisı (Gonatas) 109.

Alococerus 51.

alterego (Erionomus, Zriosternus) 15, 76.
alutaceosternus (Aceraius) 90, 93.

amazonicus (Popilius)

Analaches ae 77, 9%, 101 etc., 103, 122.
andamanensis (Basilianus, Macrolinus) 80, 83.

angulatus (Passalus, Ptichopus) 68.

angustifrons (Leptaulax) 116.

anibarbis (Leptaulax) US; 116, 18),
anipunctus (Leptaulax) 113, 116.

anna (Leptaulax) : ul, HIS

antanarivae (Ciceronius, Solenocyclus) ale

A poneleides + IE

approximatus (Passalus, Solenocyclus) 70.

arrowi (Ceracupes) | 21.

Arrox 5 32, 33, etc.

aruensis (Aulacocyclus) 18, 20.

assamensis (Aceraius) 89.

assimilis (Passalus, Veturius) 34, 38, 39.
Aulocozyclinae .. 2, 5, 9, 12, 13 etc., 120.
Aulacocyclus 5, 13, 14, 17 etc., 120, 128.
Aurelius 200103.

Auritulus 18, 21.

austeni (Ceracupes) 21.

austeni (Tiberioides) 85.

24, 27, 29.

134 | Memoirs of the Indian Museum. [ VOL.

australasicus (Mastochilus s. str., Passalus)

australiensis (Cetejus, Mastochilus Analaches)

australis (Passalus, Pelops, Protomocoelus)

barbatus (Passalus, Pentalobus)
barbicauda (Leptaulax)

basalis (Comacupes, Passalus)
Basilianus

batesi (Macrolinus)

beccarii (Leptaulax)

bicanthatus (Passalus, Taeniocerus)
bicolor (Leptaulax)

bicornis (Passalus, Spurius)

bicuspis (Aulacocyclus, Taeniocerus)
bifidus (Nasoproculus, Pseudacanthus)
binomnatus (Passalus)

birmanicus (Ophrygonius) ..

boliviae (Veturius)

borealis (Chilomazus, Tiberioides)
borneanus (Aceraius)

brachyphyllus (Pleurarius)

brasiliensis (Passalus, Paxillus)
brevioripennis (Odontotaenius, Popilius)
bucerus (Cylindrocaulus) ..

burmeisteri (Eriocnemis, Gnaphalocnemis, Pelopides

Calidas ae
camerani (Pazillosomus, Paxillus)
cantori (Ophrygonius, Passalus)
carolinensis (Gonatas)

Cassius as

catherinae (Passalus)

Caulifer

cavicollis (Verres)

cavicornis (Aulacocyclus, Comacupes)
cayor (Passalus, Pentalobus)
celebensis (Aulacocyclus)

cephalotus (Passalus, Veturius)
cephalotus (Veturius)

Ceracupes

Cetejus

cetioides (Gonatas)

championi (Proculejoides, Proculejus) . .
Chilomazus

Chondrocephalus

Page.

99, 100, 103.

77, 98, 101, 103, 122.
107.

72, 13, 74.
112, 114, 119.
19,716:

80, 86.

81, 83.

119.

INS), Ir
112, 113, 114, 119, 128.
26.

16, 17, 120.
24, 30.

64, 68.

76, 87, 88.
34, 38, 39.
85.

905922

5, 82, 84.
45, 49, 50.
24, 28, 29.
21.

94, 95.

74.

48, 50.

86, 88.

108, 110, 111.
51.

53, 55, 66.

NBs Ile

34, 40, 41, 47.
115} 16,

11.

123.

37, 39.

37.

14, 21, 120.
Tey Dt, WO, iOS}, ep

108, 110.
10, 47.
85.

2, 10, 11, 48, 44 ete.

Vin;

1918.] F. H. Grave : Passalidae of the World.

Ciceronius

clypeatus (Malagasalus)

Comacupes ore ie a
compergus (Labienus, Passalus, Vellejus)
comptoni (Aceraius, Episphenus)
Coniger À N
convexifrons (Ophrygonius)
convexus (Passalus)

cordiger (Chondrocephalus)
cornutus (Passalus, Popilius)
corticola (Passalus, Verres)

crassus (Passalus, Publius)
crenatipennis (Macrolinus)
erenatus (Paxillus)

criniceps (Veturius)

crinitus (Heterochilus, Ophrygonius)
curtus (Passalus, Petrejus)..
eyclotaenius (Leptaulax)
cylindraceus (Comacupes, Passalus)
Cylindrocaulus ..

Cyphoproculus ..

dentatus (Leptaulax, Passalus)
depressus (Macrolinus)

deyrollei (Aulacocyclus, Tazntocerus) ..
dichotonus (Spurius)

Didimoides

Didimus h

differens (Gonatas) ;
dilatatus (Mastochilus Pharochilus, Passalus)
dohrni (Aurelius, Labienus)

dorsalis (Eriocnemis, Pelopides)
duivenbodei (Macrolinus)

dunsiriensis (Ophrygonius)

duplicatus (Didimus, Pentalobus)

edentulus (Aulacocyclus, Passalus)
Epeus

Epipertinax

Epiphanus

Epiphoroneus

Epipleurothria ..

Episphenoides

Episphenus

Eriocnemis

Erionomus

Page.

HONTE

69, 70.

14 etc., 120, 123.
104, 105, 107.
85, 86.

28, 29, 124.
34, 40, 41.
34, 42.

81, 83.

45, 50, 51.
3137039)
Sip

53, 56, 66.
113, 116, 119, 128.
14, 16.

14, 21, 120.
49,

112, 113, 116, 119, 123.

80, 81, 83.
19, 20.

24, 26.

69, 72.

72.

109.

97, 98, 103.
104, 106.
94, 95, 121.
82, 83, 123.
86.

74.

1216819220;

51.
Te
78, 85 ete., 121, 122.
93.
4, 10, 69, 74, ete.

135

136 Memoirs of the Indian Museum.

Eriopterus

Eriosternus

erosus (Passalus)
errans (Aulacocyclus)
eucadorensis (Passalus)
Eumelosomus

Eumelus

exaratus (Passalus, Solenocyclus)

felderi (Aulacocyclus, Comacupes)
flachi (Episphenus)

Flaminius

Flavius ies

foveicollis (Comacupes)
fronticornis (Ceracupes, Passalus)
fur (Didimus, Pentalobus)
furcilabris (Passalus, Verres)

gelon (Plesthenus)

germari (Aceraius, Gonatas)
gigas (Eriocremis, Labienus)
glaber (Labienus)

glaber (Leptaulax, Trichostigmus)
glaber (Passalus)

glaberrimus (Passalus)
glabriusculus (Aulacocyclus)
glabriventris (Leptaulax)
Gnaphalocneminae

Graphalocnemis ss
Graphalocnemis (—Pelopides) group ..
Gonatas

Gonatas group

goryi (Passalus, Proculus)
grabowskii (Mastochilus Cetejus)
gracilis (Labienus)

gracilis (Passalus, Petrejus)
grandis (Aceraius, Passalus)
granulifrons (Chondrocephalus, Popilius)
gravidus (Pelopides)

grayi (Semicyclus, Solenocyclus)
guatemalae (Popilius)
guatemalensis (Oileus, Passalus)

helferi (Aceraius)
Heliscus
Heterochilus

Page.

22, 31.

74.

53, 64, 68.
18, 19020:
59, 56, 66.

70, 71.

15, 120, 123.

34, 40, 41.

96. .

109, 111, 122.
106, 107.

106.

113, 119.

53, 64, 68.

58, 67.

17, 20.

113, 116.

76, 121.

76, 93.

12.

78, 80, 108 ete., 122, 124.
12.

42, 43.

102, 103.

107.

56, 66.

76, 89, 92, 93.
2, 10, 44, 45, 46, 47.
76, 93, 94, 95.
72.

24, 27, 29.

53, 57, 66.

89, 93.
22, 26.
76, 86, 87.

[| Vor.

VII

1918.] F. H. Gravery : Passalidae of the World.

heydeni (Passalus, Veturius)
himalayae (Leptaulax)
himalayensis (Aceraius)
hirsutus (Aceraius)

hostilis (? Erionomus, Passalus, Sonia ace late

humerosus (Leptaulax)
Hyper plesthenus
Hyperplesthenus group

illegalis (Aceraius)
impar (Labienus)

impressicollis (Mastochilus s. lat., Pee Balin

Protomocoelus)
inaequalis (Labienus)
inaequalis (Ophrygonius, Passalus)
incertus (Passalus)
incertus (Passalus, Rhodocanthopus)
incisus (Passalus, Undulifer)
indicus (Basilianus, Episphenus)
intergeneus (Popilius, Soranus)
intermedius (Platyverres, Verres)
interruptus (Lucanus, Passalus)
interstitialis (Passalus)
invitus (Plesthenus)

jalapensis (Pseudacanthus)
jansoni (Passalus, Phoroneus)
javensis (Ophrygonius)

Kaupioloides

Kaupioloides group

Kaupiolus

klugi (Leptaulax, Paseo)
kuwerti (Aceraius)

kuwerti (Tiberioides, Tiberius)

Labienus oe
laevicollis (Aceraius, Passalus)
laevimargo (Aceraius)
lamellatus (Aceraius)
languidus (Neleus, Passalus)
laniger (Aceraius)

Lasioperix

latifrons (Passalus)

latipennis (Macrolinus, Passalus)
leachu (Paxillus)

Lepiaulacides

Page.

34, 85, 38, 39.
113, 116.

89.

SELL:
77, 103.
12.

SIL, SB:
106.

107.

104, 105, 106, 107.
86, 87, 89.

57, 66.

51, 63, 68, 124.
53, 58, 67, 124.

27, 78, 79, 103 etc., 122,

91, 93, 123.

53, 54, 66.
80, 82, 83.
45, 48, 49, 51, 124.

124,

137

138

Leptaulacinae ..
Leptaulax

Leptaulax group
Lophocephalus ..

lottinii (? Mastochilus s. lat., Passalus, Plesthenus)

Lucilius

macassariensis (Leptaulax)
Macrolininae

Macrolinus

Macrolinus group

Macrolobus

major (Gonatas)

Malagasalus

Manlius ae
manouffi (Passalus, Solenocycius)
marginatus (Passalus, Popilius)

masoni (Comacupes)

mastersi (Aulacocyclus, Taeniocerus) ..

Mastochilus, s. lat.
Mastochilus, s. str.
Microthorax

minimus (Gonatus, Omegarius)
minor (Comacupes)

minor (Gonatas)

minor (Aceraius, Ophrygonius)
Mitrorhinus

mniszechi (Proculus)

moluccanus (Labienus, Passalus)

monticulosus (Gnaphalocnemis, Passalus, Pelopides)

moorei (Episphenus)

morbillosus (Passalus, Solenocyclus)

morio (Passalus)
Morosophus

möschleri (Aceraius)
mucronatus (Passalus)

nanus (Passalus, Rhodocanthopus)
Nasoproculus

nasutus (Passalus)

naviculator (Gonatas, Passalus)

neelgherriensis (Episphenus, Passalus)

Neleides
Neleidinae
Neleinae
Neleuops

Memoirs of the Indian Museum.

Page.

12, 13, 111-ete., 128.

3-4, 111, 112 ete., 123, 124.
12.

ole

1382, UG.

12, 13, 76 etc., 120-123.

51, 76, 78, 80 etc., 120-121, 128.
12:

Gil:

109.

8, 69 etc.

Silk

Ale

24, 26, 27,- 29.
I, TG

18, 20.

77, 78, 79, 97 etc., 122.
7, 100, 122.
51.

108, 110, 122.
123.

108, 110, 111.
76, 89.

10-11, 51.

10, 43.

106, 107.

95, 96.

85, 86.

71.

53, 54, 65.
51.

91, 93.

53, 60, 67.

52, 65.

22, 30.

53, 62, 68.

108, 110, 111, 122.
86.

[Vor. VII,

1918.] Kee GRAVELY :

Neleus :

nicobaricus (Macrolinus)

Ninoides de
nitidulus (Mastochilus Pharochilus)
nobilis (Trapezochilus)
novaebrittaniae (Gonatas)
novaeguineae (Leptaulax)

obesus (Macrolinus)

obliquus (Mastochilus s. lat.)

obtusidens (Leptaulax)

oceipitalis (Epiphoroneus, Passalus)
occulidens (Aceraius) 52
occulitesselatus (Hetzrochilus, Ophrygonius)
Odonotaenius

Oeneus

Ogyges

Oileoides

Oileus

Omegarius Mn
opaeipennis (Passalus, Proculus)
opacus (Passalus)

Ophrygonius

paradoxus (Tarquinius)
Parapelopides

Parapertinax BE
parastictus (Passalus, Pentalobus)
palini (Erionomus, Passalus)
parryi (Aulacocyclus)

parvicornis (Oileoides)
Passalotaenius ..

Passalinae

Passalus

patalis (Cylindrocaulus)
Paxilloides

Paxillosomus

Paxillus

pearsoni (Episphenus)

pectinigera (Episphenoides, de
Pelopides bs
Pelopinae

Pelops ; 52
peltostietus (Aceraius, Rested ant Cetejus)

Passalidae of the World. 139

Page.
51.

83.

51.

98, 103.
94.

109.
112.

80, 82, 83.
98.
12,
53, 61, 68.
76, 92, 93.

22, 30.

i, 2, GP, BS as, al
Ape, DS, 25:

78, 108.

43.

53 109 108:

76, 79, 86 etc., 121.

9, 12, 13, 43 ete., 124.
me AEE pil Chie, 124.

10, 26.

21.

48.

8, 11, 44, 48 eic., 60, 124.
85.

98, 122.

76-77, 79, 93 etc., 121, 128.
76.

76, 107.

102, 103.

140 Memoirs of the Indian Museum.

Pentalobus

pentaphyllus (Passalus, Paxillus)
perakensis (Aceraius)

perlatus (Aulacocyclus)

perplexus (Pseudepisphenus)
Pertinacides

Pertinacinae

Pertinax A

pertyi (Passalus, De

Petrejinae

Petrejoides

Petrejus

Phanocles

Pharochilus

Phaulothoraz

Phoroneinae

Phoronaeosomus

Phoroneus

Phraortes

pilifer (Aceraius, Passalus)

planiceps (Erionomus, Passalus)
planus (Leptaulax, Passalus) ;
platypus (Aulacocyclus, Taeniocerus) ..
platyrhinus (Passalus, Veturius)
Platyverres

Plesthenus

Plesthenus group

Pleurariinae (Gravely)

Pleurariinae (Kuwert)

Pleurarius

Pleurarius group

Pleurostylus 5
politus (Mastochilus onde Pate)
polli (Passalus)

Polyacanthopus =
polyphyllus ee a s. str., Passalus)
Popilius

Proculejoides

Proculejoides group oi Passalinae
Proculejus

Proculinae

Procululus

Proculus

prominens (Passalus) on 00
Prosochtus

Protomocoelus ..

Page.

4, 10, 69, 72 etc.

45, 48, 49, 51.

Oil, GEL

Wf US, ADs

LU,

pile

51.

DIE

53, 54, 66.

51.

22, 30.

Si,

DIE

77, 97, 98 etc., 103, 122.
bile

51.

93:

90593:

70, 74, 75, 76.

113, 118, 128.

168, Ie

30, 39:

3, 5, 82, 33, 41, etc., 51.
76-77, 79, 96 etc., 121, 128.
12.

76.

51.

5, 51, 76, 78, 82, 84, 120-121.

12.

30.

SR) 2108;

53, 62, 67.
51.

97, 100, 103.

1, 10, 22, 23, 26 etc., 58, 124.

10, 22, 33, 43, 44, 47 etc.
9.

10, 22, 23, 31 etc., 48, 47, 51.

2, 12, 13, 32 etc., 124.
33, 42.

9, 10, 32, 33, 42 etc.

57, 66.

22, 31, 51.

78, 79, 107, etc., 122, 124.

[Vor. VIL,

1918.] F. H. Gravety: Passalidae of the World. 141

Page.

Protomocoelus group ae er ao dee
Pseudacanthinae Ke er sq Bb UI, I, OP Er, 122
Pseudacanthus fh aa co 2B 90) ec. 47e
Pseudepisphenus Br ar 348, 80, 111, 122, 128, 124.
Psilomus ae ie ge alle

Ptichopus 6 De Bre lee SS:
ptox (Hriocremis, han) FE .. 103, 105, 107.
ptoxoides (Labienus) ae am so 105, AO
Ptychotrichus .. xa SOs

puberilis (Epilaches, 1 este a) pe 98; LOD, 103:
pubicostis (Proculejus) .. is Aal.
Publius = je ae 55 WD, Be, a, 22, bil,
pumilio (Aceraius, Gonatas, Omegarius) 1085110122:
punctatissimus (Passalus) .. sf: >> 8, 6%, GB,
punctato-striatus (Passalus) 52, 53, 68.
punctifrons (Mitrorhinus, Passalus) .. eae BS

punctiger (Mastochilus Pharochilus) .. OOo:
punctipectis (Leptaulax, Pentalobus) .. TON UA,
purulensis (Chondrocephalus, Popzlius) EE
pygmaeus (Aulacocyclus, Taeniocerus) oo LOT.
quadricollis (Passalus, Phoroneus) ws 20 58; Ol, 0%
quadricornis (Hriocnemis, Plesthenus). . so 46 OG, 9%
quadrifer (Ophrygonius) .. ae . 86.
-quaestionis (Hpisphenoides, Mastochilus s. str.) .. 99, 100, 103.
quinquecornutus (Chondrocephalus) .. 35 Uk, GRR aueh Ar,
quitensis (Passalus, Proculejus, Prosochtus) so bil, 39,55, GG,
recticlypeatus (Passalus, Petrejus) .. 0 5. OOF
recticornis (Passalus, Popilius) bie ye) 245 265 29:
rectidens (Aceraius) ae so OR
respectabilis (Pelopides, T lu Bor
Rhagonocerus .. ai 5e HO

Rhipsaspis ae: cys ES so Ge
Rhodocanthopinae toe vol:
rhodocanthopoides acetone rss 5369:
Rhodocanthopus oe

ridiculus (Oileus) 24, 25
rimator (Oileus) ne so 25

Rimor En ae ae 7223025:
Rimoricus Me: ; ia 22,025:
robustus (Passalus, Paxillus) La so 45, 5) Gi, CA,
roepstorfi (Leptaulax) cle er 550 IR)
rotundifrons (Macrolinus) .. se 56 | CU) teh,

rugosus (Passalus) Je ae a5 Be, 58, Oi,

142 Memoirs of the Indian Museum. : [ VoL.

Page.
sambawae (Leptaulax) .. HG ld te:
sansibaricus (Passalus, Pentalobus) .. an J EE
sargi (Oileus) .. om au: = 25:
sartori (Proculejus) le ul .. 24, 32.
Scalmus ma en a 2. Ol,
schellongi (Gonatas) a een 210921117122
schraderi (Pelopides, ? Protomocoelus) en 3110:
sculptilis (Vindex) Bic 2e .. AT.

Semicyclus = re is as)
Sertorius a a oe 2 83:
Severus + De ar ... Od:
sikkimensis (Bastlianus, Macrolinus) .. 22 80; 83.
simillimus (Vetwrius) Be a 230439:
simplex (Pelopides) A ve PROD:
singapurae (Ophrygonius) .. a po (hth Ol ws
sinuatocollis (Veturius) .. Ko .. 3d.
sinuatosulcatus (Veturius) Fe oo) OL BO, HO Oo
sinuatus (Passalus, Veturius) Ae bot alle, Ws
sinuatus (Veturius) Se of ede oe
sodalis (Aceraius, Mastochilus Cetejus) 21025103.
Solenocyclinae .. na Re co ul, 1 08 mete:
Solenocyclus .. ER de 008-4, 169/010Metc-
solidus (Pseudacanthus, Triaenurgus) . .. 24, 30.
solomonis (Protomocoelus) a 107.
Soranus er He Me 222,006:
Spasalus ah eh wv 48.
spinifer (Veturius) se = so dE BO, a,

| | spiniger (Passalus, Rhodocanthopoides) 2.059160, 67.

| spinipes (Passalus) À bo „28, BY; OL
spinosus (Passalus, Rhodocanthopus) .. 50 DS) Ole
Spurius ie Be w so PR PR. 20:
sternipunctus (Verres) ue ose .. 934, 40, 41.
Stephanocephalus AG 2 .. 10-11, 51.
stoliczkae (Comacupes) .. 6. 16 US.

striato-punctatus (Passalus, Popilius) 50.28, 20%

él studti (? Ericnomus, ? Malagasalus) .. 11.69.

i subrecticornis (Oileoides) .. ae 1.02, 249120.
sulciperfectus (Macrolinus). . ne 5a 87, 68, WAR
symmetricus (Parapzlopides. Pelopides) ao OE 8,
synelytris (Vindex) a a Ay, Al.

H Synesius a me Ye a ul

4 Taenioccrus ae ae Be 3, db, 16 ee, Webs

| Taeniocerus .. ae he LT.

Tarqununae.. 38 ie UE

1918.] F. H. GRAvVELY : Passalidae of the World. 143

Page.
Tarquinius a A 53 .. 3-4, 78, 80, 111, 122, 123, 124:
Tarquinius group oe ME M 210;
Tatius a or oe eo LOS:
tavoyanus (Aceraius) 4: a: oc | OY)
tenimbrensis (Gonatas) .. oe so 10 UCSD, Lalit,
teres (Aulacocyclus, Passalus) 5 1180192720:
Tetrarachus ae ae a ol
thoreyi (Trichostigmus) .. 24 NL:
Thryptocerus .. ue Me oo all:
Tiberioides = a a no U Udy SHE Cues, Ale
Tiberius = Le Le ag MOL;
timoriensis (Leptaulax, Passalus) ns SOI DIT ROX
toriferus (Passalus) 35 2695108:
Toxeutotaenius .. a vis 5) Oil,
Trapezochilus .. bes Fe 6 (tb OB,
Triaenurgus... a a so ey BO,
Trichopleurus .. a 2 0 ll,
trichostigmoides (Erionomus) a oo dt} 70) 78, 10
Trichostigmus .. = ER om AN, I, 122),
tricornis (Aceraius) un > oo OI, OB,
tricuspis (Aulacocyclus, Tristorthus) .. so lh. Teh TG, 110) 20, =
tridens (@naphalocnemis, Passalus, Pelopides) >9 O5, OG, Teil,
tridentatus (Gonatas) > de LOS:
trigonophorus (Labienus) .. Se .. 104, 106.
Tristorthus 58 or de Sella 17
tropicus (Passalus, Popilius) SA ae 245 285029:
truquii (Proculejus) oa ae ol 8%
Truquius sh Ne ae er
Undulifer AG oe je co. Cy 28, AD) Cue
unicornis (Passalus) a ws 26 68, CB
unicornis (Veturius) dfs ays oo «6, DO, BO)
ursulus (Zeptaulax, Trichostigmus) .. oo 2, 18
urus (Macrolinus) ae + om) CB, 3B
Valerius En Ir er >> Gil
Vatiniinae TS se Jae iol:
Vatinius 5 ue as SUR OR
Vellejus ae is a so HOS,
Verres be ie at .. 8, 82, 33, 84, 40 etc., 51.
Veturius 35 LE = .. 02, 33, 84, 35 etc., 51.
vicinus (Leptaulax) a > Aber ek
Vindex ae ae ae .. 10, 11, 48, 44, 46 etc., 51.
virginalis (Aceraius, Macrolinus Ana aches) .. 98.

Vikellinus ve 3: a so 20,

144 Memoirs of the Indian Museum. [Vor. VII, 1918.]

Page.
wallacei (Aceraius, Heterochilus, Ophrygonius) .. 76, 87, 89.
waterhousei (Macrolinus) .. a 00 (hl
weberi (Macrolinus) So 6c co BL

Zosterothrix SE Se 3 oa 51.

a
= it ER er

- DER - ’ é
a ae, ROMAINS CT 8 si ©
ETS
MES

EXPLANATION OF PLATE I.

Diagram illustrating the evolution and distribution of the Macrolininae.

The somewhat isolated genera Macrolinus and Pleurarius have been omitted. The
latter is probably confined to the Indian Peninsula,! and the former to other parts of the
Oriental Region and Celebes.

The species used in the preparation of the diagram are as follows :—

Aceraius grandis

» heljeri
Episphenus comptonr
indicus
a moore
neelgherriensis

Gonatas minimus
,, naviculator

,, schellonge
Labienus compergus
= dohrni
5 gigas
a trigonophorus
Mastochilus australiensis
ER nitidulus
» peltostictus

Ophrygonius inaequalis

Pelopides dorsalis .

cs symmetricus

ae tridens
Plesthenus invitus ..

= quadricornis

Protomocoelus australis
Pseudepisphenus perplexus
Tarquinius paradoxus
Tiberioides kuwerti

Aceraius dominant sp., E. Himalayas to Borneo.
5 other spp., E. Himalayas to Borneo.
Episphenus dominant sp., Ceylon.

I: dominant sp., Indian Peninsula.
> other spp., Ceylon.
5 other spp., Indian Peninsula.

Gonatas simplest sp., New Guinea, etc.
„ only sp. shown, Moluccas.
, dominant sp., New Guinea, etc.
Labienus other spp. (left), New Guinea, etc.
5 other spp. (right), New Guinea, etc.
ss spp. with fused elytra, Moluccas.
BE simplest sp., New Guinea, etc.
Mastochilus asymmetrical sp., Australia.

5 symmetrical spp., =
ee only sp. shown, Moluccas, New Guinea,
etc.

Ophrygonius only sp. shown, E. Himalayas to
Borneo.

Pelopides other spp., E. Himalayas to Borneo.

= simplest sp., E. Himalayas to Borneo.

is dominant spp., E. Himalayas to Borneo.
Plesthenus other sp., Celebes.

iR dominant sp., Celebes.

Protomocoelus only sp. shown, New Guinea, etc.

Pseudepisphenus only sp. shown, New Guinea, etc.

Tarquinius only sp. shown, New Guinea, etc.

Toberioides only sp. shown, E. Himalayas to
Borneo.

1 See above, p. 121.

Plate I.

Mem. Ind. Mus., Vol. VII, 1918.

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Vol. VII, No. 2.

m
SW NS onIan DEL

_ OBSERVATIONS ON THE SHELLS
OF THE

FAMILY DOLIIDAE.

E. W. VREDENBURG, B.L., B.Sc., F.G.S., etc.
Superintendent, Geological Survey of India.

_ Calcutta :

PUBLISHED BY THE DIRECTOR, ZOOLOGICAL SURVEY OF INDIA.
Re AT THE BAPTIST MISSION PRESS.

À ULY, 1910.

he Rupees Eight Aunas.

OBSERVATIONS ON THE SHELLS OF THE FAMILY DOLITD

By E. W. VREDENBURG, B.L., B.Sc., A.R.S.M., A.R.C.Sc., F.G.S.,
Superintendent, Geological Survey of India. (Communicated
with the kind permission of the Director, Geological
Survey of India.)

(With Plates II—VIII.)

I1.—Nore on Dozıum (EUDoLIuUm) FASCIATUM (BRUGUIERE), AND ON THE
SUB-GENUS ZUDOLIUM.

In Volume V of the Records of the Indian Museum (1910, p. 34), Mr. H. B. Preston
has figured a remarkable specimen of Dolium from Balasore Bay, characterised by the
presence, on the body-whorl, of a varix situated at an angular distance of about
35° from the thickened outer lip. The shell has been described as a new species
under the naine of Dolium varicosum. The collections of the Indian Museum include
four more specimens exhibiting a similar feature: two from Hong-Kong, one from
Vizagapatam, and another from the collections of the Asiatic Society, the exact
origin of which is unknown. As has been pointed out to me by Dr. Annandale, three
of these shells, including one of the Hong-Kong examples, are specimens of Dolium
fasciatum (Bruguiere), to which species evidently belongs also the Balasore specimen
to which Mr. Preston has already drawn attention. The Balasore specimen had
long been dead at the time when it was collected, for it is greatly corroded, over-
grown internally as well as externally by encrusting organisms (oysters, barnacles,
Serpula, and polyzoa), while no trace remains of the coloured bands which form so
characteristic a feature of Dolium fasciatum. Amongst the four varicose speci-
mens of Doliwm fasciatum preserved in the collection, the Balasore specimen is
remarkable as the smallest, measuring 57x40 mm.,' while the dimensions of the
Hong-Kong specimen are 100 x 74 mm., of the Vizagapatam specimen 86 x65 mm.,
and of the specimen of uncertain origin 82x60 mm. It should be kept in mind
that the three localities from which varicose specimens are known to have been
obtained, that is Balasore, Vizagapatam and Hong-Kong, have also yielded normal
specimens, and also that the latter often far exceed in size the varicose individuals.
For instance, one of the specimens from Balasore Bay, in which there is no super-

! The terminal growth of this stunted specimen is abnormal : the outer lip being posteriorly distorted so as to com-
municate an unusually narrow outline to the posterior part of the aperture and to the general ventral appearance of the
shell. This anomaly does not affect the penultimate varix which possesses the normal shape of the aperture of other
specimens, so that, viewed dorsally, the shell exhibits the usua] globose-ovoid outline.

146 Memoirs of the Indian Museum. [Vor. VII,

numerary varix, measures II5x89 mm. Nevertheless, as there is never more than one
pre-apertural varix, and as, when present, it is invariably situated close to the final
aperture from which it is separated by an angular distance of from 30° to 80°, it is
evident that the shells exhibiting this character are adult, or, at least, have completed
the growth of which they were capable. In certain species of Dolium, such as
D. fasciatum, D. zonatum, D. tessellatum, D. crosseanum, the internal, and in some
cases also the external thickening of the outer lip is invariably or almost invariably
present in every specimen, quite irrespective of size, and, as it is quite inadmissible
that all these specimens should have reached the termination of their growth, it is
clear that, as in the case of many other gastropods, the animal is able to absorb the
apertural structures at each successive phase of growth. The varicose specimens of
Dolium fasciatum at present under consideration represent, therefore, individuals
which, on approaching the final term of their growth, have lost the power of resorp-
tion; and, as the average size of these specimens is below that of the normally full-.
grown shell in which the pre-apertural varix is absent, they probably represent
individuals the vitality of which has been impaired through insufficient nutriment or
some other cause.' It seems evident that the majority of the specimens of Doliwm
Jasciatum reach their final stage of growth without leaving, on the body-whorl, any
trace of this pre-apertural varix.

Nevertheless, the presence of this structure, though not constant, is of consider-
able interest as the first instance hitherto recorded of a feature generally absent from
the Doliidæ, but characteristic of many Cassididæ ; especially if we take into con-
sideration the close relationship between Dolium fasciatum and the forms that have
been referred to the sub-genus Hudolium, the resemblance of which to the Cassididæ
has frequently been commented on, and is sometimes so pronounced that certain
forms of this group have on several occasions been erroneously referred to Cassi-
daria.

The Indian Museum collections contain one more example of a Doliwm exhibiting
a supernumerary varix (text-fig., p. 147). This is a handsome specimen of Doliwm
zonatum from Hong-Kong, measuring no less than 142 x 112 mm.” The supernumer-
ary varix is situated at an angular interval of about 55° from the outer lip whose edge
has not yet received its final thickening, the growth of the shell being, apparently, still
incomplete. In this species, the apertural thickening does not affect both the external
and internal borders of the aperture as in Doliwm fasciatum, but is developed only
internally. Consequently, the supernumerary varix, in the present instance, is not
conspicuous externally where it is indicated only by a slight swelling of the surface,
bordered, on its forward side, by a linear groove. Internally it is very strongly
developed and exhibits all the typical features that characterise the labrum of a full-
grown specimen.

! Compare Dr. Annandale’s remarks concerning the varix of Hydrobioides nassa from the Inlè Lake (Rec Ind. Mus.,
Vol. XIV, pp. 172-173), in which, however, the varix, as in certain other Gastropoda, is a constant adult character.

2 Tt has not been possible to prepare a photographic reproduction of this fine specimen which came to notice only
after the plates illustrating Dolium fasciatum had already been prepared.

1919.] E. W. VREDENBURG : Shells of the family Doliide. 147

In 1869 (Journ. Conch., Vol. XVII, p. 228, pl. xii, fig. 1) Monterosato described
a deep-sea form from the Mediterranean under the name of Dolium crosseanum refer-
ring it, in 1872 (Notizie intorno alle Conch. foss. di Monte Pellegrino e Ficarazzi,
Palermo, 1872, p. 89), to a new genus Doliopsis, a name unfortunately preoccupied
by Conrad in 1865 for a rather indistinct fossil, and for which, therefore, in 1889,
Dall substituted Hudohum (Bull. Mus. Comp. Zool., Vol. XVIII, p. 232). Tryon who
was at first sceptical about the habitat of the shell suggested (Man. Conch., Vol. VII,
1885, p. 263) that it might be a specimen of Doliwm zonatum accidentally mixed with
some Mediterranean shells. There is, however, not the slightest reason to doubt that
the shell was obtained from a depth which may have been as great as 50 fathoms, by
the Palermo fishermen who gave it to Monterosato. Nevertheless, Tryon’s suggestion
prompted by the general resemblance of the shell to Doliwm zonatum appears to be

Shell of Dolium zonatum showing a supernumerary varix (4 nat. size).

to a great extent in keeping with its zoological affinities. I am not aware of the
discovery of any further specimens in the Mediterranean, but the shell has been
obtained at a number of spots in the West-Indies, from depths ranging between 90
and 300 fathoms and more.

It was described and figured as Dolium bairdi (Trans. Conn. Acad., Vol. VI,
p- 253, pl. xxix, fig. 2) by Verrill who precisely compared it with Dolium zonatum ;
its specific identity with Dolium crosseanum being recognised by Dall in 1889 (loc.
cit.). The oligocene of Liguria, the miocene and pliocene of Piedmont, the pliocene of
the Alpes-Maritimes and of the Rhone valley, and of Tuscany, contain fossil forms
which are partly the obvious ancestors of Doliwm crosseanum.

According to Cossmann’s diagnosis in the Essais de Paléoconchologie (Fasc. V,
1903, p. 138), Hudohum is essentially distinguished from Dolium, s. str., owing to its
posteriorly slightly channelled aperture, its rugose columella, its shallow anterior

148 Memoirs of the Indian Museum. [Vor. VII,

notch and the absence of an umbilicus, the latter, however, being a direct result of
the feeble depth of the terminal notch, the accretions to which consequently do not
produce the strongly twisted bulge, which leaves room for the formation of a more or
less developed umbilicus amongst the species referred to Dolvum, s. str. Cossmann
also mentions, as a differential character, the tuberculation of the spire or body-
whorl, but as the genotype, Doliwm crosseanum, is without tubercles, this character
cannot be strictly taken into consideration.

If we ignore the depth of the terminal notch and the consequent presence or
absence of an umbilicus, we find, as regards the two other differentiating features,
namely the slight posterior channel and the rugose columella, that Doliwm fasciatum,
D. zonatum and, amongst specimens of Doliwm tesselatum, all those that are not
adult, agree with Hudoliwm rather than with Doliwm, s. str. Moreover, leaving aside
again the fully adult specimens of Doliwm tessellatum, these three, species are further
characterised by an internally thickened labrum with conspicuous denticulations,
usually bifid, which is not distinctly developed in any of the other species hitherto
referred to Dolium, s..str., but which is invariably present in those that have been
referred to Hudolium, as well as in the sub-genus or genus Malea.

If then we leave aside the characters furnished by the terminal notch and
umbilicus, Dolium fasciatum, D. zonatum and D. tessellatum will have to be regarded
as members of the group Zudolium, and, what the diagnosis loses in precision by the
exclusion of the umbilical characters, it gains to an extent at least equal by including
those of the outer lip. Now, the characters of the outer lip constitute precisely one
of the features in which Æudolium recalls the Cassididæ, and, by adopting the
grouping here proposed, we moreover include within Hudoliwm precisely both those
species which occasionally exhibit a pre-apertural varix such as is observed in many
Cassididee.

The tubercles observed on some fossil forms of Hudoliwm are derived from a
pronounced axial decoration analogous to the regularly distributed web of axial lines
that characterises Pirula. Just as the characters of the aperture have prompted the
erroneous reference of certain forms of Æudolium to the genus Cassidaria, so has this
reticulated or tuberculated ornamentation, especially when combined with the
elongate columella that distinguishes certain fossil forms, caused their erroneous
reference to Pirula. The earliest portion of the spire following the protoconch generally
shows a distinct web of regularly distributed lines of growth in most species of
Dolium: but this character is particularly well marked in the case, precisely of
Dolium fasciatum in which the axial lines are at first as thick as the average of the
spiral ornaments, with which they combine to form so characteristic a network that
the early part of the spire of Dolium fasciatum might be easily mistaken for that of
ey serous (lelle wth, ile, 246)).

According to Dall (Bull. Mus. Comp. Zool., 1889, Vol. XVIII, p. 223), the
radula of Doliwm crosseanum resembles that of the Cassidide more than that of
Dolium, s. str. Apparently the only species of Doliwm, s. str. of which the radula
has been figured is Doliwm perdix (Troschel, Gebiss d. Schneck., Vol. I, pl. xix,

1919.] E. W. VREDENBURG : Shells of the family Dohide. 149

fig. 3)." The radula of Pirula, judging from that of Pirula reticulata (Troschel, op.
cit., pl. xx, fig. 12), is essentially of the same type as that of Doliwm perdix.
Further observations on the radula of various species of Dolium would be of great
interest.

In a general way, Hudoliwm seems to be somewhat of a synthetic group in
which are united some of the characters of Dolium, s. str., of Malea, of Pirula and
of the Cassididæ. Nevertheless its relationship to Dolium, s. str. is of the closest
character and entitles it to rank perhaps only as a section rather than a subgenus.
It may here be mentioned that Dolium cinguliferwm (Bronn) |= Dolium fasciatum
(Borson) non Bruguiere | of all species the one nearest related to the genotype
D. crosseanum has been referred to Dolium, s. str. by Cossmann, who nevertheless
gives a figure (Æssais Paléoconch., V, pl. vi, fig. 10) in which there is no indication
of an umbilicus, while, on the contrary, Sacco has figured a specimen regarded as a
variety also of Dolium cinguliferum (Moll. terr. terz. Piem. e Lig., 1904, part XXX,
pl. xxu, fig. 5) which shows distinct traces of the tubercles that characterise other
forms of Æudolium. The sequel of these observations deals with the case of Dolium
tessellatum which, until an advanced stage of growth, exhibits the essential characters
of Hudoliwm, but finally, when fullv adult, becomes similar to Doliwm, s. str., and I
have suggested that this may afford some guide as to the derivation of the one group
from the other, as is further indicated by their geological history. If we adopt the
erouping here proposed, Doliwm, s. str. is not known from any strata older than
upper miocene, the only recorded fossil occurrences being in eastern countries, India
and Java, while the group, at the present day, is mostly eastern. Hudoliwm already
occurs abundantly in the oligocene, having been discovered, so far, in a fossil condi-
tion, only in Europe. Prrula is much more ancient and goes back to the cretaceous.

Il.--THE SPECIFIC DISTINCTNESS OF DoLIUM MACULATUM (LAM.) DESHAYES
AND Dozıum (EUDOLIUM) TESSELLATUM, BRUGUIÈRE.

INTRODUCTION.

Amongst the beautiful illustrations to Reeve’s monograph of the genus Doliwm
attention may particularly be drawn to four, of which the two first are referred to
Dolium fimbriatum, Sowerby (species 3), the next one to Dolium maculatum, Lamarck
(species 4), and another to Doliwm costatum, Deshayes (species 8).

In his Manual (Vol.. VII, page 264) Tryon rather emphatically asserts the
specific identity of these three forms with which he further unites as synonyms a
certain number of forms described as distinct by various authors. Tryon neverthe-
less maintains the three forms costatum, maculatwm and fimbriatum as three separate
varieties or races of a species costatum. No further discussion of Tryon’s conclusions

1 Troschel (op. cit , Vol. I, p. 227) has briefly described, without figuring them, the radule of two shells which he
has referred to ‘‘ Dolium maculatum, Lam. (= tesselatum, Encycl.) and D. costatum, Desh.,” both of which seem to
resemble closely that of Dolium perdix. The exact identity of the two shells in question is unfortunately, for the present,
uncertain,

150 Memoirs of the Indian Museum. [Vor. VII,

appears to have been attempted since the date of their publication, and not only
have the three names costatum, fimbriatum and maculatum continued in use as those
of distinct species, but the same is the case with some of the other names relegated
by Tryon to the synonymy.

Having had occasion to study a large series of fossil specimens of Doliwm from
the later Tertiary formations of the Mekran, it became necessary for me to pay
special attention to the related recent forms. From a perusal of the published
information regarding the recent forms above alluded to, it was not found possible to
decide definitely whether certain of the fossils under consideration were to be
regarded as identical with some of them or rather to be interpreted as separate
varieties or species. In order to establish a trustworthy comparison, it became
necessary therefore to undertake a fresh study of the recent forms of which a rich
series is preserved in the collections of the Indian Museum. The result of this study
has been to confirm some of the identifications established by Tryon, though his final
conclusions reach too far in one direction and not far enough in another ; for, on the one
hand, there is no difference (not even varietal) between Doliwm costatum and D. fim-
briatum, both of which represent a single species which should be known as Dolium
tessellatum, Bruguiere, while, on the other hand, Dolium maculatum represents a
totally distinct species.

The supposed three forms are therefore reduced to two, which, nevertheless,
under certain conditions are apt to exhibit a superficially deceptive resemblance to
one another. Before discussing their differences and their zoological affinities, it will
be useful therefore to give a detailed description of both these forms.

DESCRIPTIONS.
Dolium maculatum (Lam.) Deshayes.
(Pl. IV, figs. 1-3; pl. V, figs. 4-6.)

1685-1692. Buccinum sp., M. Lister, Historia Conchyliorum, pl. 899, fig. 19.
1757. ‘‘ Le Minjac,” Adanson, Histoire Naturelle du Sénégal, Coquillages, p. 109, pl. vii,
fig. 6.
? 1758. Buccinum dolium, Linneus, Systema Naturae, Ed. X, p. 735.
1770. Buceinum dolium, Linn. sec. Huddestord, Martini Lister, M.D., Historie sive
synopsis methodice Conchyliorum et tabularum anatomicarum editio altera.

pl. 899, fig. 19.
? 1822. Dolium maculatum, Lamarck, Hist. nat. des animaux sans vertèbres, Vol VII, sp. 3,
p- 260 -
1831-1837. Dolium maculatum, Lam., Kiener, /conographie des coquilles vivantes, pl. iii,
fig. 4.

1845. Dolium maculatum, Lam. sec. Deshayes, An. sans vert., 2nd ed., Vol. X, p. 140.

1849. Dolium maculatum, Lam., Reeve, Monograph of the genus Dolium, sp. 4.

1857. Dolium maculatum, Lam., Küster, Systematisches Conchilien-cabinet von Martini
und Chemnitz, Vol. III, 1st section, 2nd part, p. 73, pl. Ixii, fig. 3.

1885. Dolium costatum, Menke, var. maculata, Lam., sec. Trvon, Man. Conch., Vol. VIT,
p. 264.

19197 E. W. VREDENBURG : Shells of the family Doliideæ. 151

Medium to large, globose, with depressed slightly conoidal spire measuring from
one-seventh or even less to two-ninths of the total height.

The protoconch is relatively large, its visible portion attaining a diameter of
four millimetres. This visible portion is depressed and turbinoid and consists of a
very small, flattened, coiled nucleus and of three moderately convex whorls separated
by somewhat grooved sutures; this visible portion constituting only the apical
portion of the embryonic shell which, combined with the embedded portion, would
exhibit, in this, as in ali species of Doliwm, a globose or ovoid outline ; such embry-
onic shells having, on several occasions, been described as belonging to various
genera (see Fischer, Journ. Conch., Vol. XI, 1863, p. 147). The protoconch consists
of a very highly glazed, transparent, amber-coloured, horny substance. As is usually
the case with Doliwm, the protoconch is filled with a secondary deposit of porcellane-
ous shell-substance supplying an additional support which has ensured the durability
of this delicate structure. The protoconch is strongly oblique to the axis of the
remainder of the shell.

The linear junction of the protoconch with the succeeding portion of the shell is
straight and strongly oblique, antecurrent to the posterior suture and retrocurrent to
the anterior suture. In full-grown specimens the protoconch is followed by three
spire-whorls, the height of which does not exceed one-quarter of their width, the
maximum width coinciding with the anterior margin. They are separated by chan-
nelled sutures. The first half of the first whorl following the protoconch is evenly
convex, after which the whorls become angulated at about half their height. Poste-
riorly, a primary spiral rib borders the sutural channel, while another rather more
prominent spiral rib accompanies the angulation. A third principal rib is usually
visible, at least in the later portion of the spire, along the anterior margin of the
whorls, though, in some specimens, owing to an extreme flattening of the spire, it is
overlapped and concealed by the posterior edge of the next following whorl. There
are even specimens in which the sinking of the spire is so exaggerated that the
posterior edge of the body-whorl comes to coincide with the second primary rib.
Bach of the intervals between these main ribs carries several subsidiary spiral threads,
three of which are usually particularly conspicuous, representing a median intercalary
thread of the second order, flanked by two more threads of the third order ; their
respective thickness differing but slightly. There is, in addition, especially at the
earlier stages, a more or less complete set of threads of the fourth order, many of
which tend to disappear with increasing growth. Nevertheless, in many specimens,
several of these threads of the fourth order may be continued throughout the spire
and may reach the body-whorl together with the threads of the second and third
orders, which invariably persist. Minor inconsistencies are occasionally observed.
For instance, in the space anterior to the angulation, which is narrower than the
space between the angulation and the circumsutural rib, one of the threads of the third
order may be atrophied, so that this particular interval may carry only two conspicu-
ous subsidiary threads instead of three. A singular peculiarity is observed in a
specimen from Ceylon (or ? Kachh), in which the anterior thread of the third order in

152 Memoirs of the Indian Museum. [Vor. VII,

this particular interval is of about the same thickness as the true median thread of the
second order and is shifted quite close to it, so that they both together form a
conspicuous pair occupying approximately the middle of the said interval, separated
from one another at first by a minute thread of the fourth order. The two com-
ponents of the pair gradually thicken and finally coalesce into one broad flat band,
which remains somewhat bifid and is considerably wider than either of the two other
main ribs (Pl. V, fig. 5). Nevertheless, throughout the numerous specimens that have
been studied, the characters of the spire remain remarkably constant. The lines of
srowth are straight and strongly oblique, antecurrent to the posterior suture, retro-
current to the anterior suture. At the earliest stages of growth they form an ex-
tremely delicate web, intersecting the spiral ornaments, the crowded thin raised lines,
much thinner than the three first orders of spiral ornaments, being distributed with
the utmost regularity. With increasing growth, the lines become relatively less
prominent and much less regularly distributed.

The large body-whorl constitutes the greater part of the shell. It is globose,
almost spherical, exhibiting, on the right side of the shell, a continuous convex cur-
vature which, on the left side, is interrupted by the zone of accretions of the
deep terminal notch. Viewed dorsally, the zone of accretions is almost vertical
at its rather abrupt junction with the anterior flattened termination of the basal
convexity, and then assumes a convex outline becoming gradually more oblique in -
an anterior direction towards the right of the shell. The zone of accretions winds
very steeply and bulges very feebly, which partly accounts for the narrowness of the
umbilicus. Including the ornaments continued from the spire, and omitting the
narrow ridge which posteriorly limits the terminal zone of accretions, the body-
whorl carries ten or eleven primary spiral ribs. They are broad and ribbon-like
though slightly convex. The two most posterior ribs, that is the cireumsutural one
and the one continued from the angulation of the spire, are narrower than the
succeeding ones. The surface of the primary ribs frequently carries a variable
number of fine raised spiral striations. Throughout the greater part of the shell
the intervening spaces are much wider than the ribs, the two intervals continued
from the spire, especially the most posterior one of all, being generally broader than
the remainder. ‘Towards the anterior termination, the primary ribs become more
crowded and at the same time narrower, though, as the diminution in size does not
exactly keep pace with the contraction of spacing, the three or four last intervals are
of about the same width as the adjacent ribs or only slightly broader. The number
of primary ribs remains exactly the same at all stages of growth: it is the same
in small specimens of less than 30 millimetres in height as well as in full-grown
shells of over ten centimetres. In those shells in which the spire develops an addi-
tional large rib by the coalescence of two subsidiary threads, the full-grown shell
may apparently exhibit as many as twelve main ribs, but the supernumerary rib
betrays its adventitious origin by its relative flatness, as well as by the disposition
of the subsidiary threads in the two adjacent intervals in which they are fewer than
in the true primary intervals, and lastly by the absence of the characteristic macula-

1919. ] E. W. VREDENBURG : Shells of the family Doliide. 153

tions of the genuine primary ribs. The intervals between the primary ribs are
decorated with subsidiary spiral threads. In the case of very small specimens, all
the wider intervals may contain a complete representation of intercalary threads
belonging to the second, third and fourth orders, the surface thereby acquiring a
remarkably elegant appearance. Anteriorly, as the intervals become narrower, the
intercalations may become reduced to the threads of the second and third orders, and
finally to a single intercalary thread, while sometimes the most anterior of all the
primary intervals, owing to its narrowness, is without any intercalation. These inter-
calary threads persist with increasing growth except those of the fourth order which
generally fade away. The ornamentation remains quite unaltered throughout a
wide range of successive stages of growth, but becomes somewhat altered in the
case of large, fully adult specimens, in which some of the intercalary threads broaden
out into flat. bands resembling the primary ribs in shape, and filling a considerable
portion of the available interstitial space. Various inconsistencies are observed in the
development of these bands: in most instances they are due to the broadening of
the median or principal intercalary thread, that is the thread of the second order,
which then forms a band separated on each side from the neighbouring original
primary ribs by a thin thread representing the original threads of the third order.
At other times the broadening affects not only the median thread of the second order
but also one of the flanking threads of the third order, and then the original primary
interval may contain two adventitious broad flat bands which may become quite
equal and may become shifted in such a way as to occupy a practically symmetrical
position within the original primary space. Lastly, there are instances in which the
broadening only affects one of the original. threads of the third order, and the result-
ing adventitious band is situated quite unsymmetrically with respect to the original
space. Threads of the fourth order are frequently revived on these adult specimens,
but their reappearance is very inconsistent. They are apt rapidly to assume the
same thickness as some of the threads of the second and third order, with which they
may form close-set groups of two or three spiral threads. Owing to this reappear-
ance of the threads of the fourth order, the most posterior primary interval (generally
the broadest interval) carries variously disposed groups of spiral threads of various
sizes, disposed differently in different specimens, the interpretation of which can only
be deciphered by following them towards the apex along the spire. In a general
way, these adventitious ornaments of the full-grown shell are very variable, and no
two specimens are alike in this respect. They also commence to develop at various
sizes, but usually when the shell reaches or slightly exceeds a diameter of 60 milli-
metres. Nevertheless, in one specimen from Balasore Bay (M4408), measuring 120 x
96 mm., apparently the largest in the collection, the adult characters of the orna-
mentation have scarcely commenced to appear. The terminal zone of accretions is
posteriorly bordered by a narrow, feebly prominent, but sharp ridge, adjacent to
which the accretions to the actual notch form a rather broad band, almost flat in
some specimens, moderately convex in others, carrying crowded deeply concave lines
of growth together with a few distant obscure spiral markings. On the somewhat

154 Memoirs of the Indian Museum. Yo; IUT,

convex anterior zone intervening between this band and the umbilical portion of
the columellar margin, the curvature of the lines of growth assumes a reversed
direction, with the convexity turned anteriorly or forward, and there are two or
three flat spiral bands, sometimes bifid, of about the same width as the intervening
spaces. The lines of growth, throughout the body-whorl, are crowded, fine, incon-
spicuous, strongly oblique, anteriorly retrocurrent, straight until quite close to the
anterior zone of accretions towards which they bend backwards and which they
traverse with a strongly sigmoidal curve as above described.

The first half of the first whorl following the protoconch is of a uniform brown
colour, after which the pigmentation becomes differentiated in such a manner that
the primary ribs are white with chestnut patches at regular intervals, the intervening
spaces assuming a porcelain-blue to porcelain-purple colour, best seen in the case of
very fresh specimens. The contrasted whiteness of the non-maculated. portions of
the ribs is partly due to the opaque appearance caused by the thickening of the
shell substance, while the bluish appearance of the intervals is partly caused by their
thinner substance allowing the porcelain-like effect of translucency; nevertheless,
whenever the specimens are sufficiently fresh, it can be readily ascertained that the
colour effect is largely due to pigmentation of the intervals. The resulting appear-
ance is well rendered in Sowerby’s illustration in Reeve’s Monograph. The macula-
tions may be crowded as in the case of the specimen figured in Reeve’s Monograph,
or else much wider-spaced. They are particularly crowded in-some specimens from
the Andamans and from Balasore Bay, particularly wide apart in some specimens
from Puri, but the spacing varies greatly amongst specimens from one locality, and
even at different stages of growth in a single specimen. The maculations correspond
more or less exactly from one rib to another according to the direction of the incre-
ments of growth. As a rule there are no maculations on any of the intercalary
threads. In very eXceptional cases they may be present, on the body-whorl, on
some of the threads of the second order situated at about the widest part of the
shell anteriorly to the level of the suture. The epidermis, when preserved, has the
appearance of a thin layer of yellow varnish which does not interfere with the general
appearance of the colour scheme. The pigmentation of the spire is invariably more
pronounced than that of the body-whorl. The loss of vividness of the colour decora-
tion with increasing growth in all forms of Doliwm has already been commented on
by Reeve (Monograph, sp. IT).

The large semi-circular aperture, the more interior part of which is salmon-
coloured, becoming of a pure-white to bluish porcelain-like appearance towards the
edge, is quite simple posteriorly, while anteriorly it is terminated by a deep obliquely
disposed dorsal notch without any intervening canal. The columella forms an angle
of 125° to 130° with the base of the penultimate whorl. It is slightly oblique ante-
riorly towards the left of the shell. Its general direction throughout the greater part
of its length is straight with two slight bulges of which the more posterior one coin-
cides with the inward extension of the terminal zone of accretions, the more anterior
one with the sharply reflected edge of the columellar lip surrounding the narrow

29792] E. W. VREDENBURG : Shells of the family Dolude. 155

umbilicus. Anteriorly to the umbilicus, the terminal portion of the columella is
foliaceous and gradually contracts to a point at the anterior end of the shell, its
edge being steeply oblique anteriorly towards the left. The columellar lip is not
appreciable posteriorly and becomes distinct only anteriorly where it forms the thin
flat lamina reflected over the narrow umbilicus. The outer lip is straight and
strongly oblique. When fully developed it is bordered externally by a thin though
well-defined incised straight line situated at about six millimetres trom the actual
edge of the aperture. Between this line and the actual edge, the surface expands
somewhat outward, the spiral ornaments terminating in slight fimbriations. The
external limiting line approximately coincides internally with a slight swelling which,
in some specimens, is bifid. Owing to the thinness of the shell, the external spiral
ornaments are reproduced on the inner walls in reversed order of relief as is usual in
shells of this genus. On crossing the internal swelling of the fully developed outer
lip these spiral ornaments remain unaltered in character and do not give rise to
apertural tubercles as is frequently the case on the corresponding portion of other
species of Dolium. In the great majority of specimens, even those of the largest size,
the outer lip terminates in a perfectly simple edge.

Variability.—This remarkably abundant shell is particularly constant in all its
characters. The only variable features are the more or less sunken disposition of the
spire, inconsistencies in the intercalary spiral decoration especially on reaching the
adult stage, the variable degree of crowding of the maculations, and slight differences
in outline of the body-whorl which may tend to become a little more spherical or
spheroidal or else slightly ovoid. These variations are never correlated, but all occur
quite independently of one another, so that there is no distinct tendency towards the
formation of races or varieties. The most conspicuous abnormality in the orna-
mentation is that caused by the adventitious formation of a supernumerary rib as
described above in the case of a specimen from Ceylon (or? Kachh). It is the only
available specimen exhibiting this peculiarity which perhaps represents merely an
individual aberration.

Dimensions :—
neve itis. ot co IMA im 69 mm.
Thickness .. ns tari ONE: 55 »
Heisht of spire ae : 25 IO ,„
Height of body-whorl .. TO 64 »

The larger specimen is from Ceylon (or ? Kachh), the smaller one from Puri.

Occurrence.—This is the commonest species of Dolium along the Indian coasts.
It abounds wherever the sea-floor consists of fine soft sand or mud. According to
Melville and Abercrombie (Mem. and Proc. Manch. lit. and Phil. Soc., 4th ser.,
Vol. VII, 1803, p. 32) it is a deep-sea form. Nevertheless the shells are frequently
washed on to the beach.

Owing to uncertainties in the identification of this shell and of Doliwm tessella-
tum, there is some difficulty in ascertaining the limits of its distribution from pub-
lished accounts. Judging from the material preserved in the collections of the Indian

156 Memoirs of the Indian Museum. [Vor. VII,

Museum it is found all along the coasts of peninsular India from Kachh on the western
side to Balasore in the east, and also along the coast of Arakan and the Malay
Peninsula. One specimen is labelled as coming from as far east as Amboina.

In a fossil condition, it is known from the pliocene of the Mekran coast and
from the post-tertiary formations of the Pulicat lake.

The discussion of the relationship of this shell to other species will be deferred
until the completion of the description of Doliwm tessellatum.

Dolium (Eudolium) tessellatum, Bruguiere.
(Plate VI, fig. 7; pl. VII, figs. 8-10; pl. VIII, figs. 11-13.

1789. Buccinum tessellatum, Bruguière, Encyclopédie méthodique, Vol. VE, sp. 4, pp. 236,
246, pl. 403, figs. 3a, b.
1790. Dolium tessellatum, Bruguiere, Encyclopédie méthodique, pl. 403, figs. 3a, b,
1823. Dolium fimbriatum, Sowerby, Genera of Shells, fig. 2.
1830. Dolium costatum, Menke, Synopsis methodica, 2nd ed., p. 63.
1831-1837. Dolium fasciatum, Brug. var. sec. Kiener, Iconographie des Coq. viv. Dolium, pl. iv,
fig. 6.
1831-1837. Dolium variegatum, Lam. (junior) sec. Kiener, Icon. des Cog. viv., pl. ii, fig. 3.
1845. Dolium costatum, Deshayes, An. sans vert., 2nd ed., Vol. X, p. 144.
1845. Dolium minjac, Adanson sec. Deshayes, An. sans vert., 2nd ed., Vol. X, p. 145,

n029.
1845. Dolium ampullaceum, Philippi, Zeit. Mal., p. 147.
1849. Dolium fimbriatum. Sow., Reeve, Monograph of the genus Dolium, sp. 3 ‘

1849. Dolium costatum, Desh., Reeve, Monograph of the genus Dolium, sp. 8.

1849. Dolium ampullaceum, Phil., Abbild. III, 4. Dolium, p. 12, pl. ii.

1857. Dolium costatum, Mke., Küster, Conch. Cab. von Martini und Chemnitz, Vol. III,
Ist section, 2nd part, p. 61, pl. lvi, fig. 3; pl. lvii, fig. 3.

1857. Dolium Lischkeanum, Küster, Conch. Cab., p. 71, pl. Ixii, fig 1.

1857. Dolium fimbriatum, Sow., Küster, Conch. Cab., p. 72, pl Ixü, fig. 2.

1879. Dolium costatum, Desh., Martin, Die Tertiärschichten auf Java, p. 40, pl. vil,
figs. 9, 10.

1899. Dolium costatum, Desh., Martin, Samml. des geol. Reichsmus. in Leiden, new series,
Mol p: 161 ply vos pris Te;

Medium to large, globose, slightly ovoid, with slightly conoidal, sometimes
conical depressed spire measuring from two-ninths to three-tenths of the total
height.

The protoconch is relatively small, the diameter of the visible portion not ex-
ceeding three millimetres. It consists of a horny transparent substance of dark-
brown colour. The internal secondary infilling of porcellaneous shell-substance does
not reach the apex, so that the minute, depressed, coiled nucleus, lacking internal
support, is almost always broken off. The visible portion of the protoconch is rather
prominent, semi-naticoid, that is with the appearance of a half-embedded Natica and
includes three convex whorls separated by very narrow, slightly grooved sutures.
The protoconch is slightly oblique to the axis of the remainder of the shell. The
line of junction of the protoconch with the remainder of the shell is slightly curvi-

1919.] E. W. VREDENBURG : Shells of the family Doliide. 157

linear with forward facing convexity, and is oblique, antecurrent to the posterior
suture, retrocurrent to the anterior suture.

The protoconch is followed by three to three and a half convex spire-whorls,
separated by slightly sunken sutures. Their height varies from two-ninths to a
little over one-quarter of their width, the maximum thickness coinciding with the
anterior margin. The whorls usually exhibit four ribbon-like main ribs, considerably
narrower than the intervening spaces which are slightly concave. The most posterior
main rib, which is narrower than the others, encircles the circumsutural depression.
The most anterior rib coincides with the anterior margin and is frequently more or
less overlapped by the posterior edge of the next following whorl. In rare instances,
the spire is so much sunken that the posterior edge of each whorl reaches the level
of the third primary rib of the preceding whorl. The intervals between the primary
ribs may be approximately equal, though usually the most posterior interval is
somewhat wider than the remainder. On the first whorl following the protoconch,
each interval usually carries three delicate subsidiary spiral threads, namely a median
thread of the second order flanked by two threads of the third order. Occasionally
one of the threads of the third order may be atrophied or indistinct. On the second
whorl, the threads of the third order disappear, leaving only, in each interval, the
median thread of the second order which also becomes gradually thinner and indis-
tinct with increasing growth. On the third whorl all the threads of the second order
may likewise disappear, leaving the intervals perfectly smooth, but there usually
subsists a more or less distinct remnant of the line intersecting the most posterior
interval, usually reaching even to the body-whorl. A number of extremely fine
spiral lines are usually observed on the surface of the main ribs. The extremely fine
crowded lines of growth, especially distinct and regular on the first whorl following
the protoconch, are practically straight and strongly oblique, antecurrent to the
posterior suture, retrocurrent to the anterior suture.

The large body-whorl constituting the greater part of the shell is always
strongly inflated and globose, and may be almost spherical, but is more usually
distinctly ovoid. On the right side of the shell its convexity is continued as far as
the anterior termination, while on the left side a shallow concavity intervenes
between the main basal convexity and the zone of accretions to the very deep
dorsal notch, whose outline, on the left side of the shell, viewed dorsally, is steeply
oblique anteriorly towards the right and slightly convex. The actual edges of the
notch are slightly reflected outward. Ventrally, the steeply winding anterior edge
of the terminal zone of accretions is bounded by the foliaceous termination of the
columella and columellar margin, with the formation of a narrow umbilicus. Includ-
ing the spiral ornaments continued from the spire, and irrespective of the ridge
forming the posterior edge of the terminal zone, the body-whorl, in the case of small
specimens measuring less than 35 mm. in height, carries twelve primary ribs. In
specimens measuring from 35 to 45 mm., the number of primary ribs is usually
thirteen. In the majority of specimens ranging from 45 to 100 mm., the number of
primary ribs is fourteen, though, occasionally, amounting only to thirteen. At still

158 Memoirs of the Indian Museum. [Vor. VIT,

larger dimensions, the primary ribs may increase to as many as sixteen. The
increase is due to the appearance, one at a time, of an additional rib at the anterior
limit of the convexity of the base, just along the edge of the terminal zone of accre-
tions. The ribs are ribbon-like, and become more crowded and somewhat narrower
towards the anterior limit of the base, in consequence of which the intervals, which
throughout the greater part of the body-whorl are broader than the ribs, become,
towards the anterior extremity, of about the same width as the ribs or narrower.
As in the case of the spire, the ribs carry delicate spiral lines. In the great majority
of specimens the intervals are quite without any spiral ornaments, except usually
the most posterior and broadest interval, which is generally bisected by a thin
remnant of the median subsidiary thread continued from the spire, though there
are specimens in which even this last remnant has disappeared. In the case of
very young specimens of less than 30 mm. in height, every interval throughout
the body-whorl is bisected by a very thin intercalary thread.’ With the usual
exception, as already noticed, of a feeble remnant in the most posterior interval,
all these subsidiary threads disappear before the shell has reached a height of
30 mm. Amongst the series of specimens in the Calcutta collection, the sculpture
of the body-whorl remains perfectly unaltered up to a total height of 100 mm.
It is only in quite adult specimens of still larger size that the sculpture enters
upon a new phase through the re-appearance of intercalary ribs, which may broaden
until they fill almost the whole of the available interstitial space, and assume
an appearance almost identical with that of the original primary ribs. On the later
part of the body-whorl of the largest specimen in the Calcutta collection, the seven
first intervals, counting from the posterior edge of the body-whorl, each carry an
intercalary rib. The eighth, ninth, and tenth are plain, but an intercalary rib also
appears in the eleventh. In the large individual illustrated in Reeve’s monograph
(fig. 3a), some of the intervals carry two subsidiary ribs. The scarcely bulging,
torose, steeply winding zone of accretions is posteriorly bordered by a very thin,
narrow, sharply defined ridge. It is divided into two sub-equal portions, a posterior
one corresponding with the accretions of the deep indentation of the notch, across
which the lines of growth are deeply concave, and an anterior portion corresponding
with the accretions of the anterior border of the notch and anterior termination of
the shell, across which the lines of growth are convex. This anterior portion carries
three or four spiral ribs. The band corresponding with the accretions to the notch
may also carry spiral ribs, especially in the case of small specimens, but they are less
prominent than those of the anterior sub-zone, and, in many instances, are represen-
ted merely by some obscure distant spiral lines. The anterior termination of the
anterior sub-zone, forming the anterior termination of the shell, when well preserved,
is foliaceous and somewhat palmately expanded. The thin lines of growth, over the
greater part of the body-whorl, are strongly oblique and anteriorly antecurrent, and

! Martin has figured a fossil specimen from the pliocene beds of Java (Samml. des geol. Reichs-Museum in Leiden,
new series, Vol. I, pl. xxv, fig. 372), referred to Dolium costatum, and measuring about 30 mm. in height, in which the
intercalary median thread is faintly visible in several of the intervals of the body-whorl.

1919. | E. W. VREDENBURG : Shells of the family Doliide. 159

are practically straight throughout the greater part of their course, bending backward
anteriorly only in the immediate neighbourhood of the terminal zone of accretions
which they traverse, as above described, with a pronounced sigmoidal flexure.

The large semi-circular to semi-oval aperture is anteriorly terminated by the
deep, strongly oblique dorsal notch, without any intervening canal. Its posterior
termination is slightly though distinctly channelled in the case of immature speci-
mens, but becomes simple when the shell is quite adult. The columella which is, on an
average, approximately straight, and slightly oblique anteriorly towards the left of
the shell, forms an angle of about 120° with the base of the penultimate whorl which
it joins rather abruptly. It exhibits two rather feeble winding bulges of which the
more posterior one coincides with the mward extension of the zone of accretions,
while the more anterior one represents the junction with the reflexed edge of the
columellar lip. In the case of small and medium specimens, the anterior part of the
columella carries a number of rugosities some of which are sometimes internally con-
tinued as thin spiral folds. When the specimens attain a height of about 90 mm.,
the columellar rugosities become indistinct, while, in the case of fully adult specimens,
they entirely disappear, and the columella is quite smooth. The anterior thin foliace-
ous terminal portion of the columella, anteriorly to the very small umbilicus, is very
narrow. The columellar lip spreads rather widely over the ventral surface of the
base. It is mostly so thin as not to interfere with the sculptured and coloured
decoration of the base of the penultimate whorl. Nevertheless, in the case of
immature specimens its marginal portion, of a glossy porcellaneous texture, is
frequently sufficiently thickened to become opaque white, the actual edge adhering
fairly closely to the more posterior portion of the convexity of the base, but becom-
ing semi-detached or even detached on approaching the terminal zone of accretions.
Anteriorly to the terminal zone of accretions it surrounds the small umbilicus and
joins the columella. The thickened edge gradually becomes less distinct as the
specimens grow larger. In the case of full-grown specimens the greater part of the
columellar lip ceases to be appreciable, the anterior termination, where it surrounds
the umbilicus, alone remaining distinct. In a few specimens, at the posterior
termination of the columellar lip, there is a feebly prominent though distinct oblique
ridge which contributes to define the shallow posterior channel. The outer lip is
straight and strongly oblique. When its structure is characteristically developed,
it is externally thickened and expanded, its edge is deeply fimbriated owing to the
intervals between the ribs extending much further forward than the ribs themselves,
while internally it is thickened and denticulate. All these characters, external expan-
sion, marginal fimbriations, internal thickening and denticulations, are as a general
rule most typically developed in small and especially in small-medium specimens,
especially those measuring from 55 to 75 mm. in height. In exceptional instances
they are still quite typically developed in specimens measuring as much as 80 mm.
in height, but as the shell exceeds these dimensions they become more and more in-
distinct. When the shell is quite adult, the external thickening and expansion, the
marginal fimbriations, the posterior channel, and the internal denticulations entirely

160 Memoirs of the Indian Museum. [Vor. VII,

disappear, and there only remains a slight internal thickening close to the edge. As
is usual in the shells of this genus, the external ornamentation is reproduced in the
interior of the shell in inverted relief, so that it is the sunken intervals or grooves
of the outer surface which assume the appearance of ribs on the internal walls. On
crossing the internal marginal swelling of the labrum, in those specimens in which
the apertural features are characteristically developed, each of these internal ribs
usually develops a pair of elongate denticulations or ridges which terminate externally
against the fimbriated digitations of the edge. Sometimes the most anterior internal
rib develops but a single denticulation or ridge, while frequently some of the broader
posterior internal ribs (corresponding to the broad posterior intervals of the outer
surface) may give rise to three denticulations, or, occasionally, to as many as four.

The first whorl following the protoconch is dark brown, though not so dark as
the protoconch itself. It is not uniformly tinted, the depth of the colour decreasing
considerably towards the posterior margin. On the following whorl the broad spaces
between the primary ribs are usually of a reddish or purplish, or sometimes bluish
tinge, while the main ribs themselves are white or yellowish-white maculated with
yellowish-brown. The same scheme may be continued on the third whorl with a
paler tint for the intervals, and may also extend over the entire body-whorl, the
intervals becoming still paler, until with increasing growth they may become quite
white. In certain cases, the ribs, over the whole body-whorl, may maintain the
appearance of white bands with yellow spots (as in the type of Doliwm lischkeanum,
Küster), while in other instances their coloured decoration may gradually disappear,
and, when the colour of the intervals has likewise vanished, the entire bod y-whorl
may be white. In other instances, the yellow tinge of the maculations gradually
spreads along the intervening portions of the ribs until all the spots coalesce, and the
ribs assume the appearance of continuous yellow bands. When, as is frequently the
case, the intervals have become nearly or quite colourless, the general colour scheme
becomes reversed in this sense that the ribs, instead of appearing as light spotted
bands against a darker ground, become darker bands against a lighter ground. This
is the appearance exhibited by Kiener’s Doliwm fasciatum var. Lastly, there are
specimens in which the greater part of the shell is almost uniformly tinted of a rich
orange or burnt-sienna colour, deepening to brown towards the apex. In large
thoroughly adult specimens, the intercalary ribs assume the same maculated decora-
tion as the original primary ribs. The deeper internal portion of the aperture is of
a yellowish or brownish tinge. The columella, the thickened portions of the colu-
mellar lip and inner portions of the outer lip are white and porcellaneous.

Dimensions.—The following measurements refer to a series of specimens obtained
from the Andamans, with the possible exception of No. 7, the exact origin of which
is uncertain. No. 5 is from the South Andaman.

(1) (2) (3) (4) (5) (6) (7)

Height .. 122 mM, 195 mm, 7/2 Or mm Sem mem ne
Thickness 00) 255 EEE lan SO BO on DO. 20
Height of spire 20 oe DEE DDR, 10%, LS, TO, A So

1919. | E. W. VREDENBURG : Shells of the family Doliide. 161

Height of body- (7) (2) (3) (4) (5) (6) (7)
whorl .. 105 mm. 84mm. 64mm. 56mm. 49mm. 33 mm. 32 mm.

The next specimen, of uncertain origin, is remarkable for its exceptionally
sunken spire :

(EAN es er de ate 70 mm.
Thickness En Een ve BO
Height of spire is aie eee HORS
Heisht of body-whorl oe i 65,

Variability and Development.—This shell is even more constant in its characters
than the previously described Doliwm maculatum, the only distinctly variable features
being those of the colour scheme. As regards the more essential characters, one
merely observes slight differences in the more or less sunken disposition of the spire,
and the more or less distinctly spherical or ovoid outline of the body-whorl.

At the same time, some very remarkable differences are observed in the appear-
ance of the shell at different stages of growth, especially with reference to the aper-
tural characters. While, in the case of fully adult specimens, the aperture is pos-
teriorly simple, the outer lip undifferentiated, and the columella smooth, the shell at
early and intermediate stages of growth has a strongly fimbriate and denticulate
outer lip, a posteriorly slightly channelled aperture, and a distinctly rugose columella.
Whilst the adult shell is in every sense a typical Doliwm, the posterior channel and
especially the rugose columella of the immature specimens recall the sub-genus
Eudolium. The external and internal thickenings, marginal fimbriations, and inter-
nal denticulations of the aperture, the posterior apertural channel, the thickening of
the columellar lip and the rugosities of the columella observed in these immature
specimens of Dolium tessellatum agree in every respect with the corresponding fea-
tures observed at all stages of growth in Dolium fasciatum (Brug.) which there is
reason to refer to the subgenus Zudolium, and readily account for Kiener’s interpreta-
tion of such specimens of Dolium tessellatum as representing a variety of Doliwm
fasciatum. The development of Dolium tessellatum is of great interest as suggesting
the possible derivation of the more typical forms of Doliwm from Hudolium, especial-
ly as, so far as can be judged from the information at present available, Eudolium is
geologically more ancient.

Occurrence.—With the exception of a few individuals of uncertain origin, all the
specimens in the collection of the Indian Museum are from the Andamans and
Nicobars. Shells certainly referable to this species have been obtained from the
Malay Islands and Japan.

In a fossil condition, the species is known from the upper miocene and pliocene
of Java, and from the pliocene of the Mekran.

COMPARISON OF DOLIUM MACULATUM AND DOLIUM TESSELLATUM.
Remarks on the Taxonomy.—Owing to the hopelessly confused synonymy of the
forms under consideration, the task of discovering suitable names for the two species
above described has proved extraordinarily troublesome.

162 Memoirs of the Indian Museum. VO ARE

Many of the works containing the more or less recognisable figures cited by the
earlier authors are, unfortunately, not available in India. Nevertheless there is not
the slightest doubt that they include representatives of both the species above
described in detail, while the diagnoses are generally far too concise.to elucidate the
confusion which undoubtedly exists in the synonymy.

Amongst the appellations for which any definite claim can be put forward to be
used as specific names, the oldest is probably Adanson’s “le Minjac” (1757). De-
shayes regarded the figure as representing the same species as Sowerby’s Dolium fim-
briatum, and it would correspond therefore with Doliwm tessellatum as above described.
Deshayes has therefore adopted for this species the name Doliwm minjac, Adanson.
Adanson, in his synonymy, refers to Lister’s figure which represents Doliwm macu-
latum, and to Rumphius’ which represents D. tessellatum. The description is not
sufficiently detailed for establishing which species is meant. The body-whorl is said
to carry fourteen ribs, which, in specimens of the size of the one referred to by Adan-
son, namely two inches (about 50 mm.), might suit D. tessellatum ; only, as the ribs,
over the widest part of the shell, on the figure illustrating the dorsal aspect, are shown
of two alternating sizes, of which only the thicker ones carry spots, it seems obvious
that the figure represents D. maculatum, and not D. tessellatum as conjectured by
Deshayes. As it is, Deshayes seems somewhat uncertain as to whether Adanson’s
figure represents Sowerby’s Dolium fimbriatum or Lamarck’s Dolium variegatum (An.
sans vert., 2nd ed., p. 143, footnote). The locality, Senegal, is improbable, more so
even for Dolium tessellatum. (Brug.) than for Dolium maculatum, Deshayes, since, in
the Indian Ocean at any rate, the latter extends much further west.

The authority next in date is Linneus, in the roth edition of the Systema
Naturae, published in 1758, in which d’Argenville’s genus Dolium is merged into
Buccinum, and a species recorded as Buccinum dolium. The roth edition is not
available in India. The synonymy as given by Gmelin in the 13th edition includes
the following citations :—

Rondelet, Testacea, 106, Cochlea rugosa.

Rumphius, Mus., pl. 27, fig. A, Cochlea striata s. olearia.

Calceol. Mus., 30, pl. 41.

Lister, Conch., pl. 899, fig. ro.

Bonann. Recr., 3, figs. 16, 17, 25.

Miss Kercher Mes TO, 28:

Gualtieri, Test., pl. 30, fig. E.

D’Argenville, Conch., pl. xvii, fig. C.

Seba mus. 3, pl. 68, figs. 9, 11; pl. 70, figs. I, 2, 5.

Mus. Gottwald. pl. 27, fig. 1855, fig. 188b.

Knorr) Veron, 3, ployıı to

Martini, Conch., pt. 3, pl. 116-118, figs. 1072-1075, 1082.

Out of the twelve works mentioned in this list, only five are accessible in Cal-
cutta, namely those of Rumphius, Lister, Seba, Knorr, and Martini.

The shell figured by Rumphius clearly corresponds with that here described

1919.] E. W. VREDENBURG : Shells of the family Doliide. 163

as D. tessellatum. The library of the Geological Survey contains the second edition
of Lister’s Conchology, from which it may be recognised that the figure mentioned
in the synonymy represents a shell of Doliwm maculatum, the drawing of which has
been badly interpreted by the engraver; so much so that, for anyone who had not
specially studied the species, the figure might just as well be taken for one of Doliwm
tessellatum, while the short description is too ambiguous to be of any help. In
Seba’s work, figures 9 to II, plate Ixviii, represent Dolium maculatum, while figures
I and 5, plate Ixx, represent D. tessellatum. Figure 2, plate Ixx, is too in-
distinct for identification and possibly represents a shell of another genus. Knorr’s
figure undoubtedly represents D. tessellatum. In Martini’s work, figures 1072 and
1082 undoubtedly represent D. tessellatum, while figures 1073 and 1074 undoubtedly
represent D. maculatum. The shell represented in figure 1075 is immature and
cannot be so securely identified, though it is probably referable to D. maculatum.

Amongst later authors, the one who has most fully dealt with the iconography
of the works under consideration is Küster by whom the figures published by Rum-
phius and by Knorr are referred to Dolium fimbriatum, that is to the species above
described as Dolium tessellatum ; while the figures in the works of Gualtieri and d’Ar-
genville are regarded by the same authority as representing Doliwm maculatum.

In conclusion, it is abundantly clear that the figures referred to in Gmelin’s
synonymy are about equally divided between the two species above described as D.
maculatum and D. tessellatum. The synonymy quoted by Linnzeus and Gmelin does
not help us therefore in determining which shell was intended in the Systema Nature.
The diagnosis, unfortunately, does not help us any further. It is as follows: “ testa
ovata cincta sulcis remotis : cauda prominula,” and is therefore totally insufficient to
recognise which species is meant.

In 1770, William Huddesford, in the second edition of Lister’s Conchology, adop-
ted the name Buccinum dolium for the figure referred to in the Systema. There
should be here no room for any ambiguity. Unfortunately, as already mentioned,
both the engraving and the diagnosis are very unsatisfactory and would not of
themselves materially afford any help to the identification of the species.

Bruguière, in 1789, in describing the “ buccin cordelé,” gives to a large extent
the same synonymy as Linneus for Buccinum dolium. There is nevertheless an im-
portant divergence in the reference to Martini’s work, from which Linnæus has
quoted no less than five figures of which two only are selected by Bruguiére as
representing his Buccinum tessellatum. These are figures 1073 and 1082, the second of
which, forty years later, was selected by Menke as the type of Doliwm costatum.

Bruguiére’s descriptions are of an entirely different type from Linneeus’ diagno-
ses, and, although the plates illustrating the Hncyclopédie Méthodique ' are not avail-

1 This is not the famous, though frequently inaccurate original edition of the ‘‘ Encyclopédie,” but the magnifi-
cently planned 2nd edition, initiated in 1782 under the most distinguished auspices, the several parts being entrusted
to the most accomplished and learned men of the age. The publication of this stupendous work had been so organised
as to ensure its completion in 1792. The revolution unfortunately robbed the scheme of the greater portion of its
enlighted and distinguished patronage, after which the publication lingered in a precarious and impoverished condition
until 1832, when it was finally interrupted and remained unfinished.

164 Memoirs of the Indian Museum. [Vor. VII,

able in Calcutta, the description of his Buccinum tessellatum is so complete and
precise as to render the illustrations superfluous.

Indeed, but for the praiseworthy exception of a small number of naturalists,
Bruguiere, in the admirable accuracy and fulness of his descriptions, is more than a
century in advance of his successors.

As the present enquiry has not for its object to resuscitate an obscure obsolete
name in place of others in general use and of well-established meaning, but as it is
merely an attempt to select a suitable name amongst half a dozen or more the
present usage of which is involved in the utmost confusion, it will perhaps not
appear superfluous if Bruguiére’s description is here reproduced in eatenso before
analysing it. The sentences are numbered for the convenience of reference.

(x) “ Celui-ci ne cède pas en beauté au Buccin perdrix, mais il lui est très-souvent
inférieur par son volume; celui dont je donne la description est étonnant par sa
grandeur et par sa belle conservation, il surpasse de plus d’un tiers leur proportion la
plus ordinaire ; il a quatre pouces six lignes de diamètre, la longueur de son ouverture
est de trois pouces sept lignes, et sa largeur d’un pouce dix.” (2) “Sa spire est com-
posée de sept tours complets, qui sont garnis de côtes élevées, convexes, écartées,
au nombre de quatorze sur le tour inférieur, et de quatre seulement sur ceux du
haut; ces côtes sont séparées par des sillons plats, ordinairement plus larges qu’elles,
qui sont quelquefois marqués au milieu par une ligne élevée qui suit leur direction.”
(3) “Cette coquille, ainsi que la précédente” (Buccinum perdrix) “ ne forme point
de canal enfoncé à la jonction des tours, ils appuient au contraire carrément l’un
sur l’autre, et laissent à leur jonction un rebord applati.” (4) “ Son ouverture est
très grande et cannelée dans l’intérieur.” (5) “ La lèvre droite est peu évasée
et dentée pendant la jeunesse, de manière que chaque côte est terminée par deux
lignes élevées qui disparaissent tout-à-fait quand la coquille est parvenue au volume
de celle dont je donne la description.” (6) “ La lèvre gauche ressemble a celle des
espèces précédentes par son peu d'épaisseur, qui est telle que, quoiqu’elle recouvre
les côtes du ventre de la coquille, elles n’en sont pas moins saillantes pour cela.”
(7) “La columelle est formée comme dans le Buccin cannelé, elle est tordue en
spirale et garnie à l’extérieur des côtes longitudinales jusqu’à l’échancrure de la
base ; ’ombilie est situé comme dans cette coquille, mais il a un peu moins de lar-
geur et moins de profondeur.” (8) “ Ce Buccin est ordinairement blanc à l'extérieur
ou de couleur fauve, et ses côtes sont le plus souvent marquées de grandes taches
fauves, jaunes ou orangées, qui sont presque toujours effacées sur les quatre ou cinq
tours plus anciens.” (9) “ Mais ces taches manquent quelquefois tout-à-fait, et la
coquille est alors blanchâtre ou d’une teinte faible de couleur de chair ; on en connaît
aussi des variétés qui sont toute brunes, d’autres dont les côtes sont un peu élevées,
plus écartées et presque aigues, dont la couleur tire sur le gris ou le cendré ; il ne doit
pas paraître étonnant que cette coquille offre des variétés si remarquables, puisque
son espèce occupe une étendue immense sur la surface de la terre.” (10) “ Linné a
dit qu’on la trouvait sur les côtes de la Sicile et de la Barbarie ; Bonanni dit de
même, inais il la reçut aussi des Indes orientales ; M. Adansson la trouva au Sénégal ;

1919. ] E. W. VREDENBURG : Shells of the family Doliide. 165

Rumphius a Vile d’Amboine; Martini l’indique aux îles de Tranquebar ; et Petiver
a l’île de Luçon, l’une des Philippines.”

We may now add a few comments on this admirable description.

(I) From the very first sentence, Bruguiére, in accordance with the best models
of modern descriptions, adopts the practice, too often neglected even by modern
writers, of giving detailed measurements of the species. Judging from the figures
quoted, the average diameter of large specimens should be three inches, or about
75 mm., which agrees with what is generally observed in the species above described
as Dolium tessellatum, while the exceptionally fine specimen specially selected by
Bruguiere for description measured four and a half inches, or about 115 mm. in
diameter, a size probably never attained, so far as is known, by Dolium maculatum,
and only inferior by about 20 mm. to that of the magnificent specimen figured in
Reeve’s monograph as Dolium fimbriatum. The most essential point with reference
to the present enquiry is that the detailed measurements given by Bruguiére are
those of the particular specimen about to be described. Clearly then, this description
is not a generalised diagnosis built up from a number of specimens or uniting the
separately published features of previous descriptions. It is a detailed description
of a single specimen, and obviously therefore cannot refer to more than one species.
Whatever may be that species, it is clear that the merest glance through the very
first sentence suffices to dispose of the totally undeserved criticism of Reeve, of
Kiister, and of Tryon, regarding the alleged composite character of Doliwm tessellatum.

(2) The dimensions recorded in the first sentence already exclude almost all
possibility of referring Bruguiére’s type to Dolium maculatum. The number of ribs
as recorded in the second sentence irrevocably confirms its attribution to the shell
described in the present work as Dolium tessellatum ; for even taking into account
any possibility, however improbable, of mistaking, on the body-whorl, some of the
ribs of the second order for primary ribs, the number recorded on the spire-whorls,
namely four, settles once for all the specific attribution of the shell under considera-
tion. Dolium tessellatum, as understood in the present work, does not always exhibit
as many as four ribs on the spire-whorls, but Doliwm maculatum never shows more
than three; and it is beyond all possibility of a doubt that the type described by
Bruguiere is Dolium tessellatum as here interpreted. The shape of the ribs, the
shape and size of the intervals, the occasional presence of a median line, and the
number of ribs on the body-whorl, as described by Bruguiere, are all in total agree-
ment with this interpretation. |

(3) This sentence refers to the absence of a deep sutural channel such as charac-
terises Doliwm galea or D. olearium. Sentence (4) needs no comment.

(5) Not only has Bruguiére given a perfectly precise description of the charac-
ters of the labrum, but he has recorded with admirable lucidity the history of its
development throughout the successive stages of growth of the shell. It is unneces-
sary to comment on the wonderful insight of the great naturalist at a period when
the science of zoology was still in its formative period. It is nevertheless astonish-
ing that so remarkable an observation should have passed unnoticed by all his

166 Memoirs of the Indian Museum. [Vor. VII,

successors who have either paid no attention to the ontogeny of the shell, or else have
totally misinterpreted its mode of growth.

Sentences (6) and (7) do not call for any special comment.

(8) and (9) The description of the colouring is remarkably complete, precise, and
consistent, and in several points is opposed to what is observed in Dolium maculatum ;
as for instance the remark that the maculations are usually less distinct on the early
part of the spire than at succeeding stages, just the opposite of what is usually seen
in Dolium maculatum ; or again the occurrence of non-spotted specimens, frequent
in the species described by Bruguiere, not known in Dolium maculatum; also the
occurrence of specimens uniformly tinted brown, which again entirely excludes Doliwm
maculatum.

(10) The distribution mentioned is to a large extent incorrect ; but if the name
were to be rejected on that account, it would be necessary to reject also for that
same reason an enormous proportion of the species established by Linnæus, Gmelin,
Lamarck, Sowerby and even Reeve, as, in former times, the localities from which
objects of Natural History were obtained were but too often incorrectly recorded.

In conclusion, it is very seldom, even at the present day, that a species is des-
cribed with the fulness, precision and detailed accuracy noticed in Bruguiére’s des-
cription of Buccinum tessellatum, and generally in all descriptions by that author.
If all specific distinctions depended on descriptions of similar merit, the troublesome
uncertainties of identification, of which the present imbroglio is but too common an
instance and which so seriously impede zoological research, would never happen.
There cannot possibly be two interpretations of Bruguiére’s description above ana-
lysed, and it would be inexcusable to substitute any subsequently published appella-
tion in place of his Buccinum tessellatum.

While adhering, in the text of his descriptions, to the limits of the genus Buccinum
as expanded by Linnæus, Bruguiére nevertheless classifies the species into two sec-
tions, of which the first one is explicitly stated to coincide with d’Argenville’s genus
Dolium. This must be the reason why Bruguiére has not adopted the Linnean
specific name “ dolium,” to avoid a repetition in the event of a reinstatement of
d’Argenville’s genus Dolium, the use of which was indeed resumed on the plates
illustrating Bruguiére’s descriptions.!

The next in date amongst the important works concerning the question
under consideration is Lamarck’s ‘ Histoire naturelle des animaux sans vertèbres.”
The general standard of Lamarck’s method compares unfavourably with that of
Bruguiere, though it is but fair to keep in mind the adverse circumstances that have
affected the work of all naturalists during his time and ever since; for he was not so
happily situated as Bruguiére who had the good fortune to terminate much of his
work on the very eve of that terrible catastrophy which has for ever retarded the
rate of progress of scientific knowledge and of all intellectual culture. Lamarck
deserves our admiration all the more for his undaunted perseverance amidst eircum-

! At least so I gather from the references to Bruguiére’s figures as given in the synonymy by Lamarck, Deshayes,
and Kiister. As already mentioned the plates of the Encyclopédie are not available in Calcutta.

1919. | E. W. VREDENBURG: Shells of the family Doliide. 167

stances unfavourable or even hostile to scientific research. Lamarck has followed
Bruguière in reinstating d’Argenville’s genus Doliwm. It may be noticed that,
in the generic diagnosis, Lamarck has introduced a clause “ labro per totam longi-
tudinem dentato vel crenato,’” so that, if it were desired to adhere strictly to
‚Lamarck’s definition, the name Dolium, s. str. would be applicable to the shells which
it is proposed to classify as Hudoliwm, rather than to those classified with Doliwm
galea. As however Dolium galea is the species first mentioned, it has been decided
that it is the type of the genus, and although it would be geologically and histori-
cally more logical to reserve the name Dolium, s. str. for the forms with denticulate
labrum, yet there is no distinct advantage to be gained in altering an established
usage.

In defining his Dokum maculatum, Lamarck has transcribed the synonymy of
Bruguiére’s Buccinum tessellatum, similarly differmg from Linnzeus’ and Gmelin’s
synonymy of Buccinum dolium in admitting the appositiveness of only two of the
figures from Martinis work. But for this correction, copied from Bruguiére, the
synonymy of Lamarck’s Doliwm maculatum is as hopeless a medley as that of Lin-
næus Buccinum dolium or of Bruguiére’s Buccinum tessellatum. From the synony-
my, there is no reason why Lamarck’s Dolium maculatum should represent a different
species from Bruguiére’s Buccinum tessellatum or Dolium tessellatum also mentioned
in the synonymy; no reason being given for superseding the name established by
Bruguiére. The specific name used by Linnæus is set aside by Lamarck presumably
for the same reason as by Bruguiére, that is on account of the re-establishment of
the generic name Dolium, to avoid a repetition. The description given by Lamarck
runs as follows: “ testa ovatoglobosa, ventricoso-inflata, tenui, alba; costis convexis,
distantibus, fulvo aut rufo maculatis : interstitiis stria prominula divisis.” The last
clause may indicate that it is especially the shell above described as Doliwm macula-
tum, Deshayes which Lamarck had in view, though, without any account of the
development of the shell, it is insufficient to clear up the uncertainties of the syn-
onymy. In conclusion, Dolium maculatum, Lamarck is just as uncertain as Buccinum
dolium, Linnæus.

The next important publication dealing with this subject is probably posterior
to the one of Lamarck above analysed, being the genus Doliwm in Sowerby’s
‘Genera,’ which presumably appeared in 1823. It contains an admirable figure of
a shell which Sowerby has called Dolium fimbriatum, and which tallies in every
respect with Bruguiére’s description. For those who, following the example of
Kiister, reject the specific names minjac and tessellatum, it is evident that the shell
in question must be known as Dolium fimbriatum, Sowerby.

In 1830, seven or eight years later therefore than the publications of Lamarck
and of Sowerby, Menke, in the second edition of the catalogue of his private collec-
tion, mentioned, as a new species, a Dolium costatum, with no other reference than
Martini’s figure 1082, therefore, one of the two figures specially selected by Bruguiére
as representing his Buccinum tessellatum, and by Lamarck as representing his Doliwm
maculatum. The date 1828 given by Tryon for this species is apparently incorrect,

168 Memoirs of the Indian Museum. [Vor. VII,

for, according to Küster, the species was established not in the first edition of Menke’s
catalogue (published in 1828 and not available in India), but only in the second.
The adoption of the same name for a similar shell by Deshayes in 1845 is a coinci-
dence, as Deshayes was not acquainted with Menke’s catalogue. Küster’s remark to
the effect that “for a long time previous to Deshayes, Menke had recognised this
species, fortunately under the same name, and had entered it as such in the second
edition of his Synopsis together with a diagnosis in the appendix,” is partly incorrect,
for Menke never seems to have described his Dolium costatum : the diagnosis in the
appendix referring not to Doliwm costatum, but to a Dolium tenue, from the coasts
of Syria, rightly considered by Tryon to be an immature specimen of Dolium
galea.

The first definite information subsequent to Bruguiere’s description and Sower-
by’s figure, that we can gather regarding the shells under consideration, is that
contained in the Xth Volume, published in 1845, of Deshayes’ new edition of
Lamarck’s “ Animaux sans Vertebres.’ A truly reliable character is at last noticed
by means of which Dolium maculatum:can be identified, namely the inferior number
of its primary ribs as compared with Dolium tessellatum, Bruguiere, for which Des-
hayes adopts the specific name minjac, Adanson. It is only, therefore, from the
date of Deshaye’s work that Doliwm maculatum can truly be recognised as a distinct
species. As already mentioned, Doliwm costatum is described by Deshayes as dis-
tinet from his Dolium minjac, the agreement in name with Menke, resulting, as above
noticed, from a coincidence.

Reeve, in 1849, in his monograph adorned with the superb illustrations of
Sowerby, has adopted the three species established by Deshayes, for one of which he
adopts the specific name fimbriatum, Sowerby, tessellatum, Bruguiére being rejected on
the unfounded plea that it refers both to D. fimbriatum and to D. maculatum, while
the specific name minjac mentioned in the synonymy is rejected on account of its
non-latinity. The short explanatory notices contain some inaccuracies, as in the
statement, reproduced from Deshayes, that Dolium fimbriatum is characterised “ by
the outer lip becoming strongly fimbriated on arriving at maturity,” just the opposite
of what really takes place as had already been observed by Bruguiére. In the
beautifully illustrated monograph published in 1857 as part of the Revision of Martini
and Chemnitz’ Conchilien-Cabinet, Kiister has unreservedly accepted Reeve’s con-
clusion, also including a fourth species, Dolium ampullaceum, Philippi, and adding a
fifth, Doliwm lischkeanum. The figure and diagnosis of Doliwm ampullaceum are
merely reproduced from Philippi’s work, the shell, of which only a dorsal view is
given, evidently representing a large specimen of D. tessellatum, Bruguiére. Doliwm
lischkeanum is founded on adult specimens of the same species, in which the aper-
tural thickening is therefore reduced, according to the adult characteristics already
so clearly defined by Bruguière. The name established by Bruguière is rejected for
the same unfounded reason as by Reeve, its supposed applicability to two different
species. The reason alleged for rejecting the specific name minjac is not without
some just foundation: ‘ Without necessarily ignoring all the rules of nomenclature,

1919. | E. W. VREDENBURG : Shells of the family Doliidæ. 169

the names capriciously bestowed by Adanson, cannot nevertheless lay claim to the
right of priority such as desired by Mr. Deshayes.”

Finally, in Tryon’s great work, the Manual of Conchology, the interpretation
adopted in the case under consideration is arbitrary in the extreme: D. maculatum,
Lamarck, D. fimbriatum, Sowerby, and D. costatum, Menke are regarded as mere
varieties (or even less than varieties) of a single species for which the much older
specific names minjac, Adanson, doliwm, Linneus, and tessellatum, Bruguiere are
rejected under the pretext that some uncertainty remains as to the particular variety
to which they might have been originally applied. If the various forms really did
represent mere varieties, it is obvious that the oldest name published would be that
of the species, and that the varieties would be named according to the precedence
in date recognised for each precise identification. Furthermore, not only does Tryon
reject these names on a pretext which, from his point of view, cannot be considered
valid, but, of the three other names that are allowed to stand, it is not the oldest,
Lamarck’s maculatum which is adopted as the name of the species, but Menke’s
costatum which is certainly newer by several years than either of the two others.

The foregoing lengthy discussion was unavoidable in order to arrive at some
definite conclusion regarding the names to be adopted for the two species recognised
in the present work. It should now appear sufficiently evident that the two names
to be adopted are Dolium tessellatum, Bruguière 1789, and Dolium maculatum,
Deshayes 1845.

For those who prefer archzeological erudition, however inconvenient, to the
more familiar nomenclature of long-established usage and tradition, there seems to
be no doubt that either Dolium minjac (Adanson) 1757, or Dolawm doliwm (L.) [| Hud-
desford | 1770, is the name that should be adopted for the shell above described as
Dolium maculatum, Deshayes. The adoption of such a course must carry with
it the usual warning that the pseudo-scientific security attained by the archæo-
logica] method has too often proved a delusion, for there is always the risk of an indus-
trious bibliographer discovering some forgotten monograph of earlier date than the
one relied upon as final. In any case, even as regards the substitution of Dolium
dolium to Dolium maculatum, this would be somewhat of a retrospective interpreta-
tion, -for no one who had not made an exhaustive study of the shells under considera-
tion could recognise which species is meant by Lister’s illustration. Neither the
figure nor the description of Lister nor the identification of Huddesford are of any
real help in identifying the species, and it is necessary to come down to the period of
Deshayes to obtain at last a sure method of recognising it.

If the Lamarckian appellation Doliwm maculatum be definitely adopted for one
of these species, it may perhaps be objected that to follow this specific name with
“ Deshayes ” as the author, as has been done in the present work, is not in accordance
with the recognised rules of nomenclature. The prevailing custom of appending to
the name of a species, whenever mentioned, the name of the earliest author who has
made use of such a specific name is in many cases of little scientific value. The
object of this apposition is to avoid confusion, the mention of the author’s name pur-

170 Memoirs of the Indian Museum. [Vor. VII,

porting to be an abbreviated reference to the earliest work in which a particular
specific name has received a definite meaning. If, however, the author quoted is one
who has only given an ambiguous definition, the abbreviated reference fulfils no
useful purpose. To be historically and scientifically useful, the reference should be
to the earliest author who has bestowed a definite meaning upon the name, whether
or not it may previously have been used in an unsatisfactory manner. For instance,
““ Dolium maculatum, Lamarck’ does not convey any useful information since it is
not possible, from Lamarck’s description, to ascertain which shell he had in view,
while “ Dolium maculatum, Deshayes”’ immediately gives the required clue since we .
need only refer to Deshaye’s work to know precisely which form is meant.

Characteristic differences between the two species.—From the foregoing description,
it is clear that the forms under consideration are both very well defined and remark-
ably constant, and it is easy therefore to detect the particular features that differen-
tiate them from one another. Amongst the more striking differences, the following
may be particularly mentioned.

Firstly. The protoconch in Dolium tessellatum is smaller, more prominent, less
oblique, darker-coloured than in Dolium maculatum. The secondary infilling of
porcellaneous shell-substance which occupies the whole interior of the protoconch in
the case of Doliwm maculatum does not reach the apex in the case of Doliwm

tessellatum.

Secondly. The number of primary ribs visible on the spire-whorls is three or four
in the case of Dolium tessellatum, two or three in the case of Dolium maculatum.
The intervening spaces in the case of Doliwm maculatum carry intercalary threads of
three orders, of which only those of the highest order are apt to disappear with
increasing growth. In the case of Doliwm tessellatum the intervals, in the portion
immediately following the protoconch, carry intercalary threads of two orders only,
all of which vanish entirely or almost entirely with increasing growth.

Thirdly. The number of primary ribs on the body-whorl of Dolium maculatum
is ten or eleven, and remains perfectly constant at all stages of growth. In the
case of Doliwm tessellatum the body-whorl of very small specimens carries twelve
primary ribs, the number increasing, by anterior additions, to as many as sixteen in
the case of very large specimens.

Fourthly. The intercalary decoration in the case of specimens of average size of
Dolium maculatum includes, in the broader intervals, threads of two orders, much
narrower than the primary threads. In the case of very large specimens some of
these intercalary threads may broaden into bands resembling the primary ribs, while
threads of a higher order may also appear. In the vast majority of specimens of
Dolium tessellatum there is practically no intercalary spiral sculpture on the body-
whorl. In the case of very large specimens there may appear some intercalary
bands, the number of which, in a single primary interval, never exceeds two. These
intercalary ribs of full-grown specimens of Dolium tessellatum are apt to assume the
maculated colour decoration of the primary ribs, while this is very rarely observed
in the case of Dolium maculatum,

1919. | E. W. VREDENBURG : Shells of the family Doliida. 171

Fifthly. The columella of Dolium tessellatum is shorter than that of Doliwm
maculatum. Except in the case of very large, quite adult specimens, it is always
rugose in the case of Dolium tessellatum, while it is smooth at all stages of growth in
the case of Dolium maculatum.

Sixthly. The vast majority of specimens of Dolium maculatum of all sizes have
a pertectly simple outer lip. When the outer lip is at all differentiated, it is feebly
expanded externally, feebly thickened and non-denticulate internally, with a more
or less wavy, feebly fimbriated edge. In the case of Dolium tessellatum the outer lip
is invariably differentiated, its special characteristics, in the case of the largest fully
adult specimens, being reduced to a mere internal thickening, while in every specimen
of small or medium size it is externally expanded, internally thickened and denticu-
late, and is conspicuously fimbriated all along the edge.

In conclusion, the differences are amply sufficient to justify the reference of
both forms to totally distinct species which are not even closely related. The simpli-
fication of the aperture in fully adult specimens of Dolium tessellatum, combined with
the adult reappearance of an intercalary sculpture, taken in connection with the
adult broadening of the intercalary ribs usually observed in Doliwm maculatum, gives
rise to a deceptive suggestion of convergence which, nevertheless, is quite superficial.
The history of the development of both shells clearly shows how widely divergent are
their zoological affinities.

Affinities of the two species.—Dolium maculatum appears to be an isolated species
amongst the recent fauna, and is not closely related to any other known living form.
In the characters of the outer lip, of the columella and of the umbilicus it distinctly
recalls Dolium galea, the type of the genus, as well as the closely related Dolium
melanostoma, Jay and D. variegatum, Lam., but is clearly separated by the wide-spaced
disposition of the primary ribs which, on the contrary, are close-set in Dolium galea
and the related forms. In any case Doliwm maculatum is to be classified as a typical
Dolium.

Dolium tessellatum, if we leave out of account the fully adult or gerontic indi-
viduals, is undoubtedly closely related to Dolium fasciatum, Brug., with which it
agrees as regards the posteriorly channelled aperture, the rugose columella, the
externally and internally thickened, denticulate and fimbriate outer lip, being dis-
tinguished only by the somewhat less ovoid outline of the body-whorl, and the
smaller number and wider spacing of the primary ribs, as well as some differences in
the colour decoration. It also shows a relationship though more distant to Dolium
zonatum, Green. In their apertural characters, Dolium fasciatum and D. zonatum
correspond so closely with Dolium crosseanum, Mont., the type of Hudolium, as to
justify their inclusion within that subgenus or section; the exact classificatory posi-
tion of Dolium tessellatum being rendered thereby, as already mentioned, somewhat
uncertain, since it may be regarded as a Æudolium when immature, a Doliwm, s. str.
when full grown. It is clearly at the mutual limit of the two groups, and indicates
how feebly defined is their separation.

The pliocene beds of the Mekran contain an extinct species which has provision-

172 Memoirs of the Indian Museum. [Vor. VII,

ally been named Dolium ormarense, and which, as shown by the characters of its
aperture, undoubtedly belongs to the same group as Doliwm tessellatum. The size,
number, and spacing of the primary ribs on the body-whorl is in complete agreement
with the recent form from which the fossil differs owing to the presence of a well-
developed intercalary rib of the second order in each interval at all stages of growth ;
the broader posterior primary intervals also showing indications of threads of a third
order. The spire is more sunken than in the recent form and recalls that of Dolium
maculatum.

In his first monograph on the tertiary fossils of Java (Tertidrschichten auf Java,
p. 40, 1879) Martin has mentioned the occasional presence of intercalary ribs on the
body-whorl of ‘‘ Dolium costatum,” that is therefore of D. tessellatum. On the
strength of this observation, Boettger (Die Tertiaerformation von Sumatra und thre
Thierreste, 2nd part, p. 84, pl. vi, figs. 4, 5; Palaeontographica, Supplementary
Vol. III) has referred to Doliwm costatum as a variety martini, a tertiary fossil from
Sumatra in which intercalary ribs are conspicuously developed at all stages of growth.
Nevertheless, since Martin specially mentioned that the occasional development of
intercalary ribs takes place on the body-whorl, the remark doubtless applies to the
adult character of full-grown specimens; especially as, in his latest work on the
subject (Samml. des geol. Reichs-Museums in Leiden, new series, Vol. I, 1899, p. 161)
Martin has specially described the complete or almost complete disappearance of the
interstitial decoration, with increasing growth, on the spire-whorls. In this latest
work, Martin has accepted Boettger’s identification without discussion, by simply
recording it in the synonymy. Yet the intercalary ribs, in the specimens figured by
Boettger, are far more prominent than is known ever to be the case in specimens of
Dolhium tessellatum of corresponding size. There is every reason to believe that the
fossil Doliwm martini is identical with the form provisionally named Dolium orma-
rense, especially as the spire seems to agree exactly with that of the Mekran fossil
and not with the true Doliwm tessellatum.

Another doubtful form is Doliwm modjokasriense, Martin, fossil from the later
tertiary of Java (Samml. des geol. Reichs-Museums in Leiden, new series, Vol. I,
p. 160, pl. xxv, fig. 370), in which the spire is disposed exactly in the same manner
as in Dolium maculatum, with exactly the same spiral decoration consisting of con-
spicuous ribs or threads of three orders, with indications of a fourth, this decoration
being similarly continued on the corresponding posterior portion of the body-whorl.
Anteriorly to the level of the suture, the body-whorl is said to differ from that of
Dolium maculatum owing to the presence of only two alternating orders of spiral
ornaments instead of three; only, this part of the solitary available specimen is so
poorly preserved as to permit of a doubt as to whether the ribs interpreted as
representing the first order may not partly correspond with the expanded ribs of the
second order as frequently observed on adult specimens of Dolium maculatum, in
which case the fossil under consideration would be a specimen of D. maculatwm such
as is already known in a fossil condition from the upper tertiary of the Mekran.
Otherwise, if the interpretation accepted in the original description be correct, the

1919.] E. W. VREDENBURG: Shells of the family Doliide. 173

diagnosis merges into that of Dolium ormarense. Nevertheless the prominence of the
spiral ornaments of the second and third order on the spire-whorls recalls Doliwm
maculatum more than D. ormarense. This question cannot perhaps be definitely
settled without the discovery of better preserved specimens, especially such as might
exhibit the apertural characters.

Another fossil species from the pliocene of Java, Doliwm hochstetteri, Martin
(Samml. des geol. Reichs-Museums in Leiden, new series, Vol. I, p. 162) is perhaps
also to be classified in the same group with Doliwm tessellatum, from which it is dis-
tinguished by its extremely flattened spire and the posterior inflation of the body-
whorl. The details of the aperture are unfortunately not known.

Lastly, another upper tertiary fossil from the Mekran, provisionally named
Dolium arabicum, and probably identical with Dolium townsendi, Newton (Geol.
Mag., dec. 5, Vol. II, p. 301) is probably also related to Dolium costatum, with which
it agrees in the shape and ornamentation of the spire, but from which it differs owing
to the irregular distribution of the spiral ribs on the body-whorl. In this case also,
owing to the absence of the apertural characters, the precise affinities of the species
remain uncertain.

Distribution of Dolium tessellatum and Dolium maculatum.—The areas over which
the two species are found at the present day are to a large extent distinct, though
they partly overlap. Dolium tessellatum does not seem to spread further west than
the eastern portion of the Bay of Bengal, nor Dolium maculatum further east than
the Malay Islands, while, in a westward direction, D. maculatum occurs apparently
throughout the Arabian Sea, and, in an eastern direction, D. tessellatum extends as
far as Japan. The geographical range of D. tessellatum appears to have been more
extensive in former geological times than at the present day, for its occurrence as a
fossil in the Mekran beds indicates its former presence, in later tertiary times, in
the area now occupied by the Arabian Sea.

1ll.—THE SPECIFIC IDENTITY OF DOLIUM LUTEOSTOMA, KUSTER, WITH DoLium
VARIEGATUM, LAMARCK, AND OF DOLIUM MAGNIFICUM, G. B. SOWERBY,
WITH DOLIUM CHINENSE, DILLWYN.

On a previous occasion (Journal As. Soc. Bengal, new series, Vol. XIV, 1918,
p. 449) I have commented upon the apparently discontinuous distribution of Dolium
variegatum, Lamarck, a conspicuous shell previously regarded as special to the Austra-
lian region, but which has now been shown to occur also in the northern part of the
Arabian Sea. The collections in the Indian Museum include specimens from Maskat,
and I have also ascertained the presence of the shell at Karachi. It was moreover
noticed that a shell found at Charbar on the northern shore of the Gulf of Oman, by
Mr. Townsend, and referred by Melvill and Standen to “‘ Doliwm galea var. luteosto-
mum” (Proc. Zool. Soc. London, 1901, Vol. II, p. 385), probably also represents a
specimen of Dolium variegatum.

174 Memoirs of the Indian Museum. [Vor. VII,

The specimens of Doliwm variegatum from the collections of the Indian Museum
hitherto studied were all of relatively moderate dimensions. Since the note above
alluded to was written, several more specimens have come to notice, unfortu-
nately of uncertain origin, remarkable for their large dimensions, and clearly corres-
ponding with the figured types of Doliwm luteostoma and of the synonymous Doliwm
japonicum, Dunker. At the same time, their specific identity with Doliwm variegatum
does not admit of any doubt, and hereby is explained the occurrence of Doliwm
luteostoma in the Arabian Sea as recorded by Melvill and Standen. It now remains
to be seen whether Doliwm variegatum itself is to be treated as specifically identical
with Dolium galea, since the synonymous Dolium luteostoma is regarded by Melvill
and Standen as a mere variety of that species.

With the exception of Küster, in his monograph of the genus Dolium published
in 1857 as part of the revised edition of Martini and Chemnitz’ “ Conchilien-Cabinet,”’
the authors who have dealt with these particular shells have only given short
diagnoses insufficient for identifying the various species. Even in the case of Kiister’s
work, the excellent descriptions are mostly unaccompanied by comparative criteria. It
is therefore necessary to rely principally upon illustrations and upon actual specimens
for the discrimination of the species. With regard to the possible specific distinctness
of the forms variously described a D. galea, D. variegatum, and D. luteostoma, the
question is still further complicated by the necessity to take into account a Dolium
melanostoma, Jay (1839, Cat. Mus., p. 124, pl. viii-ix), with which D. luteostoma is
regarded by Tryon (Manual of Conchology, Vol. VII, p. 261) as possibly identical.

Apart from Melvill and Standen, in the above quoted reference, none of the
authors who have dealt with Doliwm luteostoma appear to have had any hesitation
in considering it to differ specifically from Dolium galea, from which it is distinguished,
at the first glance, by its fewer and wider ribs. Tryon is the only author who has
thought it worth while to compare Dolium variegatum with Dolium galea and with
Dolium luteostoma and records, as the only distinction from the two species last
named, a relatively shallower sutural groove or rather depression, and a more elevated
spire for D. variegatum.

It need scarcely be mentioned that differences of this kind, depending on mere
matters of degree and totally ignoring individual variation, are worthless from the
point of view of precise specific discrimination. I have vainly endeavoured, with the
material now available, to discover other distinguishing features. The degree of
elevation of the spire is extremely variable from one specimen to another, as is
obvious from the two specimens of Doliwm variegatum figured in Reeve’s monograph.
Of the two specimens of Dolium variegatum as interpreted by Küster, illustrated in
plate Ixiii of his monograph, the larger one has so short a spire that it has unhesi-
tatingly been referred to Dolium luteostoma by Tryon; (the figure of the smaller
specimen being copied from Reeve). Moreover, the degree of elevation of the spire
may vary widely at different stages of growth even in the same specimen; as in
the case of the larger of the two specimens figured by Reeve, and also in the case of
a very fine specimen in the collections of the Indian Museum, measuring 168 x 139 mm.,

1919. ] E. W. VREDENBURG: Shells of the family Doliide. 175

in which the suture becomes increasingly oblique with increasing growth, so that, at
earlier stages, the spire is extremely depressed, and gradually becomes more elevated
in the full-yrown shell. In the case of such specimens, the earlier whorls exactly
agree with the diagnosis of Dolvum luteostoma, the later ones with that of Doliwm
variegatum. As to the sunken disposition of the suture in various individuals or in
various portions of one specimen, it is more or less pronounced inversely to the degree
of obliquity, but is always distinct.

We may conclude that Doliwm variegatum, Lamarck, and Dolium luteostoma,
Kiister, undoubtedly represent the same species, which should be known by Lamarck’s
designation which is older. |

The shell is undoubtedly very closely related to Dolium galea, Linn., though it
cannot be referred to the same species as has been done by Melvill and Standen who,
as already mentioned, have catalogued it from the Gulf of Oman as “ Dolium galea
var. luteostomum.” The spiral ribs, even taking full account of individual variations,
are always fewer in Dolium variegatum than in Dolium galea. At the largest stages
of growth, the number of main ribs, in Doliwm variegatum, never exceeds nineteen,
and is usually less (frequently fifteen), while Dolium galea has at least twenty ribs
even in the case of small specimens, and usually more. There is also a very distinct
difference in the general shape of both species at all stages of growth, the convexity
of the body-whorl being more evenly continuous in Dolium variegatum than in Dolium
galea, in which latter species there is a decrease in the degree of curvature along the
zone of maximum width, communicating a slightly flattened appearance to the sides
of the shell.

There remains to be considered the case of Dolium melanostoma, Jay, a remark-
ably handsome shell, the name of which refers to the dark colour pervading the
aperture, principally over the columellar lip. It seems to occur abundantly through-
out Polynesia and as far north as the Hawaiian Islands, but is not represented in the
collections of the Indian Museum. The two illustrations hitherto published, in the
works respectively of Jay (loc. cit.) and of Reeve (Monograph of the Genus Dolium,
species 2) are in every respect consistent with one another except as regards the
number of principal spiral ribs on the body-whorl, which amounts to twenty on the
specimen figured by Reeve, but which does not exceed fifteen in Jay’s original type.
The number of ribs varies therefore approximately within the same limits as in the case
of Dolium variegatum, of which, judging by this character alone, Doliwm melanostoma
might be merely a colour-variety. But there are other differences besides those of
colour. Judging from the apparently excellent illustrations, the outline of the spire
is more subulate, with less convex whorls than in Dolium variegatum. The sutures
are decidedly less sunken than in either Doliwm galea or Dolium variegatum. The
absence of spiral ribs on the anterior winding terminal bulge of Dolium melanostoma
already noticed by Reeve as a good distinction from Doliwm galea similarly distin-
guishes it from Doliwm variegatum. The spire, in Jay’s original type, is taller than
would seem ever to be the case with Doliwm variegatum, while, in the specimen figured
by Reeve, it also exceeds the average of Doliwm varvegatum ; it is therefore probably

176 Memoirs of the Indian Museum. [Vor. VIL,

more elongate, on an average, in Dolium melanostoma than in Dolium variegatum,
though further information is necessary to make certain about this point. The
narrow intercalary ribs appear to alternate with the larger ones much more regularly
in Dolium melanostoma than in either Doliwm galea or Dolium variegatum. In this
respect, as also with regard to the feebly sunken sutures, Doliwm melanostoma recalls
Dolium chinense, Dillwyn, but its ribs do not become so completely flattened out as in
the last-named species. Lastly, Dolium melanostoma, like Dolium galea, lacks the
brown maculations of Dolium variegatum and Dolium chinense.

We may conclude that Dolium melanostoma is specifically distinct both from Dolium
galea and from Dolium variegatum. There exists therefore, at the present day, a
group of three closely related species, each with its special geographical distribution :
Dolium galea inhabiting the Mediterranean and Atlantic, Dolium variegatum charac-
terising the Indian Ocean and western shores of the Pacific, and Doliwm melanostoma
in the central Pacific region. As pointed out on a previous occasion, it is improbable
that so conspicuous a species as Dolium variegatum should have been overlooked in
the Bay of Bengal and Malay region if it really lived in those portions of the ocean ;
and its occurrence in the northern part of the Arabian Sea suggests therefore a
discontinuous distribution for that species. It is known to exist along the northern
and eastern coasts of Australia, but we cannot at present ascertain whether it spreads
continuously across the intervening seas from Australia to Japan, in which latter
region it is also known to occur, this being the habitat of Dolium japonicum, Dunker,
synonymous with Doliwm luteostoma and with Doliwm variegatum. We know that
Dolium variegatum has been in existence since miocene times, and from the distribu-
tion of the fossil occurrences, we may conclude that its present discontinuous distri-
bution is due to the local shrinking of a once connected area. No fossil occurrences of
Dolium galea or Dolium melanostoma have been discovered, but our knowledge of the
later tertiary marine faunas of tropical regions is as yet too incomplete to lay stress
upon this circumstance.

11.

The second point to be discussed is the relationship to Doliwm chinense, Dillwyn,
of a form described in 1904 by G. B. Sowerby as a new species under the name of
Dolium magnificum (Proc. Malac. Soc. London, Vol. VI, p. 7, fig. 1). The specimen
described and figured by Sowerby was obtained by the late General Tripe from China,
from the very home, therefore, of Doliwm chinense.

We have evidently to deal, here, with another of the numerous instances in which,
in the genus Dolium, large individuals are apt to differ considerably in appearance from
smaller specimens of the same species; though in the present case the difference is not
of an extremely marked degree. The specimen of Doliwm chinense figured in Reeve’s
monograph measures 75 mm. in height. A specimen of nearly the same size, closely
corresponding with the one figured by Reeve, is to be found in the collections of the
Indian Museum, together with some smaller ones and a much larger one which will be
further alluded to. The maximum height of Doliwm chinense, according to Küster, is
33, therefore more than 80 mm. The type-specimen of Dolium magnificum has a

19192] E. W. VREDENBURG : Shells of the family Doliide. an

height of 110 mm., not disproportionately greater therefore than the largest previously
known specimens of Dolvum chinense. In addition to its large size, it is considered to
be further differentiated by the character of its spiral threads, said to be flatter than
in Dolium chinense. At all stages of growth, Doliwm chinense is always characterised
by its relatively flat ribs, this feature constituting a good distinction from Doliwm
variegatum, with which it has been erroneously united by Philippi and by Tryon. The
collection of the Indian Museum includes a specimen, unfortunately of unknown
origin, measuring II5 X 98 mm., even larger therefore than G. B. Sowerby’s type of
Dolium magnificum, with which it otherwise corresponds in every respect. The ribs
of the body-whorl, as in the case of Sowerby’s type, are still flatter than in specimens
of Dolium chinense of moderate size; the ornamentation, indeed, consisting of linear
grooves rather than ribs. Yet, the spire-whorls plainly show that, at earlier
stages of growth, the ribs are less flattened and are disposed exactly as in typical
specimens of Doliwm chinense. The protoconch of the specimen corresponding with
Dolium magnificum is of exactly the same size and shape as in typical specimens
of Dolium chinense; indeed, all the earlier portion of the spire is absolutely undistin-
guishable. The paired disposition of the maculated ribs observed in the type of
Dolium magnificum, and in the corresponding Calcutta specimen, is frequently matched
amongst typical specimens of Doliwm chinense. It seems obvious that Doliwm magni-
ficum is specifically identical with Doliwm chinense, of which it represents a full-grown
stage.

Even these exceptionally large specimens of Doliwm chinense do not nearly
approach the maximum dimensions of Doliwm variegatum, which reaches as much as
230 mm., and ranks amongst the largest known gastropods. Dolium chinense is there-
fore well distinguished from Doliwm variegatum by its smaller average size, its feebly
sunken sutures, and its more crowded and flatter ribs, the difference becoming
especially marked in large specimens of both species, since, in the case of Doliwm
variegatum, there is no indication, with increase of size, of the extreme flattening
characterising the full-grown stage of Dolium chinense.

NES OF THE DOLIIDÆ IN THE COLLECTION OF THE INDIAN MUSEUM.

While engaged upon the study of the tertiary fossil Doliidæ in the collections of
the Geological Survey of India, I had occasion to examine, for purposes of com-
parison, the rich series of recent shells of this family in the zoological collections of
the Indian Museum. Many of these valuable specimens had remained unnamed, and,
as the work of comparison necessitated the identification of the recent as well as of
the fossil specimens, advantage may be taken of the present opportunity to place on
record the contents of the collection of recent shells, all the more so as, in the ma-
jority of cases, we possess precise details as to the locality from which the specimens
were obtained; the information thus obtainable supplementing in several instances
that previously published.

According to the classification schemes of Fischer (Manuel de Conchyliologie,

178 Memoirs of the Indian Museum. [Vor. VII,

p. 661) and of Cossmann (Æssais de Paleoconchologie comparée, fase. V, p. 136), the
Doliidæ include two genera, Dolium and Pirula, while Dolium is itself divided into
three subgenera, namely Doliwm, s. str., Hudolium and Malea. Several authors treat
Malea as a separate genus.

: Genus Dolium, d’Argenville, 1757.

The species of Doliwm, both fossil and recent, hitherto recorded are provisionally
srouped in the following list in which are included also two fossil species from the
Mekran distriet, of which the descriptions are not yet published, and which are
mentioned with provisional names, but which probably correspond with two species
which have already been named and figured. This particular question will be fully
discussed, it is hoped, at no very distant date in the publications of the Geological
Survey of India.

The proposed limits of the subgenus Hudoliwm have been discussed in a fore-
going paper.

The species included within the three subgenera of Dolium may be grouped in
the following nine divisions:

Dolium, >. str.

It is proposed to restrict Doliwm, s. str. to those shells in which the outer lip is
simple or internally slightly thickened, and the columella smooth at all stages of
growth.

Ist Division.

This division which may be called the group of Doliwm galea (Linn.), after the
type of the genus, includes globose or slightly ovoid shells of large or very large size in
which the spiral ribs are mostly contiguous or nearly so. The following forms have
been recognized :—

Dolium galea (Linn.), in which the ribs are very numerous and the sutures deeply
sunken. There is a marked decrease in the degree of curvature along the zone of
maximum width, this portion of the shell tending therefore to assume a sub-cylindri-
cal outline.

Dolium variegatum, Lamarck, in which the ribs are fewer and the curvature more
continuous than in Doliwm galea, from which it is usually distinguished also by the
presence of brown maculations along many of the ribs. Küster’s Dolium luteostoma
and Dunker’s D. japonicum are synonymous.

Dolium melanostoma, Jay, in which the number of ribs and the shape of the body-
whorl are about the same as in D. variegatum from which it is distinguished by its
more subulate spire, the feebly sunken sutures, the absence of maculations on the
ribs, and the dark tint of the aperture.

Dolium chinense, Dillwyn, distinguished from D. variegatum by its smaller size,
more numerous and more flattened ribs, and its feebly sunken suture. Dohum
magnificum, G. B. Sowerby, represents the full-grown condition of this species.

Dolium olearium, Bruguiére, which resembles Doliwm variegatum in general out-

1919. ] E. W. VREDENBURG : Shells of the family Doliide. 179

line; sutures sunken; ribs about as numerous as in D. variegatum, only much flatter
and separated by mere linear shallow grooves, without intercalary ribs.

Dolium perdix, Linnaeus, larger and more ovoid than Dolium chinense ; sutures not
appreciably sunken; ribs quite flat and about as numerous as the primary ribs of
Dolium chinense, separated by linear grooves without the intercalary ribs that
characterise Doliwm chinense. Nevertheless, the resemblance to Doliwm chinense
becomes very close in the case of immature specimens of Dolium variegatum, which
are apt to be relatively much more globose than the adult. The resemblance is
accentuated by the fact that the intercalary ribbing is absent in the case of immature
specimens of Doliwm chinense. Nevertheless, the immature specimens of Doliwm
chinense are distinguished by the conformation of the terminal zone of accretions
which, when viewed dorsally, appears somewhat more bulging and convex, and is
separated from the convexity of the base by a slightly better defined concavity than
in the case of the immature specimens of Dolium perdix. The protoconch of Dolium
perdix is considerably larger than that of D. chinense.

Tryon regards Dolium cumingii, Hanley, D. deshayesi, Reeve and D. testardi,
Monterosato, as varieties of D. olearium; a most unlikely interpretation, for, judging
from the excellent published illustrations, the absence of a circumsutural channel, the
shape and ornamentation of the spire, the sculpture and decoration of the body-
whorl, suggest a close relationship, if not specific identity, with Dolium perdix.

2nd Division.
Large globose shells with the ribs separated by wide intervals.
Only one species can with certainty be ascribed to this division: this is Dolium
maculatum, Lamarck. The Java fossil Doliwm modjokasriense, Martin, as already
explained (p. 172), is perhaps identical.

3rd Division.

This division only includes the fossil Doliwm losariense, Martin (Samml. des
geol. Reichs-Museums in Leiden, new series, Vol. I, p. 163, pl. xxv, figs. 377, 378),
distinguished from all other species by the marked posterior angulation of the body-
whorl. So far as can be made out its outer lip is simple.

Subgenus Eudolium, Dall, 1889.
(= Doliopsis, Monterosato, 1872, non Conrad).

According to the interpretation here proposed, the subgenus Hudoliwm includes
those forms in which, throughout the greater part or the whole of the life of the
shell, the aperture is somewhat channelled posteriorly, the columella rugose, and the
outer lip internally denticulate.

4th Division.

The shells of this division, which may be distinguished as the group of Doliwm
tessellatum, include umbilicated shells in which the primary ribs are wide-spaced. In
Dolium tessellatum itself, the only form of this group in which the development of

180 Memoirs of the Indian Museum. [Voz. VII,

the shell is entirely known, the apertural denticulations and the collumellar rugosities
disappear in the case of thoroughly adult individuals, the species establishing there-
fore somewhat of a link between Doliwm, s. str. and Hudolium.

In those forms of which the apertural details are known the denticulations are
bifid. The forms which may be either definitely or provisionally classified in this
group are the following :—

Dolium tessellatum, Bruguiére, in which conspicuous intercalary ribs appear only
in fully adult shells.

Dolium ormarense, Vred., fossil, in which conspicuous intercalary ribs are seen at
every stage of growth ; Doliwm costatum var. martini, Boettger (Die Tertiaerformation
von Sumatra und ihre Thierreste, Part II, p. 84, pl. vi, figs. 4, 5, Palaeontographica,
Suppl., Vol. III) being perhaps identical.

Dolium hochstetteri, Martin (Samml. des geol. Reichs-Museums in Leiden, new
series, Vol. I, p. 162), characterised by its extremely depressed spire and posteriorly
inflated body-whorl. The details of the aperture are not known, and the classifica-
tion of this shell is therefore provisional.

Dolium arabicum, Vred., fossil, characterised by the irregular distribution of the
ribs on the body-whorl; D. townsendi, Newton (Geol. Mag., dec. 5, Vol. II, p. 30T),
being probably identical. As in the case of the preceding species the details of the
aperture are not known and the classification of the shell is hkewise provisional.

5th Division.

In this division, which may be distinguished as the group of Dolium fasciatum,
the shell is umbilicated, the ribs are crowded and close-set, the denticulations bifid.
It includes the following species :—

Dolium fasciatum (Bruguière), in which the body-whorl is practically destitute of
intercalary ribbing, and the outer lip thickened both internally and externally.

Dolium zonatum, Green, in which all the intervals carry intercalary threads and
the labrum is thickened only internally.

6th Division.

The shells of this division, which may be distinguished as the group of Doliwm
crosseanum, differ from those of the preceding group merely in the absence of an
umbilicus. The denticulations are similarly bifid. This group includes the following
species:

Dolium crosseanum, Monterosato, the genotype of Doliopsis, Monterosato and of
Eudolium, Dall, characterised principally by its crowded spiral ornamentation closely
resembling that of Doliwm zonatum.

Dolium cinguliferum (Brown) (= D. fasciatum, Borson, non Bruguiere), fossil, of
smaller size, with fewer primary ribs.

Dolium muticum, Michelotti, fossil, strongly tuberculate, with labrum thickened
externally as well as internally.

1919.] E. W. VREDENBURG : Shells of the family Dolide. 181

Dolium subfasciatum, Sacco, fossil, more delicately tuberculate than -D. muticum,
with the labrum similarly thickened externally.

Dolium stephaniophorum (Fontannes), fossil, delicately tuberculated like D. sub-
fasciatum, but without the external thickening of the labrum.

Dolium antiquum, Sacco, also fossil, is only represented by a single incomplete
cast, and its specific distinctness from D. muticum is doubtful.

7th Division.

This division only contains at present Doliwm verrillit, Dall (Bull. Mus. Comp.
Zool., Vol. XVIII, 1889, p. 233, pl. xxxv, fig. 12), which is non-umbilicated, and which
differs from all other forms of Hudolium owing to the callous thickening of its apertural
margins, a character establishing a transition towards the subgenus Malea. The den-
ticulations of the outer lip are coarse and apparently simple as is the case to a large
extent in Malea.

Subgenus Malea, Valenciennes, 1833.

The subgenus or genus Malea includes shells in which the aperture is callous and
exhibits coarse denticulations not only along the outer lip, but also along the colu-
mellar lip, both opposite the columella and opposite the base of the penultimate whorl.

8th Division.

In this division, which may be distinguished as the group of Dolium pomum, the
degree of callosity of the aperture is moderate. It includes the following species :—

Dolium pomum (Linn.), ovoid, with feebly prominent, close-set, broad ribs and
without intercalary ornaments.

Dolium orbiculatum, Brocchi, fossil, distinguished from the foregoing by the
somewhat more crowded denticulations of the outer lip.

Dolium pro-orbiculatum, Sacco (= D. denticulatum, Deshayes sec, Hoernes), more
globular than D. orbiculatum, with more even outline of the posterior part of the
body-whorl, and with the spire generally taller and more subulate.

gth Division.

This division is characterised by the excessive degree of callosity of the aperture.
It may be distinguished as the “ group of Dolium ringens.” It includes the following
species :—

Dolium ringens (Swainson), large, globose, with numerous fairly prominent ribs.

Dolium camurum, Guppy (Quart. Journ. Geol. Soc., Vol. XXII, 1866, p. 287,
pl. xvii, fig. 9), fossil, differmg from Malea ringens in its smaller size, and in the
flatter, less sharply defined spiral ribs separated by very narrow, but not deeply
sunken intervals; the external decoration therefore somewhat recalling that of the
shells belonging to the group of Malea pomum, and establishing a genealogical link
between the two groups. In the shortness and prominence of the bunch of folds
situated on the base of the penultimate whorl, and in the consequent great depth of
the embayment separating it from the anterior columellar group, the shell entirely
agrees with Malea ringens, of which it is clearly an ancestral pre-mutation.

182 Memoirs of the Indian Museum. [Vor. VII,

It has not been possible to include in the above list Dolium dunkeri, Hanley
(Proc. Zool. Soc., 1859, p. 43I), which does not appear to have been figured and
cannot be identified from the short description, nor Dolium procellarum (Bull. Soc.
Malac. France, 1885, Vol. II, p. 247), the description of which is not accessible in
India.

The geographical and geological distribution of these various shells, so far as
known, is the following :—

Oligocene.
Lower
Pliocene.
Quaternary.
Recent.

Dolium galea (Linn.) oe | ——| Mediterranean and Atlan-
tie.

melanostoma, Jay | | | ————| Pacific.

: Australia, Japan and Ara-
bian Sea; fossil in the
pliocene of Java, and of
the Mekran.

,. variegatum, Lam. ne] | | |———__ |. | |——_

chinense, Dillwyn a I-—— | ...|... |———| China; ? Malay Peninsula:
fossil in the upper mio-
cene of Java.

olearium, Bruguière .. | —— Indian Ocean, Philippines.
|

perdix (Linn.) 2. | —— Indo-Pacific region and
western Atlantic.

maculatum, Desh. sal | | SE Arabian’ Sea to Moluccas,
| fossil in the pliocene of
| the Mekran (and of
Java? = D. modjokas-
riense, Martin ?), and in
the quaternary of the
‘ Pulicat lake near Mad-
ras.

losariense. Martin de | —— Fossil in the pliocene of
| Java and of the Mekran.

ormarense, Vred. | | — | Fossil in the pliocene of

LE]

1919.]

D. (Eudolium) Hochstetteri, Mart.

a arabicum, Vred...

a fasciatum ( Brug.)

= zonatum, Green ..

u crosseanum, Mont.
sf cinguliferum
(Bronn).

muticum, Mich. ..

= subfascratum,Sacco,

2 stephaniophorum
(Font.).

> verrillii, Dall.

D (Malea) pomum (Linn.) ee

vr orbiculatum.,
Brocchi

ig pro-orbiculatum,
Sacco

en ringens, Swainson

i camurum, Guppy

| Oligocene.

MIOCENE.

Lower.

|
|

E. W. VREDENBURG : Shells of the family Doliide.

183

Pliocene.

”

Quaternary. ||

Recent.

| Fossil in
Java.

the pliocene of

| Fossil in the pliocene of
the Mekran: D. town-

| sendi, Newton is pro-
bably identical.

— | Western Bay of Bengal to
Japan.

— China and Japan: a fossil
| | variety in the pliocene
of Java.

— Mediterranean and West
Indies.

Fossil in the miocene and
pliocene of the Piedmont.

Fossil in the oligocene of
| Liguria; D. antiquum,
_ Sacco is perhaps identi-
aca

Lower (middle) miocene of
the Piedmont.

Fossil in the upper miocene
of the Piedmont, and
in the pliocene of the
Piedmont and of the
Rhone valley.

West-Indies.

“Red Sea to Society Is-
lands.”

Fossil in the upper miocene
of the Piedmont, and in
the pliocene of the Alpes-
Maritimes, of Piedmont,
and of Tuscany.

Fossil in the Vienna region.

Eastern Pacific.

Fossil in Jamaica and San
Domingo.

184 Memoirs of the Indian Museum. [Vor. VII,

The presence of Eudolium, as here defined, in the oligocene of Europe, while, in
the Eastern Seas, it is only known from upper miocene to recent times, and the
occurrence of Malea exclusively as a fossil in Europe, exclusively as a living group in
the Indo-Pacific, are typical instances of a large number of similar cases which at one
time were thought to imply an easterly migration of the tertiary fauna of Europe
into the Indo-Pacific region of the present day. It has now been shown that the
great majority of these cases resulted from our hitherto deficient knowledge of the
fossil contents of the tertiary formations of Asia, and, with increasing research, most
of these supposed instances have now vanished.

It is clear, however, that amongst the minor divisions enumerated in the pre-
ceding list some have a well-defined geographical as well as geological distribution,
The most archaie group, that of Dolium crosseanum, is entirely restrieted at the
present day as well as in former geological times to the Mediterranean and Atlantie
region. It is, so far as our present information goes, the geologically oldest group,
and that from which the other forms of Doliwm seem to have been derived. It is
possible therefore that the genus Dolium may truly have had a western origin.
Dolium, s. str., as here understood, has not been found in a fossil condition in the
west. Dolium galea is special to the Mediterranean and Atlantic. The other forms of
Dolium, s. str., as here understood, are essentially Indo-Pacific, though Doliwm perdix
has also spread to the West-Indies and to the coast of Brazil, evidently through the
former marine connection across Central America which has afforded a passage to
other Indian species, either still living in the Indian region, or known in a fossil con-
dition, as has already been pointed out in the publications of this Department
(Memoirs of the Indian Museum, Volume VI, p. 124).

As already pointed out by Sacco (Moll. dei terr. terz. del Piemonte e della Liguria,
Part VIII, 1891, p. 22), the occurrence throughout the pliocene of southern Europe
of a Malea closely related to the tropical Doliwm (Malea) pomum indicates the persis-
tence, in the Mediterranean region, of a warm climate down to the very eve of the
Glacial Epoch.

Amongst the additions to our knowledge of the distribution of these shells
furnished by the Calcutta collection, one of the most interesting is that concerning
the presence, at Maskat, of Doliwm variegatum, hitherto only known from the Austra-
lian region and Japan. While there is every reason to believe that the distribution
of Dolium variegatum at the present day is discontinuous, its occurrence in a fossil
condition in the Mekran beds as also in Java furnishes an easy clue to its present
occurrences, the distribution of the shell having been more extensive in former geological
times than at the present day. Another species, Dolium tessellatum which, at the
present day, does not appear to extend further west than the eastern part of the Bay
of Bengal, also occurs in a fossil condition in the Mekran region. Just as in the case
of Dolium variegatum, it seems now to have disappeared from the shores of the Indian
Peninsula, only, unlike D. variegatum, it has also disappeared from the northern
shores of the Arabian Sea. Consequently D. tessellatwm does not exhibit at the
present day the discontinuous distribution observed in the case of Dolium variegatum,

1919. | E. W. VREDENBURG : Shells of the family Doliide. 185

The following recent species are represented in the collections of the Indian
Museum :—
Dolium galea (Linn.)
» variegatum, Lam.
à chinense, Dillw.
Br olearium, Brug.
> pendia (Kian.)
7 maculatum, Lam.
D. (Eudolium) tessellatum, Brug.

Be fasciatum (Brug.)
a zonatum, Green.
D. (Malea) pomum (Linn.)
a ringens, Swains.

The only important species not represented are Doliwm melanostoma, Jay, and the
rare deep-sea forms D. crosseanum, Monterosato and D. verrillit, Dall, the latter of
which is at present known only from a single specimen.

The following is a list of the specimens in the Indian Museum collection :—

Dolium galea (Linnæus).

““ Europe,’ one specimen.

Dolium variegatum, Lamarck.

Maskat, two specimens.

Port Jackson, one specimen.

Locality uncertain, five specimens, three of which are of very large size.

I have ascertained the presence of this shell also at Karachi.

As already mentioned, this species is also known in a fossil condition from the

pliocene of Java, and of the Mekran.

Dolium chinense, Dillwyn.

? Singapore, three specimens.

Locality uncertain, one large specimen, measuring 115 x 98 mm., coinciding exactly
with the figured type of Dolium magnificum, Sow.

Also fossil in the upper miocene of Java.

Dolium olearium, Bruguière.

Andamans, one specimen.
Nicobars, one specimen.

? Singapore, one specimen.
Trincomali, two specimens.
Locality uncertain, five specimens.

186 Memoirs of the Indian Museum. [Vor. VII,

Dolium perdix (Linnaeus).

Andamans, four specimens.

Australia, one specimen.

Cocoa Islands, two specimens.

Mauritius, three specimens.

Gulf of Suez (Capt. R. B. S. Sewell, 8-1-1917), two specimens.
Trincomalee, four specimens.

Locality uncertain, three specimens.

Dolium maculatum, Lamarck. —

Amboina, one specimen.

Andamans, three specimens.

Off the east coast of the Andamans (Lat. 13°17’15” N., Long. 93°10’25” E.), a dead
shell dredged from a depth of 185 fathoms.

Arakan, one specimen.

Balasore Bay, three specimens (Bengal Fisheries, M 4225, M 4498, M 221°),

Bombay and probably Kachh: four specimens (M 2323).

Ceylon or Kachh, one large specimen.

Chandipore, one specimen.

Penang, one specimen.

Sandheads, one specimen, remarkable for its extremely sunken spire (A. Milner).

? Singapore, one specimen.

Tavoy coast, four specimens.

“ Indian Seas,” one large specimen.

Locality unknown, nine specimens.

I have ascertained the existence of this shell also at Puri.

In a fossil condition it is known from the pliocene formations of the Mekran and
from the post-tertiary of the Pulicat Lake near Madras. Doliwm modjokasriense,
Martin, from the upper tertiary of Java, is perhaps identical.

Dolium (Eudolium) tessellatum, Bruguiere.
Andamans, nine specimens, one of which is from the South Andaman.
Nicobars, one specimen.
Locality uncertain, five specimens.
Also fossil in the upper miocene and pliocene of Java and in the pliocene of the
Mekran.
Dolium (Eudolium) fasciatum (Bruguière).
Balasore Bay, four specimens, including the type of Doliwm varicosum, Preston.
Ceylon, two specimens.
Hongkong, two specimens.
Kachh, one specimen.
Vizagapatam, one specimen.

1919. | E. W. VREDENBURG: Shells of the family Doliide. 187

Vizagapatam, one specimen (Moti Ram).

Vizagapatam, between Dolphin’s Nose and Scandal Point, 18 Jan.—17 June, 1916
(Rev. H. Hosten, S.J.), two specimens.

Locality uncertain, six specimens.

This shell also occurs at Puri.

Dolium (Eudolium) zonatum, Green.
Hongkong, three specimens, one of which is a very large shell with a super-
numerary varix.
Locality uncertain, one specimen.
Also represented by a fossil variety in the pliocene of Java.

Dolium (Malea) pomum (Linn. ).
Andamans, four specimens.
Kachh, two specimens.
Laccadives, one specimen.
Maldives, one specimen.
Mauritius, one specimen.
Singapore, one specimen.
Locality uncertain, six specimens.

Dolium (Malea) ringens (Swainson).
Panama, three specimens.

Genus Pirula (Lamarck, 1799), Sowerby, 1823.
(= Ficula, Swainson, 1840).

The Indian Museum collections contain the following species :—
Pirula reticulata, Lam. 7

» *papyratia, Say

= tessellata, Kobelt pes of P. reticulata.

7 decussata, Wood

bi dussumiert, Val. J

nn ears (bin)

investigatoris (Wood-Mason and Alcock), E. A. Smith.

The genus seems therefore to be complete as these are apparently the only well-
defined species known living.

Unlike the genus Dolium, which seems to have reached the climax of its develop-
ment at the present day and is not known in formations older than the middle
tertiary (oligocene), the greater number of fossil representatives being upper miocene
and pliocene, Pirula is known from cretaceous times, and the fossil species are more
numerous than the recent ones. A complete review of the genus such as was attempted
in the case of Dolium would belong therefore more to the province of paleontology

188 Memoirs of the Indian Museum. [Vor. VII,

than to that of modern zoology, and would include several groups without any mod-
ern representatives. In any case a general idea of the phylogeny of the genus is as
yet unattainable owing to our deficient knowledge of the earlier tertiary faunas of the
east.

It will be noticed that the majority of the living species belong to a single group,
which may be distinguished as the group of Pirula reticulata, ancestral forms of which
are known as early as the middle eocene of Europe. Pirula investigatoris belongs
to a group of which the earliest known representative, Pirula concinna, Beyrich, has
been observed in the oligocene both of Germany and of Burma; a second fossil form,
Pirula pamotanensis, Martin, occurring in the lower miocene of Java and of Kachh.
The ancestry of Pirula ficus is at present quite unknown, and is perhaps of eastern
origin.

With regard to the substitution by Swainson, in 1840, of the name Ficula instead
of Pirula under pretext of the want of homogeneousness of Lamarck’s genus, it is to
be observed that as early as 1823, Sowerby (Genera of Shells) had already cireum-
scribed Pirula within exactly the same limits as Swainson’s Ficula, giving, as the
living type, a figure of a shell which is referred to Pirula reticulata, Lamk., but which
really represents P. papyratia, Say, while P. tricarinata, Lamk. and P. burdigalensıs,
Sow. are figured as examples respectively of the earlier and later fossil forms.

The following is a list of the specimens in the Indian Museum collections :—

1. Pirula reticulata, Lamarck.

Bimlipatam (Wood-Mason), two specimens.

Hong-Kong, two specimens remarkable for their exceptionally prominent spire.
I have carefully examined them and ascertained that they are not, as might be
thought, immature specimens ot Pirula dussumieri.

Laccadive sea, Station 248 (8°37’ N., 75°37’30” E.), at a depth of 224-284 fathoms,
in sand. (M 22°). One specimen.

Malabar coast, 45 fathoms, Marine Survey, 1891-2 (No. *421), two specimens.

Maldives, two specimens.

In Lagoon of Northern Maldive Atoll (Station 148), 13-30 fathoms, in sand,
shells, and corals, one specimen.

Persian Gulf, one specimen.

Sind, one specimen.

Vizagapatam, between Dolphin’s Nose and Scandal Point (Rev. H. Hosten,
S.J.) 18-I—17-6-16. One specimen.

“ Indian seas,” three specimens of uncertain locality, No. 2588.

Locality unknown. One fine specimen, No. 2590.

The species is also known in a fossil condition from the tertiary of Karikal and
of Java; also of Europe.

2. Pirula papyratia, Say.

Florida, four specimens, No. 2597.

1919. | E. W. VREDENBURG : Shells of the family Doliide. 189

3. Pirula tessellata, Kobelt.
Australia, two beautiful specimens, No. 2509.

4. Pirula decussata, Wood.

Panama, two specimens.

5. Pirula dussumieri, Valenciennes. '

Pirula dussumieri was originally described from Chinese specimens. In the
present collection the Hong-Kong specimens are larger than those from the Bay of
Bengal.

Balasore Bay, Bengal Fisheries, M #%1?, M *?°+, two specimens.

Bay of Bengal (20°18’ N., 90°50’ E.), 65 fathoms, Marine Survey (5337, 4423),
three specimens, of which one in spirit.

Bimlipatam (J. Wood-Mason), No. 2591, two specimens.

Hong-Kong, three splendid specimens.

Persian Gulf, Station 294, Marine Survey, No. 1410 (26°33’ N., 52°23’ E.), 40
fathoms, mud and sand, one specimen in spirit.

Persian Gulf, Station 296, Marine Survey, No. 131? (26°4’ N., 56°2’ E.), 47 fathoms,
mud and sand ; one specimen in spirit.

East of Puri, Orissa Coast, Station 69, Marine Survey, 41, 812 (19°49’N.,
86°31’ E.), 46-50 fathoms, in mud ; two specimens.

Sandheads, Gangetic delta, two specimens in spirit.

The species is also known fossil from the tertiary of Java.

6. Pirula ficus, Linnæus.

Hong-Kong, two specimens.

Kachh, two specimens.

Madras, four specimens.

Negapatam, two specimens.

Orissa Coast, Bengal Fisheries, M #3*+, one specimen.

“ Puri,’ two specimens without any mark, in a box labelled ‘“ Puri,” which
originally contained nine other specimens either from other localities or of uncertain
origin. Another box also labelled “ Puri,” and with the register slip 2589, did not
contain a single specimen certainly obtained from that locality. Nevertheless the
species does occur at Puri.

1 Sacco refers to a shell, presumably identical with Pirula dussumieri, under the name of ‘* Ficula gracilis, Crosse
(Sowerby) ” (Moll. terr. terz. Piem. e Lig., Part VIII, p. 32), without quoting the authority for this correction, for which
I had hoped to find an explanation in the Journal of Malacology for 1894 (Vol. III, p. 67) which is unfortunately not
available in India. Presumably, it may be a name retrospectively proposed by Sowerby for the shell represented in
fig. 1 of the ‘‘ Genera,” where it was erroneously referred to Pirula reticulata. The figure, however, represents Pirula
papyratia, Say, named in 1822 (Journ. Philad. Acad., Vol. 11, p. 238), earlier therefore than Sowerby’s correction,
whatever may be the date of the latter. Pirula papyratia was again described as P. gracilis by Philippi in 1848 (Zeits-
shrift für Malak., Vol. V, p. 97), the repetition of Sowerby’s appellation being presumably a coincidence. In conclusion,
the appellation gracilis is liable to give rise to confusion, while the meaning of P. dussumieri is perfectly definite.

190 Memoirs of the Indian Museum. [Vor. VII, 1919.]

Singapore, one specimen.

Tavoy Coast (Museum Collector), No. 358, one specimen.

Vizagapatam, between Dolphin’s Nose and Scandal Point (Rev. H. Hosten, 8.J.),
18-I—17-6-16, one specimen.

Vizagapatam, one specimen.

Locality uncertain, one specimen marked AS, and four marked with a query; in
a box labelled “ Puri,” with the register number 2580.

The species also occurs fossil in the tertiary of the Mekran, of Ramri Island, and

of Java.

7. Pirula investigatoris (Wood-Mason & Alcock), E. A. Smith.
Ann. and Mag. Nat. Hist., (6) XIV, 1894, p. 367 ; Illustrations of the Zoology of the Investigator.
1897, pl. vi, fig. 2.
Bay of Bengal, Station 166 (13°34’55” N., 80°32’12” E.), 133 fathoms, in brown
mud, Marine Survey No. 244-75, two specimens in spirit.
Ganjam Coast, Station 96 (I8°30’N., 84°46’ E.), 98-102 fathoms, in sand,
Marine Survey, No. 281, 8161 _ 6183, two specimens, one very large. :
Laccadive Sea, Station 258 (8°23’ N., 76°28’ E.), 102 fathoms, in sand, M 788 _
170, four specimens.

EXPLANATION OF PLATE II.

. I, a, b.— Dolium fasciatum (Brug.). Balasore Bay (M #431). Original type

of Dolium varicosum. Natural size.
2, a, b.—Dolium fasciatum (Brug.). Balasore Bay (M #222). Natural size.
3, a, b, c.—Dolium fasciatum (Brug.). Varicose specimen 6f uncertain
origin ; a, b, natural size ; c, apex enlarged.

MEM. IND. MUS., VOL. VII, 1919. Prare II.

SHELLS OF DOLIUM.

S. C. Mondul, photo. Photo.-engraved & printed at the Offices of the Survey of India, Calcutta, 1919

EXPLANATION OF PLATE III.

Fic. 4, a, b.—Dolium fasciatum (Brug.). Varicose specimen from Hong-Kong.
Natural size.

» 5,4, b, c.—Dolium fasciatum (Brug.). Varicose specimen from Vizaga-
patam. Natural size.

», 6.—Doliwm zonatum, Green. Hong-Kong. Natural size.

MEM. IND. MUS., VOL. VII, 1919, Prare III.

SHELLS OF DotıuM.

S. C. Mondul, photo. Photo.-engraved & printed at the OMices of the Survey of India, Calcutta, 1919.

=
i
On

CA RE DR ee 7
wee “ss - Mr

oe.

BEREIT

EXPLANATION OF PLATE IV.

Fic. 1, a, b, ¢.—Dolium maculatum, Desh. Andamans; specimen with fully
formed aperture. Natural size.

» 2, a, b.—Dolium maculatum, Desh. Immature specimen; Andamans.
Natural size.

» 3, a, 6, e.—Dolium maculatum, Desh. Large specimen showing adult
character of ornamentation; Andamans; a, b, natural size ;
c, protoconch enlarged.

MEM. IND. MUS. VOL. VII, 1919. Puiate IV.

2a 2b

SHELLS OF DOLIUM.

S. C. Mondul, photo. Photo.-Engraved & printed at the Offices of the Survey of India, Caleutta, 1419.

EXPLANATION OF PLATE V.

Large specimen with persistent juvenile

Fie. 4.—Dolium maculatum, Desh.
Balasore Bay (Bengal Fisheries

character of the ornamentation.

M 4428). Natural size.
5.—Dolium maculatum, Desh. Specimen with adventitious supernumer-

ary main rib. (?) Ceylon. Natural size.
6.-—Dolium maculatum, Desh. Specimen with extremely depressed spire.

Sandheads. Natural size.

MEM. IND. MUS. VOL. VII, 1919. Prare V.

, oe ze
MT EL =

SHELLS or DOoLIUM.

S. C. Mondul, photo Photo.-engraved & printed at the Offices of the Survey of India, Calcutta, 1919.

EXPLANATION OF PLATE VI.

Fie. 7, a, b, c.—Dolium tessellatum, Brug. Large specimen showing adult charac-
ters. Andamans. Natural size.

MEM. IND. MUS, VOL. VII, 1919. PA VAT:

SHELLS OF Dorıum.

S. C. Mondul, photo Photo.-engraved & printed at the Offices of the Survey of India, Calcutta, 1919

. EXPLANATION OF PLATE VII.

FıG. 8, a, b.—Dolium tessellatum, Brug. Andamans. Natural size.
» 9, a, b.—Dolium tessellatum, Brug. Small specimen. Andamans. Natural
Size:
„ 10, a, b, c, d.—Dolium tessellatum, Brug. South-Andaman; a, b, natural
size; c, protoconch enlarged; d, columella enlarged to
show the columellar rugosities.

MEM. IND. MUS., VOL. VII, 1919. Pratre VII.

SHELLS OF Do.ruM.

S. €. Mondul, photo. Photo.-engraved & printed at the Offices of the Survey of India, Calcutta, 1919,

EXPLANATION OF PLATE VIII.

Fig. 11, a, b.—Dolium tessellatum, Brug. Immature specimen. Andamans.
Natural size.
» 12, a, b.—Dolium tessellatum, Brug. Immature specimen of uncertain origin,
with fully developed outer lip. Natural size.
» 13, a, b, c.—Dolium tessellatum, Brug. Specimen of uncertain origin, with
exceptionally depressed spire. Natural size.

MEM. IND. MUS. VOL. VII, 1919: IPs) WIQUE.

SHELLS OF DOLIUM.

S, €. Mondul, photo, Photo.-engraved & printed at the Offices of the Survey of India, Calcutta, 1919.

MOIRS ©

Vol. VII, No. 3.

on COLLECTION OF OLIGOCHAETA FROM THE LESSER
_ KNOWN PARTS OF INDIA AND FROM
EASTERN PERSIA.

| BY
J. STEPHENSON, D.Sc., Li.-Col., I.M.S.,
Professor of Zoology, Government College, Lahore.

% Calcutta :

PUBLISHED BY THE DIRECTOR, ZOOLOGICAL SURVEY OF INDIA.
. PRINTED AT THE BAPTIST MISSION PRESS,

; —

APRIL, 1920.

Price F7 Rupees fight Aumas.

ur RE Ze

onisn Lacty
f “fg,

F gi \
( NOV 29 1920 |

V,
Yong Must

ON A COLLECTION OF OLIGOCHAETA FROM THE LESSER
KNOWN PARTS OF INDIA AND FROM EASTERN PERSIA.

By J. STEPHENSON, D.Sc., Lt.-Col., I.M.S., Professor of Zoology, Government
College, Lahore.

(With Plates IX—X].)

CONTENTS.
| Page
Introduction 7 ae oo 7e ae 2 7 193
Fam. Naididae.
Genus Chaetogaster.
Chaetogaster bengalensis, Annand. bp 2 #3 a 195
Chaetogaster spongillae, Annand. = > ae ss AOD
Chaetogaster punjabensis, Stephenson Ar Ag is > 100
Genus Nois.
Nois communis, Piguet var, punjubensis, Stephenson .. > 2: BLY
Neis paraguayensis, Mchlisn. .. oe er > 20 197
var. aequalis, var. nov. 2 Be We eae ak OT,
Nais pectinata, Stephenson #2 ee Ae By At EOS
Nais gwuliorensis, sp. nov. mi u. RE Le fs) OS
Genus Pristina.
Pristina longiseta, Ehrbg. 2. 2 48 Ct -- 199
Genus Stylaria.
Stylaria lacustris (L.) iy “ie In er 2 0208
Genus Branchiodrilus.
Branchiodrilus sp. .. ot “ie 2 er

Fam. Tubificidae.
Genus Branchiuro.
Branchiura sowerbyi, Bedd. 42 a a 7 22 1200
Fam. Moniligastridae.
Genus Drowidu.
Drawida barwelli (Bedd.) var. impertusus, var. nov. .. À N,
Fam. Megascolecidae.
Subfam. Megascolecinae.
Genus Pontodrilus.

Pontodrilus sp. nr + ie We Me ae Wace
Genus Megascolidzs.

Megascolidzs prashad, sp.nov... ay A 2S a 20
Genus Perionyz.

Perionys sansiboricus, Mchlsn. .. 2 mE ne a:

Perionyz mıllardi, Stephenson ,. > ae vi eg

Memoirs of the Indian Museum. |Vor. VII,

Page
Perionyx rimalus, sp. nov. e aye 30 5% 20206
Perionyx pokhrianus, sp. nov. .. Oe A as 222208
var. affinis, var. nov. ve as do a LO)
Perionyx alatus, sp. nov. de oe ee = 00. a2
Perionyx shillongensis, sp. nov. Ce se DIE de Alle
Perionyx fossus, sp. nov. de 50 ae 5% a alle:
Perionyx turaensis, sp. nov. .. ac oc or ie
Perionyx pullus, sp. nov. = se By ee eee pile
Perionyx minimus, sp. nov. De De a DR Soe al)
Perionyx igatpuriensis, sp. nov. se uP Bs 22)
Perionyx spp. ae Sc on 30 ae li
Genus Lampito.
Lampito mauritii, Kinberg ar ap ae Bh 222
Genus Pheretima.
Pheretima posthuma (L. Vaill.) 5 ae She 5h
Pheretima hawayana (Rosa) .. 5 La “i 00222
Pheretima heterochaeta (Mchlsn.) a Ei Fe TE Mn
Pheretima elongata (E. Perrier) x ae BO 22
Pheretima lianicola, Stephenson 3a Be 56 0023
Subfam. Octochaetinae.
Genus Hoplochaetella.
Hoplochaetella anomala, sp. nov. or 8 fe 0.0228
Hoplochaetella spp. i ae a ah Bale SAAT)
Genus Octochaetus.
Octochaetus barkudensis, Stephenson 5% Re ds 228
Octochaetus fermori, Mchlsn. Sr Sa aie me Se 228
Octochaetus paliensis, Sp. nov. .. AR 7: + 225
var. riparius, Var. nov. De da se RONA SIL
Octochaetus prashadi, sp. nov. .. 2 De 0 23
Octochaetus montanus, sp. nov. a oe ae .. 234
Octochaetus pallidus, sp. nov. Be ce a 002236
Octochaetus ganeshae, sp. nov. .. x: Ho we "0258
Octochaetus pachpaharensis, sp. nov. 50 Le se zoo
Genus Eutyphoeus.
Eutyphoeus incommodus (Bedd.) she ite KO .. 240
Eutyphoeus mohammedi, Stephenson cbr + So un (ood
Eutyphoeus chittagongianus, Mehlsn. ae oS of a al
Eutyphoeus waltoni, Mehlsn. .. oF se de .. 243
Eutyphoeus turaensis, sp. nov. ae Be ave .. 1244
Subfam. Trigastrinae.
Genus Hudichogaster.
Eudichogaster ashworthi, Mchlsn. on er ae wo) 246
Eudichogaster bengalensis, Mehlsn. A de 38 1248
Eudichogaster trichochaetus, sp. nov. 50 50 so Rs UW249
Eudichogaster prashadi, sp. nov. 7. ar yon 7450)
Eudichogaster falcifer, sp. nov. .. EN sk of 252
Eudichogaster pusillus, sp. nov. 5 Bie 56 .. 253

Eudichogaster kinneari, sp. nov. a ae 5% 2026255

1920.] J. STEPHENSON : Oligochaeta from India and E. Persia. 193

Page

Genus Dichogaster.
Dichogaster bolaui (Mchlsn.) .. we + de fe) ABT
var. malabaricus. var. nov, x ar oll
Dichogaster affinis (Mchlsn.) +: a os a: 258
Dichogaster crawi, Eisen os Fhe x ed 30. 28

Subfam. Ocnerodrilinae.
Genus Ocnerodrilus.
Ocnerodrilus (Ocnerodrilus) occidentalis, Eisen SE ee OS
Fam. Glossoscolecidae.
Subfam. Glossoscolecinae.
Genus Pontoscolex.
Pontoscolex corethrurus (Fr. Mill.) Se ne = 73258
Subfam. Microchaetinae.
Genus Glyphidrilus.
Glyphidrilus papillatus (Rosa) .. u ie se Sg ZB
Fam, Lumbricidae.
Genus Helodrilus.

Helodrilus caliginosus (Sav.) var. trapezoides (Ant. Dug.) a ZA

Helodrilus parvus (Eisen) an A iy, PS 7260

References to Literature ae wit me 2 Eg 220260
INTRODUCTION.

The following fairly extensive investigation is based on material from a number
of sources :—

(i) By far the largest portion consists of a collection made by Dr. Baini Pra-
shad in Central and Western India in June and July, 1917. These were
the regions of India about whose earthworm fauna we knew least; and
Dr. Prashad very kindly at my suggestion gave up a month of his vaca-
tion to making the collection. My best thanks are due to him for the
expenditure of so much time, and for the large amount of labour in-
volved in visiting so many different localities. The other collections
are much smaller, and comprise—

(ii) One from Seistan and certain localities in the North-West Frontier Pro-
vince and the Punjab, made by Dr. Annandale and Mr. 8. W. Kemp
in the course of their mollusc survey in Nov., 1918—Jan., 1919 ; this
consists largely of fresh-water worms.

(iii) One from the Western Ghats, also of small fresh-water worms, by Dr.
Annandale in March, 1918.

(iv) One from Rajputana in 1918, by the Agent to the Governor-General, Col.
J. Manners-Smith.

(v) A number of tubes brought back by Dr. Baini Prashad from the Natural
History Society of Bombay.

(vi) A small collection from the Garo Hills in Assam, by Mr. Kemp, in July-
August, 1917.

194 Memoirs of the Indian Museum. [Voz. VII,

(vii) One from the Darjiling District in the Eastern Himalayas, by Drs. Annan-
dale and Gravely in October, 1917.
(viii) A few specimens from Shillong in Assam, collected by Dr. Annandale in
April, 1918.
(ix) A few specimens sent at various times from Lucknow by Mr. G. 8. Thapar.

The collections, including the types of new species, are now in the possession of
the Indian Museum, with the exception of a few tubes which I am retaining for histo-
logical work in the future.

A considerable number of the species encountered are new, as was to be expected
from the fact that the territory explored was mostly new or little worked. I have
also met with a number of species previously described by other workers from single
or ill-preserved specimens, and have sometimes been able to improve our knowledge
of them. It can scarcely be said however that the results of the present investiga-
tion include anything of the first order of importance,—it is now too late to expect
this. There are notable additions to the genera Perionyx, Octochaetus, and Hudicho-
gaster. Perionyx must now be held to have a definite territory of its own in West-
ern India, in addition to its head-quarters in the E. Himalayas ; Octochaetus is to be
recognized as an endemic and dominant genus in West and Central India; the limits
of the Hudichogaster territory however remain where they were established by
Michaelsen in 1909 (3). Hudichogaster is the characteristic earthworm of Central
India. New species of Perionyx will still be brought from the Himalayas; new
species of Megascolex from the South, and of Hudichogaster from Central India; but
the main features of the Indian earthworm fauna are now fairly well defined.

Perionyx sansibaricus turns out to be one of the common earthworms of West-
ern India, whence doubtless it was transierred to Zanzibar, where it was first found.

The new Hoplochaetella raises some interesting points of morphology and phylo-
geny, and helps to show,—what is illustrated by other parts of the paper also, and
indeed, I suppose, by the experience of systematists in general,—that the smaller our
material, the more precise and satisfactory is our systematic work. Here as else-
where increase of knowledge brings sorrow and trouble, and where before we walked
confidently as in the daylight, we hesitate and feel befogged.

I have previously had several opportunities.—more, perhaps, than have fallen to
the lot of other workers,—of examining the curious gilled Branchiura sowerbyi, and
have been interested in meeting it once again. this time from Lucknow. Though
several workers (Beddard, Michaelsen, and myself) had previously sectioned the
animal, the possibility of the protrusion of the ectal portion of the atrium as a rela-
tively long penis was not suspected until recently. Several of the specimens trom
Lucknow had both penes protruded and visible on the surface.

A new Megascolides from the Western Ghats, and a previously imperfectly
known Glyphidrilus now found at Lucknow, are interesting in view of the rarity of
these genera in India. A Drawida, of a distinct variety, is one of the common
worms of Bombay City and neighbourhood ; the genus is otherwise almost confined
to the South and to the E. Himalayas.

1920. | J. STEPHENSON : Oligochaeta from India and E. Persia. 195

Fam. NAIDIDAE.
Genus Chaetogaster.
Chaetogaster bengalensis, Annand.

Five miles S.E. of Nasratabad, Seistan, E. Persia. Small pool in the desert; water fresh but
turbid, bottom muddy with a fairly rich growth of Potamogeton and reeds, mostly in a dying
condition. The specimens were on Limnaea gedrosiana var. rectilabrum, Annand. and Prashad,
26-xii-1918. N. Annandale and S. W. Kemp.

Peshawar, N.-W.F.P.; on Limnaea acuminata. 12-i-1919. N. Annandale. Several specimens.

Kalpani stream, near Nowshera, N.-W.F.P.; on Limnaea acuminata. 13-i-1919. N. Annan-
dale. Four specimens.

Madhopur, Gurdaspur District, Punjab; on Limnaea acuminata. 27-1-1919. N. Annandale.
Forty-nine specimens taken out of the mantle cavity of a single individual of Limnaea.

Satara Fort, W. Ghats: ca. 3,300 ft. 4-iü-1918. N. Annandale. On Limnaea chlamys,
Benson.

The specimens from the Western Ghats and from Seistan were examined in some
detail.

There is sometimes, in preserved specimens, a constriction between the pharyn-
geal region and the rest of the body. The length of the setae of segment ii is com-
monly 1204, of the rest 604 In all the batches, 15 and 16 setae were quite com-
monly found in a single bundle.

The appearance of the “crop” is rather characteristic, owing to its complete
investment of chloragogen cells. These have the arrangement of paving stones, lying
side by side, and well demarcated from each other by linear intervals. The entrance
to the crop is marked, as in the specimens from the Inlé Lake (14), by a ring of tall
cells which project into the lumen.

The lateral commissural vessels in the oesophageal region are never swollen or
heart-like. The dorsal vessel, on the other hand, is often dilated just behind the
origin of these commissures.

The species appears to have a wide distribution, and to be the prevailing com-
mensal of Limnaea. It thus takes the place in India of C. limnaer, which is found
in a similar association in Europe; Michaelsen has however found C. limnaei on a
specimen of Limnaea from the Kumaon District (Central Himalayas) (3). Last year I
identified several specimens from a sponge (Hphydatia fluviatilis) from the Inlé Lake as
doubtfully belonging to C. limnaei (14) ; through the kindness of Dr. Annandale I have
recently had the advantage of comparing my examples of this form with a specimen
of C. hmnae sent to the Indian Museum by Dr. J. H. Ashworth of Edinburgh ;
but I am still unable to say definitely that the Inlé worms either are or are not
C. limnaei. The identification of these small worms from preserved material only
is, as I have previously explained (12), often both difficult and extremely hazardous.

Chaetogaster spongillae, Annand.
Khandala, W. Ghats; from sponge (Spongilla crateriformis) in artificial tanks made by dam-
ming stream; bottom,—mud over rocks; some stones; weeds fairly abundant. 6-iii-1918
N. Annandale.

196 Memoirs of the Indian Museum. Vor. VI,

The length of a chain of two individuals was from -47 to ‘7 mm., the diameter
‘15 mm. The head is relatively small in this species, and, as has previously been
described, the upper lip projects forwards in front of the mouth, forming a short
prostomium. ‘There is a slight constriction between the pharyngeal region and the
rest of the body. The number of setae in the first bundle was five or six, in those
that follow four, and towards the hinder end three or fewer. I could not detect any
difference in the thickness of the terminal prongs (in a previous description I have
noted the distal prong as being the thinner); the length and other proportions are
the same as those I formerly gave (7). Chloragogen cells are absent on the crop.
n = 8, as before.

Chaetogaster punjabensis, Stephenson (?).
Nasratabad, Seistan, EK. Persia; water-channel in Consulate Garden. Nov. and Dec. 1918.
N. Annandale and S. W. Kemp.

A single specimen was found among a number of examples of Nais communis var.
punjabensis in association with colonies of the Polyzoon Lophopodella (see below).
The association was no doubt quite fortuitous.

Here again it is impossible to speak with certainty ; the small size, and distinct
and relatively long oesophagus of the specimen, are characteristics of C. punjabensis.
The numbers of setae per bundle appear to be rather smaller here, and I could not
follow the dorsal and ventral vessels forwards beyond the anterior end of the crop in
a glycerine mount of the specimen; I also failed to distinguish any refractile body
in the cerebral ganglion, though this is a notable feature, at least of living speci-
mens.

The specimen was in a late stage of sexual maturity ; the clitellum was present,
and there was a mass of ova in the middle of the body ; the animal was much swol-
len in this middle region. In the Punjab the Naid worms mostly become sexual at
the beginning of the hot weather,—April and May; but this specimen was taken in
Seistan in the early part of the cold weather.

Genus Nais.
Nais communis, Piguet var. punjabensis, Stephenson.

Nasratabad, Seistan; water-channel in Consulate Garden; water fresh but turbid, bottom
muddy with a scanty growth of weeds. Nov. and Dec. 1918. N. Annandale and 8. W. Kemp.
Numerous specimens.

Open pool in the reed-beds of the Hamun-i-Helmand, a few miles east of Lab-i-Baring, Seistan.
Water very slightly brackish, fairly clear, about five feet deep ; bottom muddy with a luxuriant
growth ot Potamogeton pectinatus. 8-xii-1918. N. Annandale and S. W. Kemp. Three specimens.

Peshawar, N.-W.F.P.; on Limnaea acuminata. 12-i-1919. N. Annandale. A single speci-
men.

Khandalla, W. Ghats; in algae on cliff kept wet by spray of a small waterfall.! 7-9-ii-1918.
N. Annandale. Two tubes, five specimens in- one, four in the other.

1 N. Annandale, Rec. Ind. Mus. XVI, p. 121 (1919).

1920. ] J. STEPHENSON : Oligochaeta from India and E. Persia. 197

The specimen from Peshawar occurred along with Chaetogaster bengalensis ; its
presence in this association was probably accidental.

Dr. Annandale’s note on the worms from Nasratabad runs :—“ Oligochaetes in
relatively long mucilaginous tubes intertwined with stems of weed. A colony of the
Polyzoon Lophopodella attached to each tube.” Of the two glass tubes in which the
Nasratabad specimens were sent to me, one contained worms only, the other some
fragments of weed, and several colonies of Lophopodella, each attached to a soft brown-
ish tube. I found the worms on the fragments of weed, but there were none still
remaining in the tubes to which the Polyzoon colonies adhered. I have found this
worm in numbers in tubes in Lahore (5), but the tubes in this case were appa-
rently those of insect larvae, not manufactured by the worms themselves.

The specimens from the Western Ghats are possibly a separate variety. The
dorsal needles are in the var. punjabensis finely forked ; though barely or not at all
distinguishable with the ordinary high power, the forking is quite evident on exam-
ination with the oil immersion, when the needles lie in a favourable position. In
these specimens I thought I detected a trace of bifurcation in a few cases on close
observation, but in many the forking seemed quite definitely to be absent.

Nais paraguayensis, Mchlsn.
Plate IX, fig. 1.
Gwalior, Central India; in a pond, attached to Hydrilla and other débris; 18-vi-1917. B.

Prashad. Three specimens.

This species has previously been recorded from Calcutta and from Sirsiah in
Bihar by Michaelsen, and from Lahore by me. The species seems to be rather vari-
able. The present specimens were from 4:5 to 7°5 mm. in length, and consisted of
from 29 to 56 segments, without any sign of a budding zone. The ventral setae
are three or four per bundle; in the body generally the prongs are equal in length,
but the outer is only two-thirds or even half the thickness of the inner. In the first
four seta-bearing segments both prongs seem to be longer and thinner than in more
posterior segments, but the relative thicknesses are maintained; the outer prong is
slightly longer than the inner; the shaft is also slightly thinner than in succeeding
segments.

The dorsal bundles consist usually of one hair and one needle; two needles may
occur, and also two hairs, in which case one is much shorter than the other. There
are slight variations from the typical form among the needles; in one case the smaller
outer prong was itself bifid; in another the longer prong was slightly bent outwards
towards the smaller; in one specimen the outer prong was regularly very short (fig. T,
a and b).

var. aequalis, var. nov.
Plate IX, fig. 2.

Saugor, Central Provinces; in a large lake, attached to leaves. 20-vi-1917. B. Prashad. A
single specimen, in spirit, and one preserved and flattened in glycerine on a slide at the time of
capture,

198 Memoirs of the Indian Museum. [Vor. VII,

The spirit specimen is 3°5 mm. in length, and ‘23 mm. in thickness; it has 34
segments, with a short region posteriorly in which segments are not yet differenti-
ated. There is no budding zone.

The prostomium is moderately large and long; its length is equal to its breadth at
the base, and it is rounded anteriorly. There are no eyes. The anus is dorsal.

The dorsal setal bundles begin in segment vi. They consist of one hair and
one needle seta, — never more than one of either. The hairs have approximately a
length equal to the diameter of the body. The needles are slightly sickle-shaped
(fig. 2), and forked distally; when the seta is in a good position for observation this
can be seen with the ordinary high power. The prongs are of the same length (the
outer may possibly be the least trifle longer), and join at an acute angle; the outer
seems to be slightly thinner. In length these needles are 52,, in thickness 2°5,.

The ventral setae are of the usual type, and are usually 4 per bundle; 3 and 5
were also met with. In length they are 52+, in thickness 2°54. The inner prong
is of equal length with the outer, but is twice as thick. I cannot see any difference
of type between the setae of segments ii—v and the rest; but unfortunately the one
specimen is fixed in such a position that these setae can be seen well from neither
side, and the other (the already mounted specimen) seems to be the just separated
posterior animal of a chain, in which the anterior segments have not yet fully
developed. |

There is no stomach. Coelomic corpuscles are present. The cerebral ganglion
is bifid both anteriorly and posteriorly, and consists of two more or less independent
halves, contiguous for some distance along their inner borders.

In the typical form of N. paraguayensis the outer prong of the dorsal needles is
considerably shorter than the inner, while in these specimens itis of equallength. As
however there appear to be no other essential differences, it will, I think, be sufficient
to describe them as a variety.

Nais pectinata, Stephenson.
Gwalior, Central India; in a pond, attached to Hydrilla and other debris. 18-vi-1917.

B. Prashad. A single specimen.

The specimen agrees generally with those previously described from Bheemanagar,
Travancore (6). Since the presence or absence of a stomachal dilatation is one of the
few internal marks that are used as specific characters in this genus, I may supple-
ment the original description by adding that there is none in this species.

Nais gwaliorensis, sp. nov.
Plate IX, figs. 3, 4.
Gwalior, Central India; in a pond, attached to Hydrilla and other débris. 18-vi-1917.
B. Prashad. A single specimen.

The specimen is in length 2°7 mm., in thickness ‘25 mm. There are 29 segments
with a small zone behind where segments are not yet differentiated. There is no zone
of budding. The prostomium is bluntly triangular, its length being equal to its base.
Eyes are absent. There is also no stomachal dilatation on the alimentary canal,

1920.] J. STEPHENSON : Oligochaeta from India and E. Persia. 199

The dorsal setae begin in segment vi, and each bundle consists usually of one
hair and one needle seta; two hairs are sometimes found, in which case one is shorter
and thinner than the other; once two hairs and two needles were seen,—the one
couple may perhaps have been destined to replace the other when they fell out.

The hair is as a rule not quite equal in length to the diameter of the body. The
needles are about 45” in length (hardly any can be measured accurately, owing to their
not lying flat), and bent at a very obtuse angle at a point rather distal to the middle
(fig. 3). The distal section of the seta is slightly curved in the contrary direction,—
very slightly only, so that the whole can hardly be described as sickle-shaped. There
is an indefinite nodulus at the angle in the shaft; the length of the distal to that of
the proximal section of the shaft is about as 2 to 3. The tip is bifid, the two prongs
being visible to the ordinary high power of the microscope; the angle between the
two prongs is moderately wide, and the outer, which continues the direction of the
shaft, is slightly longer and perhaps slightly thinner than the mner.

The ventral bundles, in all segments from vi onwards, consist of four or five setae,
45—53" in length and 2:54 in thickness (fig. 4a). The nodulus is distal; its exact
position on the shaft probably varies in the several setae of a bundle (cf. Stephenson,
10), though I was not able to obtain exact measurements. The prongs are equal in
length, the outer is not swollen at the base, and is only half or two-thirds as thick
as the inner.

In the first four seta-bearing segments the form differs somewhat. The shafts
of the setae are thinner and straighter; the nodulus is about the middle or a little proxi-
mal to the middle of the shaft; the outer prong is 14 times as long as the inner, two-
thirds as thick at the base, and more hooked (fig. 4b). There are four setae per
bundle; their length is 50 to 56#, and their thickness only 2-.

Remarks:—The species to which the present comes nearest are N. tenwidentis
(Walton, 15) and N. raviensis (Stephenson, 9). The distinguishing character of the
former is the very long and slender prongs of the ventral setae,—hence-its specific
name. The separation of the present form from N. raviensis depends on the characters
of the setae, both dorsal and ventral; the differences will be best realized by compar-
ing the figures given in the present paper with text-fig. 1 of my description of N.
raviensis. The most obvious are the position of the bend of the shaft of the dorsal
needles (much nearer the middle here), the relative lengths of anterior and posterior
ventral setae (the anterior are nearly twice as long as the posterior in N. raviensis), and
the characters of the prongs of the anterior ventral setae (in N. raviensis the outer is
very much the longer, and makes a very narrow angle with the inner).

Genus Pristina.

Pristina longiseta, Ehrbg.

Gwalior, Central India; in a pond, attached to Hydrilia and other débris. 18-vi-1917.
B. Prashad. One complete specimen, and perhaps one or two more in which the characteristic
proboscis or long setae were damaged.

The toothing of thehair setae was only just visible with the oil immersion lens.

200 Memoirs of the Indian Museum. [Vor. VII,

Genus Stylaria.
Stylaria lacustris (L.).

Open pool in the reed-beds of the Hamun-i-Helmand, a few miles east of Lab-i-Baring, Seis-
tan. Water very slightly brackish, fairly clear, about five feet deep; bottom muddy with a
luxuriant growth of Potamogeton pectinatus. 8-xii-1918. N. Annandale and 8. W. Kemp. Several
specimens.

Fam. TUBIFICIDAE.
Genus Branchiura.
Branchiura sowerbyi, Bedd.

Lucknow, Gaumati River. 15-iii-1919. G.S. Thapar. A single specimen.

The specimen was of fair size, 50 mm. in length, and is interesting from the fact
that both penes are protruded. As I have previously noted (14), a penis was not
described or suspected to exist in this animal, until it was found in specimens from
the Inlé Lake.

I have also received, taken on the same day and from the same source, a number
of specimens of a species of Branchiodrilus, the Naid worm with gills remarkably
like those of Branchiura, but on the anterior part of the body instead of the pos-
terior. The occurrence of the two together is a point of some interest. I much
regret not to be able to give the specific diagnosis of the Branchiodrilus, of which
three species are known, all Indian; but the worms came to hand when the present
paper was already completed, and I fear it will be some time before I have the
opportunity of undertaking detailed investigations.

Fam. MONILIGASTRIDAE.
Genus Drawida.
Drawida barwelli (Bedd.) var. impertusus, var. nov.

Bombay, Victoria Gardens. 30-vi-1917. B. Prashad. Five specimens.

Bombay, under a tree near the Fort. 30-vi-1917. B. Prashad. Seven specimens.

Elephanta Island, Bombay; high up on a hill. 30-vi-1917. B. Prashad. Three specimens.

Elephanta Island, Bombay; in a rotten tree. 30-vi-1917. B. Prashad. A single specimen.

Elephanta Island, Bombay ; on the sea-shore. 30-vi-1917. B. Prashad. A single specimen,
immature.

External Characters :—The length of fair-sized specimens is from 45 to 48 mm.,
and their diameter 3°55 mm. The colour is a rather blotchy olive, darker on the
dorsal surface than ventrally, with a still darker mid-dorsal line ; the first few seg-
ments are pale. The number of segments in two specimens was 130 and 132.

The prostomium is small, prolobous, and under cover of the first segment.

Dorsal pores are absent. .

The setae are small and closely paired, and are visible as far forward as the
second segment. The interval aa is rather less than be (# or 2bc), or may be fully
equal to it towards the hinder end; dd is about # of the circumference.

The nephridiopores are in a single line, just below the level of the setae c.

1920. | J. STEPHENSON : Oligochaeta from India and E. Persia. 201

The clitellum extends over segments x-xiii (=4); it is not well defined, and
the segments are largely unaltered.

The male pores are very prominent in furrow IO/IL, and are situated midway
between the lines of setae b and c. They are bordered by anterior and posterior lips,
and it is these lips, rather than the apertures themselves, which are the conspicuous
features; the extent of the lips is slightly variable,—from a point about in line with
the ventral pair of setae nearly to the level of the lateral pair.

On segment x in front of the male apertures are a pair of whitish papillae with
indefinite margins; the centre is whiter than the rest, and the appearance is that of
some solid organ shining through. Their exact position is variable; they may even
be near the middle line, internal to the line of the ventral setae.

The female pores are in groove II/I2, in line with setae b.

The spermathecal pores are in 7/8, immediately below the line c.

Internal Anatomy :—Septa 5/6 to 8/9 are moderately strengthened ; the rest are thin.

There are four gizzards, in segments xiv to xvii. In the second specimen dis-
sected, that in xvii was notably smaller than the others; and in xiii there were numer-
ous longitudinal shining muscular bundles on the oesophagus, forming a rudimentary
gizzard here also.

The last heart is in segment ix.

The testis sacs vary in shape ; in the first specimen dissected they were rather kidney-
shaped, with the hilus directed downwards and outwards, and the anterior ends rather
narrower than the posterior ; they were suspended by septum 9/10, almost by their middle,
the posterior portion in x being rather larger than the anterior in ix; in the second,
they were rounded smooth masses, unconstricted, mostly (practically altogether on
the right side) in segment x. When opened, the contents were with difficulty evacu-
ated, and even then only in part; a large portion of the inner surface of the sac
appeared to be proliferating the sexual cells, v.e. the testis is diffuse; a firmer though
not iridescent mass on the floor of the sac, just over the site of origin of the vas deferens,
appeared to indicate the position of the funnel.

The vas deferens is either considerably or not much coiled; passing downwards
from the under surface of the sac it runs part of its course in segment ix, and then
enters the anterior border of the prostate in x.

The prostate is of moderate size only, flattish, sessile on the body-wall, its trans-
verse rather greater than its longitudinal axis; its surface is quite soft and furry
(‘glandular ”).

Segment xi constitutes a perfectly closed annular ovarian chamber. The ovisacs
pass backwards from the hinder wall of the chamber through segments xii and xii,
and may get into xiv; their margins may have a crenulated appearance or not.

The spermathecae are situated in segment viii. The ampulla is globular or broadly
ovoid, dorsally situated in the segment, and connected by a band with the one of the
other side. The duct is much convoluted as it passes down on the posterior face of
septum 7/8. There is no diverticulum or atrial sac, not even in the body-wall, though
the duct is slightly thickened at its termination.

202 Memoirs of the Indian Museum. [Vou. VII,

Remarks :—The two species of Drawida to which the present specimens bear most
resemblance are D. bourne: (Mchlsn.) and D. barwelli (Bedd.). From the first the pre-
sent form differs (besides a few minor details) in the shape of the testis sacs (not a
great matter, since this is variable in these specimens), in its much smaller size, and
especially in the fact that the surface of the prostates is “glandular.” From the
second, it differs in a few details such as the number and situation of the gizzards
(points which are however very variable), the relative magnitude of the setal inter-
vals, and the shape of the prostates (pear-shaped in D. barwelli, almost circular, flat
and sessile in the present form); the chief difference however is the absence of dorsal
pores here,—a character which the varietal name is intended to indicate. Most of
the species of Drawida have no dorsal pores; the type form of D. barwelli, however, is
peculiar in possessing them. I have rejected the alliance of the present form with
D. bourne: (itself, according to Michaelsen, 4, a variety of D. pellucidus), because the
smooth and shiny, or soft and furry, condition of the surface of the prostate is appa-
rently a distinction of some importance (cf. Michaelsen, in the paper just quoted).

The diagnosis of the present variety may run as follows: Drawida barwell
var. 2mpertusus :—as for the typical form, with the following exceptions. Setal in-
terval aa rather less than bc, except at the hinder end. Male pores bounded by very
prominent anterior and posterior lips; a pair of indefinite but fairly large whitish
papillae on the segment in front of the male pores. No dorsal pores. Four gizzards,
in xiv—xvil. Prostates flat, sessile, almost circular.

Fam. MEGASCOLECIDAE.
Subfam. MEGASCOLECIN AE.
Genus Pontodrilus.

Elephanta Island, Bombay; on the sea-shore. 30-vi-1917. B.Prashad. A single specimen,
not fully mature.

The species was presumably the one which has been found on several parts of
the coast of India, P. bermudensis, Bedd. f. ephippiger (Rosa).

Genus Megascolides.
Megascolides prashadi, sp. nov.
Plate IX, figs. 5, 6.

Sakarwari, on the way to Mahableshwar, W. India 4-vii-1917. B. Prashad. A single
specimen.

External Characters :—Length 42 mm., diameter 4mm. The worm is unpigment-
ed, of a buff colour, which is lighter at the ends of the body and in the clitellar region.
Segments 140; the last 60 however are very short and lighter in colour, and have
perhaps been regenerated. -

The prostomium is prolobous.

The dorsal pores begin at the anterior border of the clitellum, in groove 12/13.

The setae are paired. In the middle of the body the relative size of the intervals
may be expressed by the formula ab = 2aa = 2bc = cd; behind the clitellum

1920. | J. STEPHENSON : Oligochaeta from India and E. Persia. 203

this becomes ab = 2aa = }be = #cd; and in front of the clitellum ab = faa =
2b¢ — 3cd. The mid-dorsal interval dd is in the middle of the body equal to
about half the circumference.

The clitellum is smooth, thickened, well-defined at each end, and extends over
segments xlli—xvli ( = 5).

The male pores are on segment Xviil, just outside the line of setae b. They are
small, and surrounded by only a slight whitish thickening.

On segment xix is a large flat oval papilla (fig. 5); this is not quite symmetric-
ally placed, being rather on the left side, so that while it reaches as far outwards as
the male pore on the left side, it stops somewhat short of this on the right; its centre
shows a transverse, almost groove-like, depression. On segment xx is a second
papilla, much smaller and less definite than the last, transversely elongated, with its
centre about in line with the setae a, extending inwards to about the middle line;
like the last, it is situated on the left side.

Segment xvii is delimited in front by a groove mid-ventrally, but not elsewhere,
since this segment forms part of the clitellum; on this ventral portion are seen a few
small whitish circular spots, which however do not seem to have anything to do with
setae. Setae a and b are absent on segments xviii and xix, and on the left side
on XX.

The female pore seems to be represented by a small white dot mid-ventrally
placed on xiv.

The spermathecal pores are a single pair, in groove 7/8, in or immediately
outside the line of setae b.

On the hinder border of segment viii are situated a pair of indefinite, transversely
oval papillae, in position and size corresponding to the setal interval ab.

Internal Anatomy :—Septum 4/5 is slightly thickened as compared with those in
the middle of the body; 5/6 and all the following septa down to 10/11 are moderately
thickened ; 11/12 again is only slightly strengthened.

The gizzard, in segment v, is subspherical and of moderate size. There are no
calcareous glands. The intestine begins in xv, or perhaps in xvi.

The last heart is in segment xii.

Behind the clitellum the micronephridia are arranged in transverse rows of about
eight to ten on each side; in the clitellar region they are also in transverse rows,
and somewhat larger; in front of this they are sparser, and their arrangement is
less regular. About forty segments from the hinder end the innermost nephridium
on each side in each segment enlarges, and this condition is maintained to the end;
there is thus a longitudinal row of larger nephridia on each side of the ventral nerve
cord, but I do not think that these could be described by anyone as meganephridia,
—only as enlarged micronephridia.

Testes and funnels are free in segments x and xi; the funnels were inferred from
the iridescent masses which probably enclose them; the testes were separately
identifiable in xi, while in x they were presumably continuous with a deeply attached
mass of flocculent matter (developing sperm-morulae and spermatozoa).

204 Memoirs of the Indian Museum. — [Vor. VII,

Three pairs of seminal vesicles are present. The largest are those in xii, of
moderate size and lobulated ; in segment ix is a second pair, rather smaller and also
lobulated; the third pair, in x, are smaller still, and attached to septum Io/II.

The prostates are tubular, and consist of a number of thick, adpressed opaque
coils which extend through several segments. The duct is relatively short, proceeds
almost straight inwards, and is narrow but broadens slightly towards its ectal end.

Ovaries and funnels have the usual situation.

The spermathecae are one pair. The ampulla is a large irregular sac, with much
crenulated margins (fig. 6); the duct is about as long as the ampulla, of moderate
thickness and approximately the same diameter throughout. There is a single diver-
ticulum, originating at the ental end of the duct, lobulated, about half as long as the
duct, to the side of which it is adherent.

There are no penial setae.

Genus Perionyx:
Perionyx sansibaricus, Mchlsn.
Plate IX, fig. 7.

Manmad, Bombay Pres. 28-vi-1917. B. Prashad. Numerous specimens.

Igatpuri, Bombay Pres. 29-vi-1917. B. Prashad. Two specimens.

Khandwa, Central Provinces. 23-vi-1917. B. Prashad. Numerous specimens

Kala Khund, between Khandwa and Indore, Central India. 23-vi-1917. B. Prashad. Three
specimens.

Baroda, W. India, on the banks of the river Vishvamitri. 9-vii-1917. B. Prashad. A single
specimen.

Wathur, near Mahableshwar, W. India. 6-vii-1917. B. Prashad. Nine specimens.

Londa, ten miles from Castle Rock, Bombay Pres. 6-vii-1917. B. Prashad. Eleven speci-
mens.

This interesting species has been previously recorded from Kodaikanal in the
Palni Hills (Michaelsen, 3). From the present records it appears to be common in
Western India. One of its distinguishing characters is the alternation in situation of
the terminal bladders and external openings of the nephridia ; these are placed at
about one-third of the half circumference from the mid-dorsal line, and a similar dis-
tance from the mid-ventral line, in successive segments ; as the end bladders are easily
seen on opening the animal (unlike most species, where they are small or absent),
the peculiarity is obvious in any dissection. The alternation is however, according
to my dissections, approximate, not exact ; two successive nephridia not uncommonly
end at the same level on the body-wall, and once a series of four were seen to
do so.

I append a few notes on the numerous specimens with which I have had to deal.

External Characters :—The purple colour of the dorsal surface extends partly
on to the ventral side of the animal in the most anterior segments. I found this
character of help in separating worms of this species from others when several
species were mixed together in a single catch. |

The dorsal pores, which Michaelsen found to begin in groove 9/10 or 8/9 in his

1920. | J. STEPHENSON : Oligochaeta from India and E. Persia. 205

original single specimen, and in 4/5 in the examples from the Palni Hills, begin here
in 3/4, remarkably far forward.

The region of the male pores is characteristic (fig. 7). Segment xviii is somewhat
lenethened in the mid-ventral region, its anterior and posterior limits being bulged
forwards and backwards respectively. The male apertures are small transverse
slits close to the middle line; while in front and behind the pores there is a cres-
centic depression, the convexity of the depressions facing forwards and backwards
respectively, and causing the bulging, forwards and backwards, of the intersegmental
erooves; the pores are thus situated on a transverse ridge between the crescentic
depressions. A characteristic feature is that the setal ring is not interrupted by the
pores, but is continued across the segment, on the ridge but just behind the
male pores.

Internal Anatomy :—I find the gizzard to be in segment vi (Michaelsen in v) ;
it is very rudimentary. The intestine is somewhat swollen in xiii, but not specially
vascular, nor are there any ridges in the interior,

Testes and funnels are free in segments x and xi. Besides the seminal vesicles
in segments xi and xii, there was in one specimen dissected a minute structure in
ix, on the left side and attached to the anterior face of septum 9/10, consisting of
two small ovoid lobules, and perhaps representing a rudimentary vesicle. The
vesicles in xii were larger than those in xi, and were somewhat lobulated; those in xi
were markedly so. ;

The ampulla of the spermathecae is pear-shaped, narrowing towards the ectal end
to a duct; this is short, relatively narrow, cylindrical, one-third as long and one-
fourth as wide as the ampulla. There is a single diverticulum, on the inner side of
the duct, consisting of a few indistinct seminal chambers aggregated together on a
short stalk; the diverticulum is one-fourth the length of the ampulla.

Perionyx millardi, Stephenson.
Plate IX, fig. 8.

Bombay, Malabar Hill. 1-vii-1917. B. Prashad. Three specimens.

Talegaon, on the way to Poona. 2-vii-1917. B. Prashad. A single specimen.

Kalyan, a short distance inland from Bombay, Bombay Pres. 7-vii-1917. B. Prashad.
Four specimens.

Virar, N. of Bombay, Bombay Pres. 7-vii-1917. B. Prashad. A single specimen.

The present species was described by me (11) from three specimens from Malabar
Hill, Bombay ; these were however much softened and in bad condition, and a few
supplementary notes may therefore be of interest.

The lengths of the specimens which have now come to hand show rather greater
variation than those of the original batch; these are from 35 mm. (though this was
perhaps regenerated at its hinder end) to 90 mm.

The ventral break in the setal rings is very small,—about 14 ab,—but it is regu-
lar from the clitellum backwards. The dorsal break is very irregular, small or absent,
and often not in the middle line. The setae are slightly closer set, and certainly more

206 Memoirs of the Indian Museum. [Voz. VII,

regularly placed, ventrally than dorsally. The numbers counted were :—ix/40, xii/41,
xix/48, and in the middle of the body 41.

No septa are thickened.

The gizzard, in segment vi, is of some size, but its walls are thin and soft.

The seminal vesicles of segment xii are larger than those of xi, and bulge back
the posterior septum of the segment; they may even extend backwards so as to bulge
back septum 13/14. Both pairs are smooth and scarcely or not at all lobed.

The prostates may take up fully two segments,—xvili and xix, bulging septum
17/18 forwards and 19/20 backwards. The border of the gland is not cut up into
lobes, except by an indentation on the anterior margin, and by a deep notch from
which the duct emerges. The duct is quite straight, soft and only slightly shiny, and
of equal diameter throughout; it passes directly inwards.

The chief difference between these specimens and those formerly described is in
the spermathecae; all the specimens of the present series which were dissected pos-
sess a diverticulum, thus differmg from the previous examples, in which I found
none. In these, the ampulla is a large ovoid sac; the duct is much shorter than the
ampulla, from which it is distinctly marked off, narrow and of the same diameter
throughout. There is a single diverticulum, which varies in its appearance; regard-
ing the specimens from Malabar Hill, Bombay (whence the original examples of
the species also came), all that can be said is that there appears to be a small scale-
like diverticulum from the base of the ampulla; in that from Virar the diverticulum
arises from the junction of duct and ampulla, is small, rather scale-like and flattened,
and lies against the base of the ampulla (fig. 8a); in that from Talegaon it is rather
flat, sessile, somewhat cauliflower like, showing a number of small seminal chambers
(fig. 8b). In the specimens from Kalyan the whole organ is of a rather different
appearance; the ampulla is somewhat lobed, and has a fairly broad base, from which
the duct issues; the diverticulum is larger than in the previous specimens, and is
divided into three lobules lying side by side; indeed the incisions between the lobules
appear to be so deep that each lobule has its own attachment to the upper end of
the duct, i.e. there are really three diverticula (fig. 8c). Since however the whole
of the anatomy, including the penial setae, is the same in these latter specimens as in
the others, it scarcely seems allowable to separate them. The absence of any diverti-
culum in the original specimens is perhaps due to their relative immaturity.

Perionyx rimatus, sp. nov.
Plate IX, fig. o.
Jor Pokhri, 4,800 ft.. Sitong, Darjiling Dist., E. Himalayas. 22-28-x-1917. N. Annandale
and F. Gravely. Two specimens, one mutilated.

External Characters :—Length 80 mm. ; diameter 45 mm. Colour in the anterior
part of the body a light rather blotchy purple on the dorsal side, pale in the posterior
half except for a purple middorsal stripe; ventrally pale throughout.

The body is rather flattened, and the clitellum is narrowed. Segments 107.

1920.] J. STEPHENSON : Oligochaeta from India and E. Persia. 207

Prostomium epilobous +; a faint groove delimiting the prostomium behind.

Dorsal pores from 4/5.

The setal rings are often closed both dorsally and ventrally; sometimes there is
a small break, but it is irregular and varying in extent,—not more than 2ab or 2yz,
and often less. The setae are rather small,--smaller and closer set on the ventral
than on the dorsal surface. The following numbers were counted :—v/59, ix/63, xii/64,
xix/56, and in the middle of the body 56.

The clitellum extends over segments xiii-xvi = 4; it is narrowed, rather lighter
in colour, with visible dorsal pores and intersegmental grooves.

On segment xviii is a deep transverse crack with corrugated anterior and
posterior lips, situated in the middle of the length of the segment and extending
transversely over the middle half of the ventral surface. It is difficult to see the
male pores in this groove, but they appear to be near its centre, and only separated
by a slight median elevation in the floor of the groove.

The female area is a median whitish circular patch anteriorly on segment xiv.

The spermathecal apertures are two pairs, in grooves 6/7 and 7/8; they are
small, with slightly puckered lips, rather close together, about in line with seta c or
the space cd.

There are no other genital marks.

Internal Anatomy :—Septa 4/5, 5/6 and 6/7 are thin, but increase slightly in
thickness progressively; 7/8 is somewhat strengthened, and 8/9 moderately so.
Thence to the prostatic region all are slightly, but none much, strengthened,—10/11 is
perhaps least so.

The gizzard is small and rudimentary, in segment v. The intestine begins
behind the prostates.

The last heart is in segment xiii.

The nephridia end in the same line.

Testis sacs are present in segments x and xi. In x the sac is lobed, distinctly
though not deeply, and presents the appearance of a number of ovoid lobes lying
side by side in transverse series; it is continuous below the oesophagus from side
to side, encloses both oesophagus and hearts, and appears to be divided dorsally by a
median septum above the alimentary canal. In xi the sac is smaller, and lies
posterior and ventrally to the vesiculae seminales.

The seminal vesicles, in segments xi and xii, have a granular surface but are not
otherwise lobed. Each pair is fused dorsally over the alimentary canal, so as to form
a single sac in each segment; that in xi overlies the testis sac deeper in the segment,
—it is not an extension of the testis sac (as is described for P. himalayanus by Michael-
sen, 3), but has an independent attachment to the posterior face of septum 10/11.
In an earlier stage of development, exemplified by the second of the two specimens,
the sac of segment x is smaller, and does not include the alimentary canal and hearts ;
it grows up round them, it would appear, during its formation.

The prostates are large, occupying segments xviii and xix, and it may be part of
xvii also; they are deeply cut up by the septa, and also otherwise much indented.

208 . Memoirs of the Indian Museum. | [Vor. VII,

The duct is much twisted and relatively thin, not firm and shining ; its ectal end is
rather stouter than the rest.

There are two pairs of spermathecae (fig. 9). The organs occupy the whole
available space in segments vii and viii, and meet dorsally. The ampulla is a large
irregular sac, with bulgings here and there; the duct is moderately stout, half as
long as the ampulla, not firm or shining. The diverticula are in the form of a few small
warts on the duct a short way below the base of the ampulla; these form a cluster,
about half a dozen in all, of which one appears to be larger than the rest; the smaller
hardly project at all, but the iridescent spermatozoa shine through.

There are no penial setae.

Remarks :—I was at first inclined to identify this form as Perionyx himalayanus
(Michaelsen, 3), on the ground of its possessing testis sacs,—a rather unusual feature
in this genus. There is also the fact that the present specimens come from the same
district (Darjiling Dist.) as P. himalayanus, and that both are much paler in colour
than is general.in this genus. But, neglecting slighter differences such as the distri-
bution of the thickened septa, the character of the prostatic duct, the numbers of
the setae, and the extent of the dorsal pores, there remain essential differences in
the extent of the clitellum, the spermatheca! diverticula, and especially the configu-
ration of the male area. In P. himalayanus the male pores are situated on large
round papillae which are quite at the sides of the ventral aspect, one-fifth of the
circumference apart from each other; and a number of setae intervene between the
two papillae. Here the pores are situated near each other in the depth of a trans-
verse crack.

Though both species possess testis sacs, it is possible that their relations to the
seminal vesicles are not the same. In P. himalayanus the sperm sacs are apparently
prolonged laterally to form seminal vesicles in segments x and xi; here there is no
such prolongation in segment x, seminal vesicles being absent in this segment; and
the seminal vesicles of xi are independent of the testis sacs, and have their own at-
tachment to the anterior septum of the segment. —

Perionyx pokhrianus, sp. nov.
Plate IX, figs. 10, II.
Jor Pokhri, 4,800 ft., Sitong, Darjiling Dist., E. Himalayas. 22-28-x-1917. N. Annandale
and F. Gravely. A single specimen (along with the last).
External Characters :—Length 65 mm.; diameter 3 mm. Colour pale violet
dorsally,—posteriorly paler than in front; a mid-dorsal darker stripe; ventral surface
unpigmented. Segments 96.
Prostomium epilobous +, the tongue (posterior portion projecting into segment 1)
not cut off by a groove behind.
Dorsal pores begin from groove 4/5. :
The setae are in rings, which are almost closed both dorsally and ventrally, and
may be quite closed in the hinder part of the body; aa or zz = It or 13 times ab or

1920. ] J. STEPHENSON : Oligochaeta from India and E. Persia. 209

yz. The setae are set slightly closer ventrally than dorsally. The following numbers
were counted :—v/50, ix/58, xii/54, xix/48, and in the middle of the body 44.

The clitellum is narrowed, and extends over segments xiü-xvi (= 4). It is
lighter in colour than the neighbouring segments, and the intersegmental grooves and
setae are visible.

The male field occupies the middle of segment xviii. Here are seen a pair of
papillae which take up the greater part of the length of the segment (fig. 10); they
are bounded both in front and behind by a common transverse or slightly crescentic
groove, the anterior groove being the better marked, and are separated from each
other by a longitudinal groove in the middle line. On their outer margins the papil-
lae are not delimited from the surrounding area. The male pores are near the middle
line, and nearer the posterior than the anterior border of the papillae.

The female aperture is situated in a small median circular depression, close to
the anterior border of segment xiv. |

The spermathecal apertures are in grooves 6/7 and 7/8, very close together, near-
ly in line with seta b. _ |

Internal Anatomy : —The first few septa (4/5-6/7) are very thin; none are missing,
and none are markedly thickened, though 8/9, 9/10, 12/13, and 13/14 are slightly
stronger than the others.

The gizzard, in segment v, is large and barrel-shaped; it is rather soft, but by
no means vestigial. The intestine begins in xviii, but is compressed between the
prostates in xviii and xix.

The last heart is in segment xiii.

The terminations of the nephridia are apparently in the same line.

Testis sacs are present in segments x and xi; both are continuous dorsally over
the oesophagus and dorsal vessel. That in segment x is very delicate, and has some-
what the appearance of a seminal vesicle; that in xi, also delicate, is covered by the
seminal vesicles of the segment, to which the sac is adherent, though it can be
separated.

The seminal vesicles, in xi and xii, are large, and have a granular surface, but
are not otherwise lobed. In xi the two vesicles are adherent in the middle line,
though they are separable without damage; those in xii merely touch each other.

The prostates are large, and take up the whole of the three segments xvii-xix ;
they are deeply incised by the septa, and also otherwise indented. The duct is rather
short, soft, irregularly twisted, and thin, but somewhat dilated at its extreme ectal
end.

The female organs have the usual position.

The spermathecal ampulla is a very irregularly lobed sac (fig. 11). The duct is
short,—one-fourth or one-fifth the length of the ampulla,—and marked off by a con-
striction at its beginning. The diverticula are small swellings, three or so in number,
side by side on the upper half of the duct; they have a metallic appearance, due to
the iridescent spermatozoa shining through.

There are no penial setae.

DO Memoirs of the Indian Museum. [Vor. VII,

Remarks :— Here again, in the pale colour and the testis sacs, there is a resem-
blance to P. himalayanus. But here too the differences seem to be too great to allow
us to unite the two forms in a single species. Apart from less important features such
as the numbers of the setae and the extent of the dorsal pores, we have to consider
the extent of the clitellum, the much greater development of the gizzard in the pre-
sent form, and especially the quite different configuration of the male field and the
much greater approximation of the pores.

var. affinis, var. nov.
Plate mies Ar re
Sitong Ridge, alt. ca. 4,700 ft., Darjiling Dist., E. Himalayas. 22-28-x-1917. N. Annandale
and F. Gravely. Two specimens, one not fully mature.
Jor Pokhri, 4,800 ft., Sitong, Darjiling Dist., E. Himalayas. 22-28-x-1917. N. Annandale and

F. Gravely. A single specimen.

External Characters :—Length 55 mm. ; diameter 225 mm. A slight slaty or
purplish tinge dorsally, with well-marked median darker stripe; pale ventrally.
Segments 105.

Prostomium epilobous 2, the tongue not cut off behind.

Dorsal pores begin from 4/5.

The setal rings are almost unbroken both dorsally and ventrally ; the ventral
break is absent or practically so throughout, and there is no dorsal break in the
hinder part of the body ; anteriorly a small break exists on the dorsal side, where 22
—14yz. The setae are closer set ventrally than dorsally. The following numbers
were counted :—v/38, ix/44, xii/45, xix/37, and in the middle of the body 36.

The clitellum extends over four segments, xiii to xvi; it is narrowed, but not
much modified otherwise.

The male field (fig. 12), on segment xviii, consists of a depression with sloping
sides; on these sides are placed the papillae with the male pores, so that these face
somewhat inwards. The papillae are rather wider transversely, are delimited by
grooves in front and behind, the grooves in front being the better marked; and are
separated in the middle line by a slight interval, while laterally they fade away into
the general surface without any definite delimitation. The apertures are small trans-
verse slits, their centres in line with seta c.

The female aperture is marked by a median roundish pit anteriorly on segment
xiv, abutting on groove 13/14.

The spermathecal pores are in grooves 6/7 and 7/8, opposite the interval cd.

Internal Anatomy :—A number of septa in the anterior part of the body are
slightly thickened, but none more than slightly.

The gizzard is in segment v, and is of moderate size and fairly firm. The intes-
tine begins perhaps in xviii, but is narrow in xviii and xix, where it is compressed. by
the prostates.

The last heart is in xii.

The ducts of the nephridia appear to end at different levels on the body-wall ;

1920. ] J. STEPHENSON: Oligochaeta from India and BE. Persia. 211

but, since there are no end-bladders as in P. sansibaricus, it isnot always easy to see
where exactly they end, and in any case there is no regular alternation.

It is difficult in some of the examples of this and the last few species to demon-
strate the testis sacs to complete satisfaction. Here they seem to be present in seg-
ment x, the mass of developing spermatozoa being covered over by a thin filmy mem-
brane ; the sacs of the two sides are probably continuous beneath the gut. In seg-
ment xi the testis sac is continuous on each side with the seminal vesicle.

The seminal vesicles are in segments xi and xii; they are large, taking up the
whole available space, with a granular surface but not otherwise lobed; there is no
indication of separation dorsally,—the pair in each segment is completely fused.

The prostates are large, and take up the whole of three segments, xvii to xix;

‘they are deeply indented by the septa, and also otherwise much cut up. The duct is
moderately long, is bent with the angle directed backwards, is soft and rather thin in
its ental, thicker and shining in its ectal portion.

The female organs have the usual situation.

The spermathecae, in segments vii and viii, have each a large irregularly lobed
sac-like ampulla, which is as broad as long. The duct is stout, slightly shiny, well
marked off from the ampulla, about half as broad as the ampulla but considerably
longer,— 14 times as long (fig. 13). The diverticulum appears as a rounded knob at
the ental end of the duct, with two small masses of iridescent spermatozoa shining
through.

There are no penial setae.

Remarks :—The above is the description of the more mature of the two specimens
from Sitong Ridge. The specimen from Jor Pokhri differed slightly; thus the papil-
lae bearing the male pores, and so the male pores themselves, were rather further
apart,—opposite setae d instead of c, and separated by perhaps one-seventh of the
circumference as against one-twelfth in the specimen described above ; the depression
containing the papillae was less marked; and the spermathecal pores were wider
apart,—opposite setae d or e instead of the interval cd.

The closest relative of the present form is certainly the species last described.
In this, the numbers of the setae are smaller, and the configuration of the male field
and the degree of separation of the male and spermathecal apertures also differ. The
most important points however are the spermathecae (the figures show the great
difference in the relative length of the duct), and the position of the last heart (in xii
here, in xii in the last form).

It is difficult, in cases such as this, where a number of related forms have appar-
ently arisen recently, or are possibly yet in process of differentiation, to know when
the degree of differentiation which justifies the creation of a new species has been
attained. Had this form been found at a distance from P. pokhrianus, its separation
as a different species might perhaps have been justifiable; so far as the recollection
of my Own experience goes, the position of the last heart does not, I believe, vary
within the limits of recognized species. (But see description of Octochaetus prashadi,

post. p. 233).

212 Memoirs of the Indian Museum. [Vou. VII,

Perionyx alatus, sp. nov.
Plate IX, figs. 14-10.
Sitong Ridge, alt. ca. 4,700, ft. ; Darjiling Dist., E. Himalayas. 22-28-x-1917. N. Annandale
and F. Gravely. Three specimens.

External Characters :—Length 84 mm.; diameter 3 mm. Colour dusky purple
dorsally; pale, slightly brownish below. Segments 123.

Prostomium epilobous À, tongue not closed behind.

Dorsal pores begin from groove 4/5.

The setal rings are closed dorsally and ventrally; the setae are closer set ven-
trally. The following numbers were counted :—v/50, ix/55, xii/ca. 54, xix/50, and
in the middle of the body ca. 52.

The clitellum extends from segment xi as far as the anterior third of xvii
(=44). It is slightly lighter in colour, and is the same diameter as the rest of the
body; setae and intersegmental grooves are visible.

The male field takes up the whole of the ventral surface of segment xviii (fig. 14).
Its. rounded lateral borders occupy the ventro-lateral region of the segment, and its
anterior and posterior borders correspond with the limits of the segment. Its chief
feature is the presence of a pair of large transversely elongated papillae, conjoined
by a narrow neck in the middle line, their outer ends rather narrower, and their
margins crenulated. The conjoined papillae are surrounded by a deep groove or
moat. The male pores appear as transverse grooves in the broader, inner part of
the papillae; the actual openings seem to be at about the middle of the grooves,
the distance between them being approximately one-fourth of the transverse extent of
the visible ventral surface, as seen looking down on it. The penial setae are seen as a
number of black dots in the grooves.

The female aperture is represented by a mid-ventral small depression anteriorly
on segment xiv.

The spermathecal pores are two pairs, in grooves 6/7 and 7/8, the same distance
apart as the male pores, about in line with the setal interval de.

Internal Anatomy :—No septa are visible in front of 5/6, which is very thin ; none
are notably thickened, though 6/7, 7/8, and 8/9 are slightly so.

The gizzard, in segment v, is large, long from front to back, cylindrical, and
rather soft. The intestine begins behind the prostates, in segment xx.

The last heart is in segment xiii.

The nephridia end in the same line.

Testis sacs are present in segments x and xi. In segment x1 they le beneath
the seminal vesicles, to which they are adherent, though it does not seem that their
cavities are continuous ; the membrane which constitutes the sac is fine, and rather
indefinite, and there is some matting together of all the structures, including the
hearts, as if all were enclosed in organizing connective tissue. In segment x the con-
dition is similar ; the testis sac is adherent to the seminal vesicle, which spreads over
it from the segment behind.

1920. | J. STEPHENSON : Oligochaeta from India and E. Persia. 213

The seminal vesicles belong to segments xi and xii. Those of a segment are fused
together dorsally over the alimentary canal without any hint of a division; as already
mentioned, the vesicle of xi extends forwards dorsally over x as well.

The prostates are large, occupying segments xvii to xix. They are cut up by the
septa, and also otherwise indented into numerous lobes. The duct is irregularly twis-
ted, soft, moderately long, and its ectal portion is wider than its commencement.

The female organs have the usual situation.

The spermathecae lie in segments vil and vili; the posterior pair are the larger.
The ampulla is a considerable sac, with smooth surface, not indented. The duct is
separated by a constriction, and is slightly swollen below the constriction; it is very
stout,—one-third as thick as the ampulla, or even at its upper end quite half as thick :
its length is about two-thirds that of the ampulla. Its swollen upper part corresponds
to the diverticulum, and spermatozoa can be seen shining through in small patches,
but there are no definite seminal chambers (fig. 15).

The penial setae (fig. 16) are 1:08 mm. in lensth, and 20: in thickness. The shaft
is almost straight for the greater part of its length, but has a sharp curve at its proxi-
mal end, resembling that of a hockey stick, and is more gently curved towards the
tip. The point is blunt; and the ornamentation consists of irregularly scattered
minute spines on the distal portion of the shaft.

Remarks :—This species belongs to the same group as the preceding (presence of
testis sacs, and character of the spermathecal diverticula). It is however well differ-
entiated by the presence of penial setae, and the characteristic conformation of the
male genital field; the specific name refers to the large wing-like papillae of this
region,

Perionyx shillongensis, sp. nov.
Plate IX ner
Shillong, Assam ; 4,500-5,000 ft. 16-20-iv-1918. N. Annandale. Two specimens

External Characters :—Length 66 mm. ; diameter 3 mm. Colour a dusky purple
dorsally, passing through violet to the pale unpigmented ventral surface. Segments
120. The animal is circular in transverse section (the species of the genus are usually
more or less flattened dorso-ventrally).

Prostomium epilobous #, tongue not cut off behind.

Dorsal pores begin from groove 3/4.

The setal rings present a very small and rather irregular dorsal break (z2—=I4yz),
absent altogether in the hinder part of the body; and a small ventral break (1}ab),
more regular than the dorsal but absent in the posterior third. The setae are rather
more closely set ventrally. The following numbers were counted :—v/42, ix/46, xii/49,
xix/48, and in the middle of the body 48.

The clitellum extends over five segments, xiii to xvii; it is very slightly swollen,
and there is a slight difference in tint, but this region is otherwise unaltered.

The male field is a white, rather swollen, transversely elongated oval area on the
ventral surface of segment xviii, about two and a half times as broad as long, and

214 | Memoirs of the Indian Museum. [Vor. VII,

taking up the length of the segment. The pores are fairly conspicuous, rather close
together, about in line with the setal interval cd.

The female pore is apparently situated in a small whitish area anteriorly on seg-
ment xiv.

The spermathecal apertures are situated in grooves 7/8 and 8/0, about the same
distance apart as the male pores or slightly closer, in line with the interval be.

Internal Anatomy :—The septa of the anterior part of the body as far back as the
prostatic region are slightly thickened, with the exception of the first, 4/5, which is
very thin; perhaps 6/7-9/10 are most thickened.

The gizzard, in segment vi, is of fair size; the walls are somewhat, but not very,
soft, and it must be reckoned as well developed for a species of this genus.

The intestine begins in segment xvi; no calciferous glands were seen.

The last heart is in segment xii.

The ducts of the nephridia end in approximately the same line.

Testes and large funnels are free in segments x and xi.

The seminal vesicles are in xi and xii; they are large, and bulge out the segments
in which they are contained, those in xii pressing back septum 12/13 to the level
of 13/14. They are smooth and not cut up into lobes. Those in xii meet dorsally
but do not fuse ; those in xi actually fuse in the posterior part of the segment.

The prostates are of moderate size, and are confined to segment xviii, though
bulging the septa forwards and backwards to some extent. The surface is indented
so as to form a number of lobes. The duct comes off from a hilus on the inner side,
is short and straight, and though moderately stout is soft and without muscular glitter.

The ovaries and funnels are in segment xiii. Tiny white subspherical appendages
by the side of the alimentary canal on the anterior wall of segment xiv appear to be
ovisacs.

The spermathecae, in segments viii and ix, are large, and fill out their respective
segments. The ampulla is an ovoid sac. The duct is half as long and almost half as
thick as the ampulla,—that is, relatively very stout. The diverticula are present as
two clusters of seminal chambers on the duct just below the base of the ampulla;
each cluster is cauliflower-like, and sessile by a ‘broad base; the clusters leave the
postero-internal and anterior or antero-internal aspects of the duct free (fig. 17).

The penial setae are ‘87 mm. long, and 20 thick. The shaft is straight, the tip
slightly bowed and bluntly pointed. The ornamentation consists of about eight rings
of rather fine spines ; there may be a very fine ninth or even tenth ring.

Perionyx fossus, sp. nov.
Plate IX, fig. 18, IQ.
Shillong, Assam, 4,500-5,000 ft. 16-20-iv-1918. N. Annandale. A single specimen.
External Characters :—ULength 86 mm. ; diameter 3:5 mm... Colour a dusky pur-
ple dorsally, pale and unpigmented ventrally, the upper tint shading off rather gradu-
ally into the lower. Segments 136. The body here again is circular in transverse
section.

1920.] J. STEPHENSON : Oligochaeta from India and E. Persia. PS

Prostomium epilobous 3, tongue cut off behind.

Dorsal pores begin from groove 4/5.

The setal rings are interrupted dorsally ; zz=2yz behind the genital region, but
is rather less elsewhere,—in general about I4yz ; the interval is regular, and the setae
z are in longitudinal lines. There is no ventral break. The ventral setae are much
closer set than the dorsal. The following numbers were counted :—v/52, ix/56, xii/56,
xix/52, and in the middle of the body 54.

The clitellum extends over half of segment xiii in front and one-third of xvii
behind (=3}). It is smooth, rather lighter in colour, and retains both the setae and
intersegmental grooves.

The male field is a deep squarish depression midventrally on xviii, which takes
up the whole length of the segment, and is about one-third of the ventral surface in
transverse extent. The anterior wall of the depression is vertical, the posterior slopes
more gently ; the sides are steeper than the posterior, but not so steep as the anterior
wall. Across the floor and sides of the excavation extends a transverse groove—very
narrow, a crack only. The male apertures appear to be situated in this crack, at the
junction of the floor and side walls of the depression ; they are thus fairly close to-
gether, and about in line with seta d or e.

The female pore is indicated by a slight depression in a rather pale circular area,
between the setal zone and anterior margin of segment xiv.

The spermathecal pores are two pairs, in grooves 7/8 and 8/0, fairly wide apart,
—separated by about a quarter of the circumference, or in line with about the ninth
seta from the middle line.

Internal Anatomy :—No septa are notably thickened; perhaps 9/10 is most so.
The first is 4/5.

There is a rather large but soft gizzard in segment vi. There are no calcareous
glands, but the oesophagus shows the traces of transverse vascular channels in segment
xiii, though the tube is not wider here than in neighbouring segments. The intestine
begins in xvi.

The last heart is in segment xiii.

The nephridial ducts end in the same line.

Testes and funnels are free in segments x and xi.

The seminal vesicles in segment xi are quite fused together dorsally, and fill out
the whole of the segment. Those which belong to segment xii are similarly fused,—
in their hinder parts at any rate; they pass beyond the limits of segment xii and
form a large mass which extends to the hinder end of xii.

The prostates are large compact masses, which bulge the septa of segment xviii
forwards and backwards, and in this way take up a space of three or four segments.
The gland is but little cut up into lobes. The duct emerges from a deep cleft on the
inner surface ; it is narrow and bent once or twice while still within this cleft, broad-
ens on emerging, and becomes fairly stout and shining in its ectal portion.

The spermathecae fill out all available space in segments viii and ix. The am-
pulla is irregularly ovoid in shape, and the duct is half as long and quite one-third as

216 Memoirs of the Indian Museum. Bloom WL,

thick as the ampulla. The diverticula are rather inconspicuous ; they are small flattish
swellings on the duct at about the middle of its length, two in number, sessile, and
lobulated (fig. 18).

The penial setae (fig. 19) are very little modified. In length they are ‘45 mm.,
in thickness 18,; their shape is that of an ordinary seta, with the usual double curve.
The tip is fairly sharply pointed, and there is no nodulus. The margin shows a few
small indentations near the tip.

Perionyx turaensis, sp. nov.
Plate X, figs. 20, 21.
Above Tura, Garo Hills, Assam ; 3,500-3,900 ft. ; under bark. July-August 1917. S. Kemp.
Five specimens and two fragments.

External Characters :—Length variable, but the differences are probably due to
fragmentation. The animal seems to break up easily ; one specimen shows a regene-
rated zone at the hinder end. The longest specimen, apparently complete, measured
74 mm.; diameter 2 mm. Colour dark brownish purple dorsally, with still darker
median stripe; ventral surface unpigmented. Segments 132. The ventral surface is
somewhat flattened.

Prostomium epilobous 3 or rather more, with squarish posterior tongue, which
may or may not be demarcated behind.

Dorsal pores from groove 4/5 or 5/6.

The setal rings are almost closed ventrally; on the dorsal side zz may be twice
yz in front of the clitellum, but is less behind. The setae are closer set ventrally,
and the ventral setae appear smaller than those on the dorsal side. The numbers are
difficult to count, and there are intervals where some seem to have dropped out,
so that the following are estimates only :—v/48, ix/56, xti/54, xix/44, and in the
middle of the body 55.

The clitellum is rather narrowed; it includes two-thirds of segment xii and
extends back to the hinder border of xvii (=42). The intersegmental furrows are
not obliterated.

The male pores are on xviii, close together near the middle line, on small round
papillae. The papillae touch each other or almost so, and are situated in a slight
common depression, rectangular or transversely oval in shape, which does not take

up the whole length of the segment.

The female pore is single, and appears as a simple depression or as a transverse
slit in front of the setal zone of segment xiv.

The spermathecal apertures are two pairs, in grooves 7/8 and 8/9, close together
near the middle line.

Internal Anatomy :—No septa are specially thickened, though 8/9 and in an even
less degree 7/8 are slightly so.

The gizzard, in segment vi, is rudimentary.

A pair of calcareous glands are present in segment xiii as well-defined ovoid
swellings in which the vascular channels run longitudinally. The intestine begins
in xviii.

1920. | J. STEPHENSON : Oligochaeta from India and E. Persia. 21

The last heart is in segment xii.

The arrangement of the anterior male organs was not quite clear. There are
seminal vesicles in segments xi and xii,—large square masses filling out the segment,
attached to the anterior septum of the segment, those of xi perhaps partly fused
together in the middle line, those in xii contiguous only. In x there is a similar
structure; in the specimen first dissected it was definitely noted to be a sac, not
merely a compact mass of coagulum; the funnels appeared to be contained within
them, so that they would be testis sacs. In another specimen (in which however,
as noted below, the male organs had an abnormal position) there were no sacs in the
corresponding segment,—only a cleanly detachable mass of coagulum ; and the fun-
nels were free in this and the next segment.

The prostates are compact cubical masses in segment xviii, the septa of which
are not bulged backwards or forwards. The short and moderately stout duct passes
transversely inwards from the hilus.

The ampullae of the spermathecae have a peculiar shape; the anterior border
is deeply indented, so as to form two or three rounded lobules (fig. 20), the lowest
of which may simulate a diverticulum. The duct is thick, short, and not definitely
marked off from the ampulla. What I take to be the real diverticula are a few small
rounded knobs at the ental end of the duct, which apparently are not always present.

The penial setae (fig. 21) are 5 mm. long, and 11 thick at the middle. The
shaft is straight with a slight curvature at the distal end, and tapers rather rapidly.
The tip however is cut off squarely, and carries five or six finespines. There are also
six circles of fine spines on the curved distal portion of the shaft, just above the tip.

The abnormal specimen mentioned above had the genital organs two segments
further forwards than the normal. ‘Thus the male pores were on xvi, the posterior
seminal vesicles in x (extending back however as far as xii), the anterior vesicles in
ix, the spermathecae in vi and vii. The male funnels were free in segments viii
and ix.

Remarks :—The nearest relative of the present species is P. parvulus (Stephen-
son, 12), from near Ghoom in the E. Himalayas; but the penial setae and form of
the spermathecae prevent the union of the two. These two, with P. excavatus, P.
fulvus, and perhaps P. bainii, seem to form a closely allied natural group.

Perionyx pullus, sp. nov.
Plate X, fig. 22.

Belgaum, Bombay Pres. 4-vi-1917. T.R Bell. A single specimen, the posterior end broken
off.

External Characters :—Length of the fragment 62 mm. ; diameter max. 3°5 mm.
Colour dark grey on both surfaces, scarcely any difference between dorsal and ventral.
Segments present 165. Ventral surface concave except at anterior end, thus present-
ing a longitudinal groove bordered by prominent ventro-lateral ridges.

Prostomium epilobous 3, the tongue being triangular; from its backwardly
directed point a groove is continued in the middle line back to the clitellum.

218 Memoirs of the Indian Museum. [Vor. VIT,

Dorsal pores begin in groove I/2; this must be about the extreme anterior limit
which they ever attain, though I could not say that it is unique. Beddard (1)
states that dorsal pores are never found on the first one or two segments of the
body.

The setal rings are interrupted dorsally; in front of the clitellum zz=3yz, and
behind=about 2yz; but behind the clitellum the interval itself, as well as the inter-
setal intervals on each side, is irregular. The ventral break is absent, or small and
irregular. On the ventral surface the setae are small and very close together, —almost
as close as they can be. For this reason, and also because of the dark colour of the
animal, they are very difficult to count; behind the clitellum there are about 60,
and further back 64.

The clitellum is extensive, from xi to xx (=10); it is slightly swollen, less
marked or absent ventrally, rather darker in colour, and dorsal pores are absent.

The male field is situated on segment xix (whether normally, or exceptionally in
this particular specimen, cannot be said). It consists of a mid-ventral rectangular
area, rather broader than long, which takes up most of the length of the segment ;
it is delimited at the sides by slight grooves and in front and behind by deep tren-
ches, which coincide with the intersegmental furrows; the trenches are however
much broader than the intersegmental furrows, and encroach anteriorly and post-
erlorly on the surface of the segment. The pores are apparently on two small whitish
papillae very close together near the midventral line.

The female pore was not visible.

The spermathecal apertures are two pairs, in grooves 7/8 and 8/9, close to the
middle line.

Internal Anatomy :—No septa are specially thickened.

There is no gizzard, even vestigial. The pharyngeal glands are especially bulky,
and extend as large masses on the alimentary canal backwards to segment ix. The
oesophagus is bulged laterally, and its walls are vascular, in segments x-xui. The
intestine begins in xvil.

The last heart is in segment xii.

The nephridia end in approximately the same line.

The male funnels (presumably testes also) are free in segments x and x1.

Seminal vesicles occupy segments xi, xii, and xiii. They are relatively small,
are attached to the anterior wall of their segment in each case, and have a racemose
appearance owing to their being composed of a number of small lobules; indeed the
extremely racemose appearance of those in segment xiii caused them momentarily to
be mistaken for ovaries. There appeared to be an additional minute pair of racemose
vesicles in xiv also.

The prostates, in segment xix, are small, and consist of a number of elongated
finger-like lobes, the whole forming a bushy cluster. The duct is small, soft, not
much thickened, and runs transversely inwards.

Ovarian funnels were identified in xi.

The spermathecae (fig. 22) are situated in segments viii and ix. The ampulla is

1920. | J. STEPHENSON : Oligochaeta from India and E. Persia. 219

very irregular in shape, and narrows below to become the duct without any distinet
delimitation. If the beginning of the duct is taken to be just below the diverticula,
it is about as long as the ampulla; it narrows gradually towards its ectal end. The
diverticula, about three in number, are small rounded sessile chambers, situated at
what may be considered as the lower part of the ampulla.

There are no penial setae.

Perionyx minimus, sp. nov.
Plate X, fig. 23.
Belgaum, Bombay Pres. 4-vi-1917. T. R. Bell. Numerous specimens.

External Characters :—The worms are very small; a long one measures 45 mm.
in length, while the thickness is only I mm., or as a maximum 12 mm. The colour
is a medium brown dorsally, and a rather lighter brown ventrally. Segments roo.
The ventral surface is flattened.

Prostomium epilobous 4 or nearly so, tongue cut off behind; both prostomium
and first segment divided in the mid-dorsal line by a longitudinal groove.

Dorsal pores from 4/5; or a rudimentary pore in 4/5, and the first well developed
pore in 5/6.

The setal rings are almost closed ventrally; the dorsal interval is well marked,
—2yz. The setae are much closer set ventrally than dorsally. On the twentieth
segment there are about 26 setae, in the middle of the body about 36.

The clitellum extends from the middle, or perhaps the anterior end, of xiii
backwards to include segment xvii (=4$ or 5); it is rather narrowed, of a somewhat
darker colour, and the separate segments are easily distinguishable.

The male pores are on conspicuous round papillae on xviii. The area between
these papillae is depressed, the depression extending longitudinally from the middle
of xvii to the anterior third of xix; this area is encroached on laterally by the
papillae, so that the depression has a dumbbell shape, the dumbbell being placed
longitudinally ; the apertures look somewhat inwards.

The female pore (or pores) are situated in a transverse groove anteriorly on
segment xiv.

The spermathecal pores were not distinctly seen, but appeared to be about a
quarter of the circumference apart, in grooves 7/8 and 8/0.

Internal Anatomy :—No septa are thickened.

There is apparently a rudimentary gizzard in segment v. The oesophagus is
slightly bulged in segments xiii and xiv, with longitudinal vascular striations. The
intestine begins behind the prostates, in segment xix. The “pharyngeal glands”
extend back to segment vii as definite lobes on each side filling out the segments.

The last heart is in segment xii.

Relatively large seminal funnels are present in segments x and xi, embedded in
a mass of very adherent flocculent matter simulating seminal vesicles; the testes
were not separately identified. |

Seminal vesicles are present in segments ix and xii; both pairs are brown in

220 Memoirs of the Indian Museum. [Vor. VII,

colour. Those in xii are large and lobulated, and meet but do not fuse with each
other dorsally ; the vesicle on the right side in segment ix was moderately large, but
that on the left was small (? had been in part detached and washed away).

The prostates occupy more than one segment, —xviii and part of xvii on the
right side, xviii and part of xix on the left; they are somewhat loosely lobulated.
The duct is short, narrow and rather soft; it is covered over by the lobules of the
gland, and is not visible till these are separated.

The ovaries, relatively very large, and funnels are in segment xiii. In segment
xiv are conspicuous ovisacs, containing large ova.

The spermathecae are of moderate size, in segments viii and ix. The ampulla
is rounded, and rather flattened between successive septa. The duct is of about the
same length as the ampulla; at its ental end, between the two diverticula, it is
rather narrow, but becomes thicker below them (fig. 23). The diverticula are small,
subspherical, and attached to the ental end of the duct by short stalks; they have
rather a cauliflower-like appearance, but this is not due to any lobulation,—it seems
to be occasioned by wisps of iridescent spermatozoa shining through the wall.

There are no penial setae.

Perionyx igatpuriensis, sp. nov.
Plate X, fig. 24.
Igatpuri, Bombay Pres. 29-vi-1917. B. Prashad. Three specimens, one consisting of only
the anterior end.
Elephanta Island, Bombay ; on the sea-shore. 30-vi-1917. B. Prashad. Three specimens.

External Characters :—Length 40-52 mm.; diameter 2-2°5 mm. Colour dark,
purple dorsally, pale ventrally; clitellum slightly lighter dorsally than the rest of
the dorsal surface. Segments 150-170.

Prostomium epilobous $, tongue broad, cut off behind.

Dorsal pores begin from groove 4/5.

The setal rings are unbroken dorsally ; the ventral break is small and irregular,

or altogether absent. The setae are closer set ventrally than dorsally. The following
numbers were counted :—v/36, ix/48, xii/46, xix/4I, and in the middle of the body 44
(in the specimen examined from the Bombay material the numbers were somewhat
greater in the anterior segments).

The clitellum extends over xili-xvii (only to xvi in Bombay specimen) (=4 or 5);
setae are present, and the intersegmental grooves can be distinguished.

The male pores appear as depressions, each in the middle of a small circular area
with slightly raised lips. near the middle line in segment xviii; these areas are sepa-
rated only by a groove in the mid-ventral line, and in length take up nearly the
whole segment. Black dots, which may be seen in the depressions, are the penial
setae.

The female pore is represented by a whitish dot in a transversely oval depression
anteriorly on segment xiv. a

The spermathecal pores are two pairs, in grooves 7/8 and 8/9; they are conspicu-

1920.] J. STEPHENSON : Oligochaeta from India and E. Persia. 221

ous round apertures near the middle line, the same distance apart as the male
pores.

Internal Anatomy :—The first septum appears to be 5/6; all are present, and all
are thin.

The gizzard, in segment vi, was soft and rather small in the original specimen
from Igatpuri; in the specimen from Bombay however it was of large size, but soft
and yielding easily to pressure with a needle; in shape it was rather cylindrical, but
somewhat narrower posteriorly.

There are no calciferous glands; the gut may be bulged laterally in segments
xiii and xiv. The intestine begins in xxiii.

The last heart is in xiii.

The endings of the nephridia are in the same line.

Testes and funnels are free in segments x and xi; these segments are full of
flocculent material (developing spermatozoa).

Seminal vesicles occupy segments xi and xii; they are large paired sacs, not
fused in the middle line. They are rounded in form, their borders not indented or
lobed (in the Bombay specimen they appeared somewhat granular, as if made up of
minute lobules). Those in xii are the larger, pressing back septum 12/13.

The prostates are compact masses occupying segment xviil, the limiting septa of
which are much bulged apart by their bulk; each consists of an anterior and poste-
rior lobe, from between which the duct issues. The duct is short, straight, and narrow,
and passes transversely inwards.

The large ovaries and the funnels are in segment xui.

The spermathecae, in segments viii and ix, present an elongated ovoid or irregu-
lar ampulla with a short stout duct, a third or a quarter as long and a third as wide
as the ampulla. There is a single diverticulum, attached to the base of the ampulla,
sessile, cauliflower-like and consisting of a number of small seminal chambers ; in ex-
tent it is one-third or a quarter as long as the ampulla, against the lower part of
which it may be flattened (fig. 24).

The penial setae are 44-52 mm. long, and 15. thick in the middle. The shaft
is almost straight, very slightly bowed,—more so at the distal end ; the tip is simply
and rather bluntly pointed; there are about six rings of rather small spines near the
tip. (In the Bombay specimen the setae were a little larger, —'68 mm. long, and 204
thick, with seven well-marked rings of small spines, and two or three rings only very
faintly indicated proximal to these).

Remarks :—The present species resembles P. millardı,— very closely in many res-
pects. The distinctive difference is in the spermathecae, which there have no diverti-
culum. Minor differences are the setal rings,—the breaks dorsally and ventrally being
smaller or mostly absent here; and perhaps the penial setae, which have fewer rings
of spines in the present species.

Perionyx spp.
In the various collections examined there were a number of specimens of this
genus which could not be referred to any species on account of immaturity. Some

222 Memoirs of the Indian Museum. [Vor. VII,

such were obtained from above Tura, in Assam ; from the Western Ghats; from Pa-
shok in the Darjiling Distriet; from the Sitong Ridge, and from hill streams near
Sitong, also in the Darjiling Distriet. All these regions are already in the recognized
area of distribution of the genus, and the specimens may be dismissed with a couple
of remarks on their habits. Of a batch of specimens found above Tura, Mr. Kemp
remarks, ‘This worm is found coiled up on the upper or under sides of leaves in dense
jungle, forming a compact gelatinous mass. When touched it springs to life, perform-
ing somersaults and other acrobatic feats.” The second noteworthy feature is the
aquatic habitat of the worms from hill streams near Sitong; they were found living
in water under stones.
Genus Lampito.
Lampito mauritii, Kinb.

This worm is one of the commonest in India,—absolutely the commonest in the
present collections ; and being so widely distributed it is scarcely necessary for the
future to note the precise details of each capture.

The following are the localities from which I have received it :—In the Central
Provinces and Central India, from Nemar Kheri on the way to Indore, Katni, Gwalior
and Jubbulpore ; in S. Rajputana, from Dungarpur and Banswara ; and in the Bom-
bay Presidency from Bombay (where it is very common), Broach, Surat, Ahmedabad,
Nadiad, Dhanu, Sirvai Madhopur, Baroda, Palchar, and Joshachivir on the way to
Mahableshwar. :

Genus Pheretima.
Pheretima posthuma (L. Vaill.).

Also extremely common; from Khulna and Dattapakur in Bengal; Udaipur in
Rajputana ; Gwalior in Central India; Bindra Ban, near Muttra, United Provinces ;
and Baroda in the Bombay Presideney.

Pheretima hawayana (Rosa).

A common Indian worm ; from Bindra Ban, Udaipur, and Bombay.

Pheretima heterochaeta (Mchlsn.).

Common in India; from Sureil, 5,000 ft. in Darjiling District. An immature
specimen, probably of this species, from Pashok, 3,500 ft., also in Darjiling District.

Pheretima elongata (E. Perrier).

Manmad, Bombay Pres. 28-vi-1917. B. Prashad. Several specimens.

Palia, between Indore and Ujjain, Central India. 27-vi-1917. B. Prashad One specimen,
a mutilated anterior end.

Indore, Central India. 23-vi-1917. B. Prashad. Three specimens.

Indore, banks of River Kan. 25-vi-1917. B. Prashad. Several specimens, immature, but
probably of this species.

Ujjain, Central India. 26-vi-1917. B. Prashad. Several specimens, with others, probably
of the same species, but immature. |

1920. ] J. STEPHENSON: Oligochaeta from India and E. Persia. 223

Namkana, Sunderbans, Bengal ; near the banks of a reclaimed island. 10-xi-1918. B. Pra-
shad. Several specimens.
Caleutta. Nov. 1913. E. C. Dormieux. A single specimen.

The length of the specimens from Manmad (the only ones subjected to a complete
examination) varied; a long one was as much as 230 mm.

Large testis sacs, enclosing alimentary canal, hearts, and dorsal vessel, as well as
the testes and funnels, were present in segments x and xi; and seminal vesicles in xi,
xii and xili, as noted by previous observers. The seminal vesicles of xi are contained
within the testis sac of that segment ; those of xii are large, and meet mid-dorsally ;
those of xiii are small and rounded.

It may be noted that the last heart was in segment xii, in the specimen dissec-
ted. I found no spermathecae (this condition has previously been noted in the spe-
cies).

Remarks :—This worm has not hitherto passed for common in India. It was re-
corded from three localities by Michaelsen (3) as a result of his examination of the
extensive collection of the Indian Museum, —from Hyderabad, Deccan; Kandy in
Ceylon; and Karachi in Sind. I had not myself met with it previously. Like others
of the genus, the present species has a wide distribution outside India—indeed it
might be called a ‘“ world-wanderer.”’

The worm has generally gone under the name of P. biserialis ; Michaelsen was
able, by an examination of the types of P. elongata, to establish its identity with the
latter (4).

Pheretima lignicola, Stephenson.
Bombay. June, 1915. N.B. Kinnear. Several specimens, not all mature.
Bombay. 7-xi-1914. N. B. Kinnear A number of specimens.

The colour appears to be variable; olive-green and brown have been noted in
previous-specimens; these,—the first of the two batches at least,—were a metallic
bluish purple dorsally, and pinkish ventrally.

The setae of segments ii-ix were larger than those on the rest of the body. The
intestinal caeca were crenulated on their dorsal margin.

Subfam. OOTOCHAETINAE.
Genus Hoplochaetella.
Hoplochaetella anomala, sp. nov.
Plate X, figs. 25-29.
Belgaum, Bombay Pres. 5-vi-1910. N. B. Kinnear. Eight specimens, and a fragment con-
sisting of the posterior end. In bad condition.

External Characters :—Length about 85 mm. ; diameter 3 mm. The worms are
apparently unpigmented or almost so; in their present condition, with much thinned
body-walls, the colour is given by the intestinal contents. Segments ca. 105.

Prostomium epilobous À, tongue broad, sides slightly converging behind, not cut
off by a transverse furrow.

224 Memoirs of the Indian Museum. [Vor. VII,

Dorsal pores begin from 4/5.

The setal rings are almost closed ventrally ; aa=14-2ab. The dorsal break is
about 4yz in front of the clitellum, 3yz behind it, and 2yz in the middle of the body.
The setal intervals decrease somewhat outwards from the middle line, so that aa > ab >
be>cd. The numbers of setae are difficult to ascertain, since the worms will bear
scarcely any manipulation, and doubtless many setae have dropped out; I could only
count the intervals between the muscle bundles :—viii/36-44, xii/40-46, and in the
middle of the body 40.

" The clitellum extends from $xiii to }xvi (=3); it is smooth, brownish, swollen
and well defined; setae are present, but no intersegmental furrows.

The male area (fig. 25) presents two pairs of well-marked depressions, crater-like,
with thickened and rounded margins, on segments xvii and xix; these are rather
oval in a transverse direction. The lip is less definite on the inner side, where it
merges into a mid-ventral thickened area between the two depressions of a pair. The
depressions extend transversely between the lines of setae b and A, and with their lips
and the median thickened area take up the whole transverse extent of the ventral
surface ; longitudinally also they occupy the whole of the length of their respective
segments, without however encroaching on segment xviil.

The male apertures are in the inner portions of the excavations, between the
lines of setae c and d.

The female aperture is single, in a small circular whitish area just in front of the
setal zone of segment xiv.

The spermathecal apertures are two pairs, on small papillae on segments vii and
ix; they are in line with seta c, and about one-fifth of the circumference apart. In
segment ix the papillae are in the setal zone, in viii in front of it.

The above description is that of the specimen of which a complete examination,
internal and external, was made. In some of the other specimens there were a few
variations in the external genital marks which deserve mention.

In one, there was a cup-shaped shallow depression, also with a thickened rounded
margin, on the left side of segment xviii; this was rather smaller than those on xvii
and xix, much shallower, and slightly internal in position to them. In another, there
were depressions on both sides of xviii. Or the depressions containing the male pores
may be smaller than above described, and circular in shape, not extending so far out-
wards; or the depressions may be reduced by the filling up of their outer part, the
lip being much thickened here, almost like a papilla to the outer side of the pit.

A displacement of the setal line in the spermathecal region has been noticed in
other species of the genus. This was not the case in the specimen first examined,
though setae cde appeared to be absent on both sides of segments viii and ix. In
another, cdef were displaced forwards on both sides in segment viii; cd were displaced
backwards on the right side in ix, while on the left side the setal line was irregular.

Internal Anatomy :—The septa were all softened ; they are apparently distinguish-
able as far forwards as 3/4, and 11/12 and 12/13 are perhaps somewhat thickened.

The gizzard is of large size, regularly ovoid, firm, in segment vii. The calcifer-

1920.] J. STEPHENSON : Oligochaeta from India and E. Persia. 225

ous glands are in segments x-xiii, small, ovoid, set off from the alimentary canal; in
segments x and xi they are within the testis sacs. The intestine begins in xvi.

The last heart is in segment xii. There is no large transverse vessel in xiii, but
anteriorly in xiv a pair of stout vessels are given off from the dorsal vessel,—these
may however only go to the alimentary canal.

In the first nineteen segments only micronephridia are present; some of these
are scattered, and others form large tufts anteriorly by the side of the pharynx.
Meganephridia begin from segment xx; probably micronephridia coexist, but have
become unrecognizable.

Testis sacs are present in segments x and_xi as large chambers which enclose the
dorsal vessel and alimentary canal as well as the testes and funnels; those of seg-
ment x enclose in addition a pair of seminal vesicles.

The seminal vesicles are in segments ix, x, and xii; those in x, within the testis
sacs, are attached to the posterior wall of the segment ; those in ix and xii are large,
and only slightly indented into lobes.

The prostates are two pairs of coiled tubes. The anterior occupy segments xvii
and xviü, the posterior segments xix to xxi. The duct is in each case stouter than
the gland, shining, straight and rather short, thinner at its ental end; each passes
obliquely forwards and inwards. The ends of both pairs of ducts are surrounded by
a soft white cushion on the inner face of the body-wall.

The vasa deferentia are two on each side, and pass backwards side by side beyond
the termination of the anterior prostatic duct (fig. 26); shortly behind this, the outer
duct of the two doubles back, after crossing underneath the inner, and ends immedi-
ately behind the termination of the anterior prostatic duct; the other vas deferens
goes on, and ends immediately in front of the termination of the posterior prostatic
duct.

The ovaries are in segment xiii; there are small ovisacs in xiv.

There are two pairs of spermathecae ; the ampulla is a sac of very irregular form
(fig. 27),—in some cases a portion is almost constricted off. The duct is stout, near-
ly as long as the ampulla, dilated and not shiny in its ental part, narrower and shining
below; at its thickest part it is half as broad as the ampulla; it is separated from
the sac above it by a constriction. The diverticula are two, opposite each other,
attached to the duct immediately below the dilated part; each consists of a few
rounded seminal chambers, the whole being sessile on the duct. The endings of the
ducts, as seen on the inner side of the body-wall, correspond in position to the
papillae seen externally.

As in other species, there are a number of accessory glands. These are clusters
of finger-shaped structures, three to five in each cluster, of various lengths, the
longest about equal to the duct of the spermatheca or rather longer. Hach cluster
ends near the termination of a spermathecal duct. The accessory glands themselves
are solid, but have a distinct and fairly long duct with a lumen, and they are
connected, where they pierce the body-wall, with the modified copulatory setae of
segments villi and ix; fig. 28 is a rough sketch of gland and seta extracted together.

226 Memoirs of the Indian Museum. [Vor. VII,

These copulatory setae are in length ‘61 mm., and in thickness at the middle
224 : they are almost straight, with a slight curve at the proximal end, tapering
and bluntly pointed distally. There is scarcely any ornamentation,—only a few very
fine oblique lines, or semicircular markings convex proximally, near the tip
(fig. 29).

There are no penial setae in the segments of the male apertures.

Remarks :—The above interesting form has obvious relations with those I have
previously described (13) under the name Hoplochaetella; it has, besides the same
arrangement of prostates and spermathecae, the same displaced and modified setae in
the neighbourhood of the spermathecal apertures, the same distribution of calciferous
glands, and the same peculiar nephridial system. It does not however agree in all
points with the emended definition of the genus which ! gave.

Of the points of difference, one of the most interesting is the manner of ending
of the vasa deferentia; in the other species of the genus the vasa deferentia unite,
and then open in common with the anterior prostate on the seventeenth segment,
while here the vasa are separate, and open, one with the anterior prostate on segment
xvii, and the other with the posterior on xix.

In the Megascolecidae, what may be described as an attraction between the ter-
minal portions of the genital organs is of very general occurrence. The primitive
condition in the family is that known as the “original Acanthodriline,” where the
prostates end on segments xvii and xix, the vasa deferentia (after joining together)
on xviii, and the spermathecae in furrows 7/8 and 8/9. In the Megascolecinae, one
pair of prostates has disappeared, and the other has been “attracted” to join the
termination of the vasa deferentia in xviii, an intermediate stage being seen in the
genus Diplotrema. In the other species of Hoplochaetella, and in Hrythraeodrilus, the
conjoined vasa deferentia have been attracted forwards (to continue the same figure)
to join the anterior prostatic duct on xvii. In the other species of Hoplochaetella also,
the two pairs of prostates are approaching each other, and the two pairs of sperma-
thecae show the same tendency. The very frequent union of the original pair of
female pores on segment xiv is perhaps to be referred to the same group of pheno-
mena. In Hutyphoeus (Octochaetinae) the single vas deferens on each side ends in
common with the prostatic duct on xvii. Similar instances can be quoted from the
Ocnerodrilinae.

In the present form, the attraction has taken a different course; the two vasa
deferentia of each side have been as it were pulled apart, one towards the anterior, the
other towards the posterior prostate, while these maintain their original position on
the middle of segments xvii and xix.

In regard to the above point of difference, neither the present nor the former
species of Hoplochaetella can be said to be primitive as compared with the other;
evolution has merely taken a different course in the two. In certain features how-
ever the present form appears to show a more primitive condition than the species
previously described. Thus the two pairs of spermathecal apertures, which in the
other species are both on segment viii, are here more widely separated, on segments

1920. | J. STEPHENSON : Oligochaeta from India and E. Persia. 227

viii and ix respectively. So too the prostatic apertures are in previous species pos-
teriorly on xvii and anteriorly on xix, or actually in the furrows 17/18 and 18/19;
while here they maintain their original position at the middle of xvii and xix.

Lastly, the testes and funnels are free in segments x and xi in the former species
but are contained in testis sacs in the present form. Here I am not clear as to which
is the more primitive condition. As a rule, of course, the free condition is to be
looked on as primitive, and that in which the testes and funnels are enclosed in sacs,
—separated-off portions of the coelom,—as secondary. But in the previous species
of the genus the testes and funnels are not free in the usual sense ; segments x and xi
are very harrow, and septa 9/10, 10/11, and 11/12 are fused together at their periphery
so as, at first, to give the impression of one enormously thickened septum. What
has happened is that these septa have become approximated, and the contained seg-
ments very much contracted ; whether originally, before the contraction took place,
they contained testis sacs cannot now be decided,—the sac-walls (if the sac originally,
as in the present form and often elsewhere, contained alimentary canal, hearts and
dorsal vessel) may have simply fused with the walls of the segment.

I have previously (13) shown that Hoplochaetella is to be regarded as the ances-
tor of Hrythraeodrilus. In some ways the present form may represent that ancestor
more Closely than any of the previous species; thus the condition as regards testis
sacs is the same in this form and in Hrythraeodrilus, and similarly with regard to the
number and position of the seminal vesicles (three pairs in this form and in Hrythrae-
odrilus, in segments ix, x, and xii; two, in ix and xii, in the other species of Hoplo-
chaetella). The distinctive difference between Hoplochaetella and Erythraeodrilus is
the presence of two pairs of prostates in the first, of one pair only in the second; in
this, the present form agrees with Hoplochaetella, along with which I propose to in-
clude it, widening the previously given definition of the genus (as regards the endings
of the vasa deferentia, the positions of the prostatic and spermathecal apertures, and
the free testes and funnels) for the purpose.

Hoplochaetella spp.
Daman Road, N. of Bombay (between Bombay and Surat). 7-vü-1917. B. Prashad. Five
specimens, none sexually mature.
Bombay, Malabar Hill. 1-vii-1917. B. Prashad. Numerous specimens, none mature.
Bombay, Elephanta Island, high up on a hill. 30-vi-1917. B. Prashad. Numerous speci-
mens, none mature.

In the specimens from Malabar Hill the spermathecae couid be seen on dissection
to be just forming ; they appeared to end in the furrows 7/8 and 8/9; if so, the ap-
proximation of the two pairs has not gone so far as in the majority of the species,
where both pairs are on segment viii.

The specimens from Daman Road presented one point of interest to me. The
type of the genus Hoplochaetella is Bourne’s Perichaeta stuarti ; and in identifying my
former five species as belonging to this genus (13), one point which came up for dis-
cussion was that Bourne described certain diverticula of the intestine which I did

228 Memoirs of the Indian Museum. [Vor. VII,

not find in any of my specimens ;—* In somites xxiii-xxvi (?) there are four pairs of
special diverticula on the dorso-lateral portions of the intestine.” In the example of
the Daman Road batch which I dissected I noted that the segmental swellings and
intersegmental constrictions of the intestine were very marked dorso-laterally in its
anterior part; and from about segment xxiii onwards for ten or a dozen segments
there were very distinct lateral (not however dorso-lateral) sacculi, but their extent
was rather indefinite. I did not, in my previous paper, consider the feature as of
generic significance ; still it is perhaps some slight confirmation of my identification
to find something similar (though not quite identical) in a worm which certainly be-
longs to the same group as those I there described.

Genus Octochaetus.

Octochaetus barkudensis, Stephenson.
Barkuda Island, Chilka Lake, Ganjam Dist., Madras Pres. ; at base of tree. 25-vii-1917 to 4-
viüi-1917. N. Annandale. Four specimens.
Same locality, date, and collector. Under stones on shore of island. One larger and a number
of smaller specimens.

The papillae on segment viii may be joined in the middle line, and so appear as
a dumbbell-shaped thickening.

In addition to the median oval papilla on segment xxii there may be a similar
one on xxi.

Octochaetus fermori, Mchlsn.
Dhanu, a little distance N. of Bombay. 7-vii-1917. B. Prashad. Several specimens, imma-
ture.
Surat, W. India. S-vii-1917. B. Prashad. Several specimens.
Ahmedabad, W. India. 11-vii-1917. B. Prashad. Several specimens.
Baroda, W. India; by the banks of a tank. 9-vii-1917. B. Prashad. A single specimen.
Same place ; on a small hillock. 10-vii-1917. B. Prashad. One specimen, with perhaps two
others, immature.
Gwalior, Central India : bank of a stream. 17-vi-1917. B. Prashad. Five specimens.
Same place ; under flower-pots. 17-vi-1917. B. Prashad. A single specimen, rather imma-
_ ture.
Octochaetus paliensis, sp. nov.
Plate X, figs. 30-33.
Palia, between Indore and Ujjain. Central India. 27-vi-1917. B. Prashad. A single speci-
men, not in good condition.
Indore, Central India. 23-vi-1917. B. Prashad. Several specimens.
Poona. 3-vii-1917. B. Prashad. Two specimens, one smaller, not fully mature.
Bina, Central Provinces. 19-vi-1917. B. Prashad. A single specimen.

The description which follows is of the specimen from Palia, the one which was
first examined. A few differences in the Poona specimens and in the one from Bina
will be mentioned subsequently.

External Characters :—Length 45 mm.; diameter 2°75 mm. Colour yellowish
grey, not darker on the dorsal surface ; clitellum browner. Segments IAI.

1920. ] J. STEPHENSON : Oligochaeta from India and E. Persia. 229

Prostomium apparently proepilobous (buccal cavity everted).

Dorsal pores from furrow 12/13.

Setae paired ; in front of the clitellum ab = jaa = less than half (say 2) be = 2cd ;
behind the clitellum ab = 3aa = half be = cd; in the middle of the body ab = 2aa =
3bc and is slightly less than cd.

The clitellum extends over segments xii-xvii (=5). Furrows are visible ventral-
ly, but not dorsally ; dorsal pores are absent.

The male field shows two elongated trench-like depressions, on segments xvii and
xix respectively ; these take up the whole length of the segments, and are thus sepa-
rated by a transverse ridge which represents the ventral surface of xviii; the tren-
ches extend from a little outside the line of setae b to a corresponding point on the
other side, and their lateral portions are rather deeper than the middle.

The prostatic pores are in these lateral portions, in line with setae b, on small
white papillae. The seminal grooves pass longitudinally and straight between the
pores of the same side.

The female pores could not be distinguished.

The whole of the ventral surfaces of segments viii and ix are thickened and some-
what flattened, forming a couple of low papillae, transversely much elongated, and
with their lateral ends rounded ; these extend on each side to some distance outside
the line of setae b. The spermathecal apertures appear to be just in front of the site
of setae a of these segments, though setae a and b are not visible.

Internal Anatomy :—The first septum is probably 4/5; this is moderately
thickened. Behind this is a space in which lies the gizzard; the next septum is 7/8 ;
this, and all the succeeding ones to 11/12 are somewhat thickened ; thereafter the
thickening decreases as far as 14/15, after which all are thin.

The gizzard, in front of septum 7/8, is spherical, and not very hard. There is a
pair of large calciferous glands in segment xv, on the right side getting also into xiv ;
each is kidney-shaped, with its convex margin indented. The intestine begins in xvil.

The last heart is in segment xii.

The excretory system is micronephridial.

Male funnels are free in segments x and xi; testes were only doubtfully identi-
fied.

The seminal vesicles are in segments ix and xii. In the latter segment they are
small, somewhat flattened, and lie within the curve of the hearts on each side. In
segment ix I found none on the right side ; but on the left there was a fairly large, very
deeply indented and lobed vesicle.

The prostates are two pairs of opaque, moderately thick, convoluted tubes, in
segments xvii and xix, which bulge apart the septa. The duct is thinner, shining,
wavy in its course, transverse in direction, and thinner in its first part than after-
wards.

The ovaries are in the usual situation. :

The spermathecae are two pairs. The ampulla is elongated, somewhat conical,
of moderate size; the duct is half as wide and one-third as long as the ampulla, not

230 Memoirs of the Indian Museum. [Vor. VII,

sharply marked off, and not shining. The diverticulum is single, club-shaped, with-
out distinct stalk, and in length one-third or one-fourth of the whole main pouch
(ampulla plus duct) ; it arises from the ectal end of the duct (fig. 30).

The penial setae are in length -65--76 mm., and their thickness at the middle is
16; the shaft is straight, the distal end slightly curved, and the tip bluntly pointed ;
the ornamentation consists of about eight circles of small spines near the tip (fig.
31a). A second shape also occurs, with a somewhat sinuous outline at the distal
end, and more sharply pointed tip (fig. 316); this is possibly due to such setae being
younger, and not fully straightened out.

The copulatory setae of segments vili and ix are in length 76-82 mm., and 22»
thick in the middle. The proximal half is straight, the distal portion bowed ; the tip
is rather sharply pointed, and somewhat claw-shaped, with a slight swelling just
proximal to the point. On the bowed distal portion of the shaft the convex and
concave borders are furnished each with a row of spines, or incisions (fig. 32; in the
example illustrated the spines stand off remarkably clearly ; usually they are closely
adpressed to the shaft).

In the specimen from Poona, the setal intervals were a little different :—in front
of the genital region ab = 2aa = half be = cd; behind the clitellum ab = aa = be
— cd; in the middle of the body ab = ?aa = bc = cd; dd is approximately + of
the circumference.

The trenches on the male field of the previous example are here contained within
a somewhat thickened area, which extends posteriorly to the middle of segment xx,
where it becomes joined to a transversely much elongated papilla ; this papilla covers
the posterior half of xx and the anterior half of xxi, and transversely is of equal
extent with the trenches in front.

There seemed to be a very thin septum in front of the gizzard, corresponding to
6/7 ; 5/6 seemed to be the only missing one. Small ovisacs were present in segment
XIV.

The spermathecal ampulla here has the form of an ovoid sac ; the duct is hardly
distinguishable. The diverticulum is a rather cauliflower-like cluster of small seminal
chambers, with a short stalk (fig. 33).

In some of the penial setae the tip, instead of being rounded, appears to have
sharp lateral edges. On the copulatory setae also there seems to be a lateral, rather
thick ridge on the claw-like tip ; this was more or less distinct in the previous speci-
men also.

In the specimen from Bina, the prostomium might be said to be compounded of
the prolobous and tanylobous types, —a tanylobous prostomium with a transverse
furrow at the anterior end of the “ tongue ” cutting it off from the projecting lobe in
front. The relations of the setal intervals are scarcely sufficiently different from the
type to deserve mention. The seminal vesicles, in ix and xii, were fairly bulky, and

their margins were only slightly lobulated.

1920. ] J. STEPHENSON : Oligochaeta from India and E. Persia. 231

The spermathecae were, as seen in dissection, of the simplest possible form,—
elongated sacs, narrowing somewhat at their ectal end, without diverticulum. On
microscopical examination however a diverticulum was seen, bound down to the main
sac ; this was an ovoid simple appendage, showing no seminal chambers, entering the
main sac rather nearer the ectal than the ental end; it was a little narrower towards
its attachment, but there was no stalk; its length was a quarter, its thickness less
than half, that of the main pouch.

The penial and copulatory setae were exactly those of the variety next to be
described. ees,

var. riparius, var. nov.
Plate X, figs. 34, 35.
Gwalior, Central India ; on the bank of a stream. 17-vi-1917. B. Prashad. Four specimens.
Same place ; inagarden. Same date and collector. Numerous specimens.
Same place ; under flower-pots. Same date and collector. Two specimens.

External Characters :—The largest specimen examined was 90 mm. long, and 3°5
in diameter. Colour buff or yellowish grey, no difference between dorsal and ventral
surfaces ; the clitellum may have a reddish tinge. Segments 135.

Prostomium tanylobous or almost so; furrows at the sides of the tongue parallel,
but may be only slightly marked.

Dorsal pores begin at the hinder border of the clitellum.

The setae are paired. The relations may be expressed thus:—In front of the
clitellum ab = 4aa = 3bc = 3cd; behind the clitellum = aa = 2bc = 2cd; in the mid-
dle of the body = 3aa = 2bc =cd; dd in the middle of the body is equal to 3 of the
circumference.

The clitellum extends over segments xiii-xvii; it is thickened, well defined at
both ends, the furrows obliterated but setae just indicated.

The male area is rather square, with thickened borders ; it includes segments
xvii-xix and the anterior half of xx, extending transversely across the whole ventral
surface. This area presents a dumbbell-shaped depression, expanded on segments
xvii and xix, the narrowed handle of the dumbbell being on xviii (2e. the dumbbell
is placed longitudinally) ; the narrowing of the depression on xvili is caused by two
large flat papillae, continuous at their outer margins with the thickened edge of the
general male area (fig. 34).

The prostatic pores are in the line of setae b, in the broadened ends of the dumb-
bell; the seminal grooves are convex inwards, skirting the inner border of the flat
papillae.

The whole male field nay be compared with that of the typical form by suppos-
ing that the lateral papillae of the present form have there extended mesially and
joined, thus producing two separate depressions, one on xvii and one on xix (what I
_have called the “ trenches’), with an intervening transverse ridge on Xviil.

The female pores (or pore) are doubtless situated in a transversely oval area in
front of the setal zone of segment xiv, which extends from a point between a and b
on one side to a corresponding point on the other.

232 Memoirs of the Indian Museum. [Vor. VII,

The ventral surfaces of segments vill and ix are thickened, as in the typical form ;
these thickenings show each a pair of shallow depressions, approximately in the situa-
tion of the ventral pairs of setae (which however are not to be seen), and connected
each with its fellow across the middle line by an irregular shallow trench. The sper-
mathecal pores are in the depressed areas, rather in front of the middle of the seg-
ment, in the line of setae a, or between a and b.

Internal Anatomy :—Septum 4/5 is moderately thick; the next is 7/8, which is
somewhat thickened, as are all as far backwards as 13/14. The thickest of the series
are 10/11 and 11/12; the thickening decreases by degrees in front and behind these ;
there is some slight thickening even as far back as 16/17.

The gizzard is large and ovoid, occupying part of the space between septa 4/5 and:
7/8. There is one pair of calciferous glands, in segment xv, with lobed outer margins.
The intestine begins in xviii.

The last heart is in xii.

The micronephridia are small, numerous, and scattered irregularly.

Testes and funnels are free in segments x and xi; these segments are filled out by
masses of flocculent matter. The seminal vesicles, in ix and xii, are small.

The prostates are much coiled, in xvii and xix; the relatively short duct is thin-
ner than the glandular part, and passes inwards with an irregular bend at its begin-
ning. A number of muscular bands, similar to those described by Michaelsen in O.
surensis (4), are sufficiently prominent to attract attention.

The spermathecae are so exactly similar to those of the previously described
specimen from Poona that no further account is necessary.

The penial setae, 7 mm. long, 18» thick in the middle and 20. at the proximal
end, resemble very closely the second shape found in the type form; the number of
rings of spines is about a dozen.

The copulatory setae are 52 mm. long, and 2or thick in the middle. They have
the same bluntly pointed, slightly swollen and claw-shaped tip as the typical form.
The convex and concave borders of the distal third of the shaft are furnished with
thin serrated ridges (fig. 35), somewhat as if the spines of the typical form were bound
to the shaft each by a delicate web. In addition, on the face of the seta which is
presented to the observer under the microscope, a series of semicircular markings is
seen; but I am not quite clear what these actually represent.

In one, but only one, of the numerous specimens from the garden, an additional
marking was present. This was a large slightly hollowed transverse papilla on the
posterior half of segment xx and anterior half of xxi, extending from between a and b
on one side to a corresponding point on the other, and joining the thickening of the
male field along its anterior border (compare the specimen of the typical form from
Poona).

Remarks :—The species is a very variable one,—in the characters of the pro-
stomium, the external genital markings, the spermathecae, the penial and copulatory
setae, Thus there are three well-marked kinds of spermathecal diverticulum, —simple

1920. | J. STEPHENSON : Oligochaeta from India and E. Persia. 233

and attached to the ectal end, chambered and attached to the ectal end, and simple
and attached (not merely attached but bound down by connective tissue) to near
the middle of the sac. The variations are however so distributed that it is impossible
to describe all the combinations as separate forms; the one that I have named (var.
riparius) is distinguishable on external examination by the marked difference in the

male genital area.
Octochaetus prashadi, sp. nov.

Plate X, figs. 36-38.
Kalvan, near Bombay. 7-vü-1917. B. Prashad. A single specimen.
Sakarwari, on the way to Mahableshwar, W. Ghats. 4-vii-1917. B. Prashad. Two speci-
mens.

"External Characters :—Length 51-61 mm.; diameter 2°5-35 mm. Colour buff,
no pigmentation, no difference between dorsal and ventral surfaces. Segments ca.
150; v and vi biannular, some or all of the rest up to the clitellum triannular.

Prostomium epilobous in varying degrees.

Dorsal pores from furrow 12/13 (there may be a rudimentary pore in I1/I2).

There are some slight variations in the setal relations in the various specimens,
but they are not very different from the following :—In the anterior part of the body
ab = ?aa = ?bc = 3cd, and the same behind the clitellum; in the middle of the body
ab = taa = half bc = $cd. The dorsal interval dd = + of the circumference.

The clitellum is absent, or very faint and indefinite.

The male field shows a quadrangular thickening which includes part of xvi and
extends backwards to the hinder border of xx; laterally it reaches to the line of setae
c. On segments xvii and xix are transverse trench-like depressions, deeper in their
lateral portions, where the “prostatic pores are situated on rounded papillae in line
with setae b. The general aspect of the male area is thus not unlike that of the last
species.

Here again is the same difficulty. One of the specimens from Sakarwari, though
corresponding closely in all other points, and especially in the peculiar penial and
copulatory setae, differs markedly from the other examples in the configuration of the
male field. The thickening is less extensive; it does not get on to segment xx, and
laterally does not reach the line c; there are no transverse depressions. Just possibly
the difference is due to the animal not being sexually as advanced as the others.

The female pores are perhaps represented by a pair of small whitish dots near the
middle line on segment xiv, and nearly at the middle of the length of the segment.

The ventral surfaces of segments viii and ix are thickened, especially, it may be,
round the sites of setae a and b; these setae may not be visible, or they may be seen,
rather closer together than usual and shifted forwards nearer the anterior border of
the segment. The spermathecal apertures are in furrows 7/8 and 8/9, conspicuous,
in line with setae b, or between a and b.

Internal Anatomy :—Septum 4/5 is somewhat thickened. 5/6 is thin, and 6/7 absent.
There is then some thickening as far as the clitellar region, most marked, perhaps, in
septa 9/10 and 10/11, and decreasing in front of and behind these.

234 Memoirs of the Indian Museum. [Vor. VII,

The gizzard is in front of septum 7/8, relatively large, globular, but not very firm.
There is a large calciferous gland on each side in xv, projecting backwards also into
xvi; each is divided into an anterior and a posterior lobe, and on the whole is kidney-
shaped ; the posterior ends approach each other, and are at a higher level in the seg-
ment than the anterior. The intestine begins in xvii or xviii.

The last heart was in segment xili twice, in xii once.

The micronephridia are numerous and small.

_ Testes and funnels are free in segments x and xi; the funnels are of large size,
or at least the iridescent mass which adheres to them is. The seminal vesicles are in
segments ix and xii; they are slightly lobulated; both pairs may be of moderate size,
or those in 1x may be much larger than the posterior pair.

The prostates, in segments xvii and xix, are relatively large, and bulge apart the
septa of the containing segments. The glandular part is a rather thick and opaque
tube, closely coiled; the duct is much thinner, shiny, passing inwards with a bend or
loop at its origin, or with a wavy course.

Ovaries and funnels occupy the usual situation; there were in one specimen
minute empty ovisacs in segment xiv.

The spermathecal sac has a very stout duct which is not sharply marked off; the
diverticulum is of considerable relative size, and has a very thick stalk; it arises from
the spermathecal duct where the latter enters the body wall (fig. 36). So much can
be said of all three specimens; but all three differ in details. The ampulla may be
irregular in shape, or simply ovoid; the diverticulum may be almost as long as the
main sac, or considerably shorter; it may be quite simple, or it may be slightly lobed,
and on microscopical examination a few small seminal chambers may be distinguish-
able.
The penial setae (fig. 37) are in length 1°5 mm., and 404 thick in the middle;
the tip is slightly hooked, and rather hollowed or spoon-like on its concave side. The
ornamentation consists of a large number of close-set rings of fine spines, which extend
nearly half way along the shaft.

The copulatory setae are ‘8 mm. long, and 26. thick at their middle. They are
slightly bowed, and the tip is pointed ; the distal portion of the shaft is marked by a
number of scar-like depressions, with a general semicircular shape, and sharply defined,
elevated and notched proximal margin (fig. 38).

Remarks:—In a number of details the present form remarkably resembles the
last. The penial and copulatory setae however are very characteristic. There is no
doubt however that this and the last are closely related species.

Octochaetus montanus, sp. nov.
Plate X, figs. 39, 40.
Panchgani, W. Ghats, 12 miles from Mahableshwar ; alt. 4,000 ft. 5-vü-1917. B. Prashad.
A single specimen.
External Characters :—Length 60 mm. ; thickness 3°5 mm. Colour buff, non-pig-
mented, slightly blotchy in places. Segments 158.

1920. | J. STEPHENSON : Oligochaeta from India and E. Persia. 238

Prostomium epilobous 3, the tongue separated from the projecting lobe in front,
but not delimited behind.

Dorsal pores begin from furrow 10/11.

The setal relations may be expressed as follows :—On segment vii ab = aa = half
bc = cd; behind the clitellum ab = taa = 2bc — half cd; in the middle of the body ab
= taa= be =#cd; dd — almost 3 of the circumference.

The saddle-shaped clitellum takes up nearly eight segments, from near the anter-
ior border of xii to xix inclusive.

The male field is a rectangular whitened area which includes the ventral surfaces
of segments xvii-xix, and extends laterally to just outside the lines of setae b. The
seminal! grooves run longitudinally in line with setae a; the prostatic apertures, at
the ends of the grooves, are not separately visible; the intersegmental furrows are
visible, intersecting the seminal grooves at right angles.

An elongated, transversely oval genital papilla is present behind the male area,
over the situation of furrow 21/22, depressed in its centre, where it covers the anterior
half of xxii and the posterior of xxi; laterally it reaches on each side to the line of
setae b,

Segment xiv presents a whitish pad mid-ventrally, on which the female pores
are perhaps represented by a couple of darkish dots at about the middle of the length
of the segment.

The spermathecal apertures seem to be in furrows 7/8 and 8/9, as a couple of
whitish dots in line with a.

Internal Anatomy :—Septum 4/5 is moderately stout; 5/6 to 7/8 are very thin;
8/9 is somewhat thickened, 9/10 to 11/12 moderately so, 12/13 very slightly so.

The gizzard is in segment vi, of moderate size, squarish. The calciferous glands
are two pairs, in segments xv and xvi, ovoid, antero-posteriorly compressed, and situ-
ated dorsally on the alimentary canal by the side of the dorsal vessel. The intestine
begins in xvüi.

The last heart is in xii.

The excretory system consists of scattered micronephridia.

Testes and funnels are free in segments x and xi. Two pairs of large seminal
vesicles occupy ix and xii, those in xii meeting dorsally above the alimentary canal ;
both pairs are much lobulated,—indeed they might be described as racemose.

The prostates are two pairs, rather small. The glandular part is a rather thick
opaque tube, with a few undulations but generally transverse in direction; the duct
is very small, short and thin, also transverse in direction.

The large ovaries have the usual situation.

The spermathecal ampulla is an irregular sac; its duct is large, stout at its begin-
ning and narrowing towards its ectal end, as long as the ampulla and fully half as
thick in its ental portion. There is a single diverticulum, which arises from the
duct at or above the middle of its length; it is finger-shaped on the whole, slightly
swollen at its blind end, where a few seminal chambers are indistinctly seen (fig. 39).

The penial setae are in length up to 1°5 mm., but very thin,—only 6» in thick-

236 Memoirs of the Indian Museum. [Vor. VII,

ness at the middle. The shaft is rather bowed, and slightlv undulating towards the

tip; it tapers very gradually, and the tip is simply pointed ; there is no ornamenta-

tion (fig. 40). 73
There are no copulatory setae in the spermathecal region. ;

Octochaetus pallidus, sp. nov.
Plate XI, figs. 41, 42.
Panchgani, W. Ghats, 12 miles from Mahableshwar; alt. 4,000 ft. 5-vii-1917. B. Prashad.

Two specimens, one damaged in the clitellar region. :

Mahableshwar, W. Ghats, 5,000 ft. 5-vii-1917. B. Prashad. A single mature specimen.

External Characters :—Length 40-44 mm.; diameter 2°5 mm. Colour pale yel-
lowish, quite unpigmented, no difference between dorsal and ventral surfaces. Seg-
ments 166 ; segments vi-ix indistinctly triannular,—indeed this secondary annulation
extends faintly as far back as the clitellum.

The prostomium differs in the two specimens from Panchgani; in the first it is
prolobous, in the second somewhat epilobous, with a very broad angle behind, which |
is continued back by a median groove through segment 1.

Dorsal pores from furrow 10/11 ; the first is small.

The setal relations may be expressed as follows :—In front of the spermathecae
ab = + aa = half bc = cd; behind the genital segments ab — 4 aa = 3 bc — half cd;
at the middle of the body ab = ? aa = nearly half bc = 3 cd; dd is equal to half the
circumference or rather less at the middle of the body, but at the hinder end is only
about one-third of the circumference.

The clitellum is saddle-shaped, and extends over xiii-xvii (= 5).

The male field is represented in the undamaged Panchgani specimen by a thicken-
ing on the ventral surface of segments xvii-xix, which extends outwards to a little
beyond the lines of setae 6. The prostatic pores are in the line b, and the seminal
grooves lie just outside this line, straight and longitudinal in direction, curving inwards
at their extremities to end at the pores; the grooves are close to the edge of the
whitish area.

In the specimen from Mahableshwar the field was rather more extensive, and
circular in shape, it embraced portions of segments xvi and xx, and reached outwards
toc. The prostatic pores were situated each in a small transverse groove.

The female pores are paired, on minute papillae a little internal to and in front
of the site of setae a.

The spermathecal pores are two pairs, at the site of setae a on segments viii and
ix; they are represented by small round papillae, white at their summits, but
definite apertures were not present.

Internal Anatomy :—Septum 4/5 is thin, 5/6 and 6/7 very slightly strengthened,
7/8-11/12 all somewhat thickened, 12/13 only very slightly so; the rest are thin. —

The gizzard is barrel-shaped, in segment vi; the oesophagus is distinctly streng-
thened in segment v also, where shining longitudinal muscular bands are seen. This
seems to be the beginning of a double gizzard, such as is seen in Eudichogaster,

1920 ] J. STEPHENSON : Oligochaeta from India and E. Persia. 237

Dichogaster, and Trigaster ; i do not however suggest at present that any of these
genera are derived from this species, or indeed from the genus Octochaetus at all.

There are no calciferous glands. The intestine begins in xvi.

The last heart is in segment xii.

The excretory system is micronephridial. In the postelitellar segments the organs
are few in each segment and of moderately large size; they are placed in a transverse
row in each segment, about seven on each side, and in each row they increase in size
from the ventral end of the row as far as the fifth nephridium, but the two most
dorsal are smaller again. This was the condition in the anterior part of the animal,—
in that part usually opened for dissection; on opening the posterior part of the
specimen the difference in size was found to disappear towards the hinder end.

A curious modification of the nephridia, which I have not seen before, was found
in segments xi and xii. In each of these segments a ventrally situated pair of
structures attracted my attention ; these appeared at first to be flattened bags, rather
oval in shape, taking up approximately the whole length of the segment from one
septum to the other, and attached by a stalk at their inner ends to the body-wall
near the middle line. They could be detached from the body-wall except where they
were attached by the stalk; there was nothing to correspond externally. They were
present in the same segments and the same position in the second worm from
Panchgani. Microscopically they were found to be masses of micronephridial tubules,
arranged in a compact series of parallel loops, the loops running transversely to the
long axis of the oval mass.

Testes and funnels were free in segments x and xi, along with much flocculent
matter.

There are two pairs of seminal vesicles, in ix and xii. Both pairs are of moder-
ate size; those in xii are racemose, those in ix have almost smooth borders.

The prostates are two pairs; they are tubular, of moderate size, the glandular
part consisting of a series of apposed loops. The duct is very narrow at its begin-
ning, but soon widens; it makes a complete bend, and then passes inwards and
slightly forwards; it is of some length, straight after the initial bend, stout and
shining.

The ovaries are in segment xiii; ovisacs are present in xiv.

The spermathecal ampulla is elongated, narrower towards its blind end, swollen
near its base, of moderate size; the duct is quite short, and appears dilated, so as to
be subglobular. The diverticulum is single, stalked, rounded, about equal in size to
the duct, to the side of which it is attached; the diverticulum shines with contained
spermatozoa, but is not chambered (fig. 41).

The penial setae (fig. 42) are 79 mm. long, and 7 to 8; thick at the middle.
The shaft is slightly bowed, and tapers gradually towards the tip; the point is fairly
sharp, and there is no ornamentation, but the distal end of the seta has a curious
wavy outline.

Remarks :—The present form appears to be related both to the last (smooth
penial setae), and also to the anomalous and somewhat problematical O. bishambari

238 Memoirs of the Indian Museum. [Vor svat:

which I described some time ago (9); in additional to having smooth penial setae the
latter species has, like the present one, no calciferous glands.

Octochaetus ganeshae, sp. nov.
Plate XI, figs. 43-45.
Ganeshkhind, 4 miles from Poona, 3-vü-1917. B. Prashad. Three specimens.
Londa, 10 miles from Castle Rock, W. India. 6-vii-1917. B Prashad. One specimen, an
anterior fragment, not fully mature.

External Characters :—Length 43 mm.; diameter 2°5 mm. Colour unpigmented,
pale except where matter in the alimentary canal shows through the body-wall.
Segments 150.

Prostomium epilobous to a somewhat varying extent, about 2; sides of tongue
parallel, or (once) converging behind to form a Y-shaped figure.

Segments v and vi biannular, thenceforwards as far as the clitellar region
triannular.

Dorsal pores from furrow 12/13.

Setal ratios may be expressed as follows :—Behind the genital region ab = laa =
2bc = 3cd; in the middle of the body ab = laa = 2bc = 3cd; but apparently
different specimens show slightly different ratios. In front of the genital region the
setae are small and difficult to see. The mid-dorsal distance dd is almost two-thirds
of the circumference.

The clitellum is absent, or not developed in these specimens.

The male field is a slightly raised whitish area,—more raised towards its lateral
margins,—rectangular in shape, including segments xvii-xix and extending out-
wards on each side rather further than half way between the lines of setae b and c.
The prostatic pores are between the lines of a and b, and the seminal grooves are
almost straight, slightly bowed inwards.

The female pores are marked by a pair of minute indistinct papillae anteriorly
on segment xiv and internal to the line a.

The spermathecal apertures are minute slits on segments viii and ix, just in front
of and between the two setae of each ventral couple.

Internal Anatomy :—Septum 4/5 is moderately strengthened; 5/6 and 6/7 are
absent, 7/8 and 8/9 are slightly thickened, 9/10 to 11/12 considerably so, 12/13, 13/14,
and even 14/15 slightly so.

The gizzard is of moderate size, rounded, and situated in the space between
septa 4/5 and 7/8; probably it is morphologically in vi, since there are soft-walled
portions of the canal both in front of and behind it within these limits.

The calciferous glands, in the specimen from Ganeshkhind first examined, are
in xv, of moderate size, confined to this segment, somewhat kidney-shaped, slightly
lobed, attached to the gut by the hilus, and meeting dorsally over the gut. In the
specimen from Londa the glands took up part of two segments, xv and xvi, and were
mainly in the latter; they were deeply incised by the septum,

The intestine begins in xvii or xviii,

1920. | J. STEPHENSON : Oligochaeta from India and E. Persia. 239

The last heart is in xil.

The micronephridia are arranged in a single transverse row in each segment.

Testes and funnels are large and free, in segments x and xi. Seminal vesicles
occupy segments ix and xii; both pairs are of moderate size, and slightly lobed.

The tubular prostates consist of a few coils only. The duct is about half the
thickness of the glandular portion, of the same diameter throughout, soft and
semitransparent, passing with an undulating course transversely inwards.

The female organs have the usual situation.

The spermathecal ampulla is an elongated sac, narrower towards its blind end,
almost sessile on the body-wall, so that a duct cannot be separately distinguished.
There is a single diverticulum, which may appear simple, or may show a few small
lobulations; it is small, and attached by a short stalk to the base of the ampulla
where this joins the body-wall (fig. 43).

The penial setae (fig. 44) are ‘42 mm. long (possibly longer when fully developed),
and rop thick at the middle. The shaft is almost straight, slightly bowed towards
the distal end; the tip is pointed and slightly hooked; the ornamentation consists
of a few circles of fine spines near the tip, principally visible as fine irregularities of
the edges of the seta.

The copulatory setae of the spermathecal segments are in length ‘27 mm., but
here also the full length may not have been attained; they are 104 thick at the
middle. They are not very different in type from the penial setae; the shaft is
straight for the most part, slightly bowed towards the tip, which is bluntly pointed
and somewhat claw-shaped. The ornamentation consists of a number of fine spines
on the convex and concave borders of the terminal portion of the shaft; in the
specimen from Londa a few incomplete rings were seen (fig. 45).

Octochaetus pachpaharensis, sp. now.
Plate XI, figs. 46, 47.
Pachpahar, about 40 miles S. of Kotah, Rajputana. 13-vii-1917. DB. Prashad. Five

specimens.

External Characters :—Length 28 mm., diameter I mm. Unpigmented. Seg-
ments 05.

Prostomium broad, slightly epilobous, tongue not cut off behind.

Dorsal pores from furrow 7/8.

Setal relations not easy to determine in such a small worm,—perhaps in general
ab =? aa = 2 bc = 3cd; dd is slightly less than half the circumference.

The clitellum extends over segments xiii— 3 xviii (= 45). It is saddle-shaped
except on xiii or xii and xiv, where it is complete.

The prostatic pores, and the straight seminal grooves, are between the lines
a and b.

The female pores are apparently paired, on the anterior part of segment xiv.

The spermathecal apertures were not visible externally; on dissection they
appeared to be in grooves 7/8 and 8/0.

240 Memoirs of the Indian Museum. [Vor. VIT,

Internal Anatomy :—Septum 5/6 is somewhat thickened, 6/7 considerably, 7/8,
8/9 and 0/10 much thickened, considering the size of the worm ; 10/11 to 13/14 are
also somewhat strengthened.

There is a rudimentary gizzard in segment vi, of fair size, barrel-shaped, but
soft ; it appears as shining longitudinal bundles of muscular fibres. The large lobes
of the pharyngeal glands occupy much space in segment v. There are no calciferous
elands. The intestine begins in xiv.

The last heart is in xii.

The excretory system is micronephridial. Behind the genital region there are
three nephridia on each side in each segment, with the form of flattened coils (not the
flattened plate-like organs of Dichogaster, where similarly there are only a few
nephridia on each side in each segment). In front of the prostatic region there are
even fewer, perhaps only one on each side in some segments ( ? ).

Testes and funnels are free in segments x and xi (judged from the flocculent
masses in these segments, which pass deep in the segments into the iridescent covering
of the funnels ; testes were not separately identified), Seminal vesicles are present
only in xii, as rounded masses which may meet dorsally over the alimentary canal.

The prostates are two pairs, tubular, of fair length and sometimes extending
beyond their own segment, bent several times. The duct is much thinner than the
glandular portion, almost straight, and shining.

Ovaries are present in segment xiii, and ovisacs in xiv.

The spermathecal ampulla is an exceedingly irregular and deeply lobed sac
(fig. 46) ; the duct is as long as or longer than the ampulla, but relatively narrow, of
the same diameter throughout, firm and shining. What I think is the diverticulum
(since in one specimen it contained iridescent spermatozoa, which the main sac never
does) is a saccule attached to the ental end of the duet, much resembling one of the
lobes into which the ampulla is ineised.

Two spermathecae were found on the right side in segment vili, opening side by
side; this is presumably an individual anomaly.

The penial setae are bent into about two-fifths of a circle, and measure across the
bend ‘7 mm.; the thickness at the middle is 12«, but at the proximal end 204 The
shaft tapers gently towards the distal end, which does not quite continue the direction
of the curve, but is slightly bent in the contrary direction. The tip is somewhat
wavy, and ends in a fairly sharp point. The ornamentation consists of a number of
irregular rings of small teeth on the distal portion of the shaft, the terminal portion
however being smooth.

Remarks :—The absence of caleiferous glands, with the small number of nephridia
per segment, allies this form to the group of pallidus and bishambart.

Genus Eutyphoeus.

Eutyphoeus incommodus (Bedd).

Bharatpur, E. Rajputana. 15-vii-1917. B. Prashad. Two specimens,

1920. | J. STEPHENSON : Oligochaeta from India and E. Persia. 241

Eutyphoeus mohammedi, Stephenson.
Rawal Pindi, N. Punjab. June-July 1917. R. Hodgart. A single specimen.

This worm has been recorded only once previously, from Allahabad (9). I add a
few notes on the external characters of the present specimen; the internal anatomy
corresponds exactly with that of the type.

The length of the present example was only 39 mm.; its maximum diameter,
behind the clitellum, where it was perhaps unnaturally swollen, was 4°75 mm. ; the
worm is thus short and stout. Segments ca. 160, all very short behind the clitellum.
The prostomium showed no projecting lobe,—perhaps this was withdrawn within
the buccal cavity; a couple of parallel grooves on the dorsal surface of the first
segment indicated that it was tanylobous.

The setal intervals were as follows:—ab = laa = 2be = ÿcd ; but in front of the
clitellum the mid-ventral distance aa becomes much smaller,

The male pores were situated on distinct papillae.

Eutyphoeus chittagongianus, Mchlsn.

Above Tura, Garo Hills, Assam ; 3,900 ft. On paths after rain. July-Aug. 1917. S. W.

Kemp. Two specimens W !7—, W 17°.
Above Tura, Garo Hills, Assam ; 3,900 ft. July-Aug. 1917. 8. W. Kemp. Two specimens

W 161, W 198,
Sureil, Darjiling Dist. ; 5,000 ft. 11-31-x-1917. N. Annandale and F. Gravely. A single

specimen.
Nam Ting Pokri, Darjiling Dist. ; 4,600 ft. June-July 1918. $8. W. Kemp. A single speci-

men.

This worm has been described by Michaelsen (3) from two specimens in a bad
state of preservation. I therefore add a few notes to supplement the original
account.

External Characters :—The length is various, from 182 to 405 mm. ; the maximum
diameter is as much as 10 mm. The colour may be a medium olive dorsally, lighter
below ; or the worm may be almost unpigmented throughout.

Secondary annulation is well marked on the anterior segments; iv and v are
biannular, vi has two chief and two subsidiary furrows, and succeeding segments as far
as the clitellum are primarily triannular, with secondary furrows on the first and last
annuli; the post-clitellar segments may also be triannular.

Detae seem to be sometimes absent on the first four or five segments.

The male pores, each sunk in a separate pit in Michaelsen’s specimens, are often
contained in a large transverse furrow which extends across the ventral surface ; this
furrow has rounded but not tumid lips, and on looking into it the male pores are
seen on small transversely oval papillae in the line of setae b.

The spermathecal apertures, in b in Michaelsen’s original specimens, and in some
of mine, may be between b and c.

The genital markings, described in the original account as “ intersegmental

242 Memoirs of the Indian Museum. [Vor. VII,

areas,” are seen in better preserved specimens to be clean-cut depressions, mainly on
the posterior annulus of the anterior of the two segments with which they are in rela-
tion. These may occur also on furrow IO/II.

In some of my specimens there is a tendency for the post-genital depressions to
divide into two (as in H. kempi; see below, Remarks); and low flat papillae may be
present within some of the depressions.

Internal Anatomy :—The dorsal vessel, as in many species of the genus, does not
extend forwards to the anterior end of the body; it gives off two pairs of lateral
commissures in front of septum 8/9, and is not traceable beyond this,—it does not
get on to the gizzard.

The micronephridia behind the genital region are arranged as a transverse row of
small organs in each segment just behind the attachment of the septum ; from the
prostatic region forwards they are numerous and irregularly scattered.

Testis sacs, about which Michaelsen was in some little doubt, are present, large
and subspherical, in xi, lying against septum 10/11; they are contiguous, and ap-
parently communicate with each other, beneath the alimentary canal.

The spermathecal duct is practically absent, the ampulla being sessile on the
body-wall and attached by a portion of its under surface; if a duct is described, it
would be very short and stout. The broad fan-shaped diverticulum is often divided
into two ;—not always, and when undivided it may appear like a flange surrounding
the greater part of the attachment of the ampulla to the body-wall. The number of
seminal chambers appears to be about twenty.

The penial setae vary in length from 2 to 5 mm., and in thickness from 30 to 404;
some of the difference in length may be due to the growth of the shorter not having
been completed. The extent of distribution of the small teeth on the distal end
varies. Even when fully formed setae were projecting from the male pores, the dis-
tal ends seemed to be often soft and perhaps deformed; Michaelsen had the same
difficulty. The typical form of the tip seems to be broadened and perhaps spoon-
shaped.

Remarks :—I have come to the conclusion that #. kempi, which I described from
the Abor country (8), must be identified with the present form. The difference in
form and distribution ot the genital markings, and in the penial setae, are not suffi-
cient to justify its separation, when one takes into consideration the variations that
are now known to occur.

It might be justifiable to separate the specimens from Sureil and Nam Ting
Pokri as a distinct variety, on account of (i) absence of pigment, (ii) spermathecal
apertures midway between b and c, (iii) genital markings as pairs of oval depressions,
those of a pair being contiguous in the middle line in furrows 19/20, 20/21, and 21/22,
(iv) the tip of the penial setae free from ornamentation and smooth, while on the
other hand, the minute spines extend further up the shaft than in the typical form.
But the forms I previously described as #. kempi show an intermediate condition as
regards (ii) and (iii), and in them too the tip of the penial setae is free from spines,
though these do not extend far up the shaft.

1920. | J. STEPHENSON : Oligochaeta from India and E. Persia. 243

Eutyphoeus waltoni, Mchlsn.

Delhi. 15-vii-1917. B. Prashad. Eight specimens.

Gwalior, Central India : on the bank of a stream. 17-vi-1917. B. Prashad. Four specimens.

Ahmedabad, Bombay Pres. 11-vii-1917. B. Prashad. Numerous specimens.

Baroda, W. India. 9-vii-1917. On banks of Vishvamitri River. B. Prashad. Two speci-
mens, one immature.

Same place; in a garden. 9-vii-1917. B. Prashad. Several specimens, all or mostly imma-
ture.

Same place ; by the side of a tank. 9-vii-1917. B. Prashad. Four specimens.

Same place; on a small hillock. 10-vii-1917. B. Prashad. Five specimens.

Same place ; in a garden. 10-vii-1917. B. Prashad. A number of specimens, mostly imma-
ture.

Navli, between Baroda and Ahmedabad. 10-vii-1917. B. Prashad. Four specimens, imma-
ture.

Caleutta ; banks of Hugli River, in partly saltish water. 23-viii-1918. B. Prashad. Four
specimens, not fully mature.

The species is common in India, and is already well known. The present large
number of specimens, of varying degrees of maturity, has led me to the conclusion that
E. bengalensis Mchlsn. (3) has no separate existence, and is only an immature form
of E. waltom. In going over the above batches of specimens I had at first no suspi-
cion of this, and diagnosed those from Delhi, Navli, and three of those from. Baroda
as H. waltoni, while T put down the worms from Gwalior, Ahmedabad, and two of the
batches from Baroda as E. bengalensis ; I began to have doubts however during the
progress of the work, and with regard to one of the batches from Baroda, of worms
of varying degrees of maturity, I could feel no certainty. The Calcutta specimens I
at first thought to be a new species. It seems to be the case that H. waltoni produces
its penial setae early, and that these may be well developed before the clitellum and
characteristic genital markings show themselves.

I may perhaps be allowed to discuss shortly the differences between the two
species, according to Michaelsen’s original descriptions (3).

E. waltoni would appear to be a much larger worm; the lower limit of length
for E. waltoni, however, is not very different from the length of Michaelsen’s single
specimen of #. bengalensis (waltoni 90-230 mm., bengalensis 72 mm.—hy a misprint
given as I2 mm.).

The prostomium of waltoni is tanylobous, of bengalensis prolobous. I have pre-
viously described the prostomium of waltoni as combined pro- and tanylobous,—
tanylobous with a transverse groove in front of the tongue; Michaelsen does not
distinguish this form of prostomium from the typical tanylobous, without the trans-
verse groove. In one of my specimens which I put down as bengalensis without
hesitation, the prostomium was tanylobous.

As may be seen from a number of the species referred to in previous pages of
this paper, small differences in the setal ratios are of no importance; I may mention
however that similarly in one of the present series of specimens, which I had no
doubt about identifying as bengalensis, the ratios were exactly those given for waltont.

244. Memoirs of the Indian Museum. [Vor. VII,

Michaelsen does not mention the clitellum of bengalensis, and it was presumably
absent. This I take to be a sign of immaturity. The same may be said of the
absence of the genital markings.

The original account of bengalensis has “testes and sperm-duct-funnels free (?)”’;
my note on the specimens which I did not doubt, at the time, were bengalensis runs
“I think sperm-sacs ; the membrane is certainly very thin, but to the best of my
observation it exists.” HH. waltoni has a common sperm-sac; so that there is pro-
bably no difference here.

One chief difference is that the penial setae of waltoni are beset with minute
spines near the tip, while those of bengalensis are smooth (the shape is much the
same). I have already noted (9) that I could not identify distinct spines even with
the oil immersion lens in some specimens of waltont ; and in the present series, among
those which are undoubtedly waltoni, I have twice found the spines absent from the
spoonshaped tip, though a few were seen further up the shaft.

Lastly the spermathecal diverticula are said to be on opposite sides of the duct
in bengalensis, side by side in waltoni ; in some of the present series of waltoni how-
ever I have found them opposite each other.

Eutyphoeus turaensis, sp. nov.
Plate XI, figs. 48, 49.
Above Tura, Garo Hills, Assam ; 3,500-3,900 ft.; in rotten wood. July-Aug. 1917. S. W.

Kemp. Four specimens, one somewhat damaged. W 1x.

Same place, date, and collector. Under bark. Three specimens, immature. W 175,

External Characters :—Length 100 mm.; maximum diameter 3°5 mm. Colour
light grey, unpigmented; no difference between dorsal and ventral surfaces. Seg-
ments 171; segments long in front of, short behind the male apertures. Segments
v and vi biannular or indistinetly triannular, vii triannular, viii and ix triannular
with other secondary furrows also, x triannular, xi indistinctly so.

Prostomium small, tanylobous, tongue broader behind than in front.

Dorsal pores from furrow 11/12.

Setae are scarcely visible on segments ix and x, while they are enlarged on iil.
vi. The intervals may be expressed thus:—Anteriorly ab =4aa=half be=äcd; behind
the clitellum ab=taa=2bc=3cd; in the middle of the body ab=taa=half be=jcd ;
dd is nearly equal to two-thirds of the circumference.

The clitellum is very slightly marked; all that is to be seen is some thickening
ventrally on segments xv to xvii.

The male pores, on segment xvii, are in a pair of narrow transverse depressions
which extend over an interval greater than ab, the limits of the depressions being
inside the line a and outside b; penial setae, marking the site of the aperture, pro-
trude in the line b.

The female apertures were not visible.

The spermathecal apertures are in furrow 7/8, in the line b.

1920. | J. STEPHENSON : Oligochaeta from India and E. Persia. 245

Genital markings are faintly visible in furrows 14/15, 15/16, and 16/17, in line
with a, as inconspicuous slightly pigmented spots surrounded by circular grooves.

Internal Anatomy :—Septa 4/5 and 5/6 are moderately stout; the next is 8/0,
which is somewhat displaced backwards; this septum and the two following are
rather close together, and are all somewhat strengthened ; 11/12 is absent, as in #.
waltoni and many others of the genus ; 12/13 is very thin, and is bulged backwards
to the level of 14/15 by the seminal vesicles ; the rest are also thin.

The gizzard is barrel-shaped, situated in the long space between septa 5/6 and 8/0.
Calciferous glands are represented by an ovoid dilatation of the alimentary canal in
segment xii, with transverse vascular striations. The intestine begins in xv.

The last hearts are in xiii; those of xi are bound down by connective tissue to
the walls of the oesophagus. The dorsal vessel seems not to be continued forwards
beyond the gizzard, ending by giving a pair of commissures at the anterior end of
this organ.

The micronephridia are arranged in a transverse row in each segment behind the
clitellum ; in the dorsal half of each segment there are pretty regularly three on each
side.

The testis sacs are one pair, in segment xi; they are separate from each other
and longitudinally elongated, lying by the side of the nerve cord. They are connec-
ted with the seminal vesicles in segment xii; these are large, two-lobed, the lobes being
anterior and posterior; each lobe is subdivided by further indentations; they extend
back to the level of 14/15 by bulging back the septa.

The prostates are small, occupying segments xviii and xix; the coils are closely
packed. The duct is not markedly different from the glandular part; it is scarcely
narrower, is soft, not shining, and passes forwards and inwards with an undulating
course.

The female organs were not identified.

The spermathecae (fig. 48) are elongated sacs lying longitudinally on the body-
wall, to which they are attached by a portion of their under surfaces; a duct can
thus hardly be described,—it would at any rate be called very short and stout. The
diverticula are two, one on each side, attached to the junction of sac and duct by a
short relatively stout stalk; each shows a lobulation which indicates the presence of
three or four seminal chambers.

The penial setae (fig. 49) are in length up to. I°5 mm., and in thickness 35, ;
the distal half is slightly curved, and the tip bluntly pointed and rather claw-like.
The ornamentation consists of very fine dot-like markings over the distal eighth or
tenth of the length of the shaft, including the tip.

Remarks :—This species appears to be allied to #. chittagongianus ; the internal
anatomy is remarkably similar, and the chief difference between the two is in the
genital markings. Although the present specimens were possibly not quite mature,
it seems fairly safe to say that the markings differ in both character and position in
the two forms,

246 Memoirs of the Indian Museum. [Vor. VII,

Subfam. TRIGASTRINAE.
Genus Eudichogaster.
Eudichogaster ashworthi, Mchlsn.
Plate XT, figs. 50, 51.

I have lately received, by the kindness of Dr. J. H. Ashworth, two specimens
of this species, of the same batch which furnished the types of the species described
by Michaelsen (2). I need only add very few notes on the peculiarities exhibited by
these.

The papillae of the spermathecal pores are not always symmetrical; in the speci-
men to which I devoted most attention, the papilla on the right side of segment viii
took up about the middle two-fourths of the segment, that on the left side the anterior
half; those on ix took up the anterior two-fifths of the segment, and encroached
somewhat on the intersegmental furrow.

The gizzards in segments v and vi are short, and do not include the whole length
of the oesophagus in these segments; a soft ring is thus left between the two, and
another between the second and the hinder limit of its segment.

The specimen differed from Michaelsen’s in the seminal vesicles; instead of
vesicles in ix and xii, I find a pair in xii, a pair in x attached to the posterior septum
of the segment, and a single vesicle in ix, on the left side only (one specimen only
dissected).

I think Michaelsen’s paper contains a slip where he speaks, in the diagnosis and
again in the detailed description, of the diverticulum of the spermatheca entering
the distal end of the duct. Michaelsen always uses the word distal to mean “nearer
the surface of the body”? (in such a case as the present, when speaking of an internal
organ); the diverticulum however enters the duct at the ental end of the latter, just
below the ampulla, and the same is the case in the numerous specimens to which
reference is made below. '

The copulatory setae are, according to Michaelsen, found on the papillae of the
male field (but not in connection with the prostatic apertures), and doubtfully in the
neighbourhood of the spermathecal apertures; they are of a well defined type. In
the specimen which I dissected I found them in the neighbourhood of the spermathecal

! There is a difference in the use of the words “ proximal” and “distal” by English and German
writers. I was taught to use them for ‘‘ nearer to’ and “ remoter from, the fixed point of attachment”’ ;
and Beddard, for example, among writers on this group, uses the words in the same sense. Thus in
such an organ as the spermatheca, which is attached by its duct to the inner surface of the body-wall
and projects freely inwards into the coelom, a diverticulum attached to the duct near the body-wall
would be proximal, and one attached to the duct near the ampulla would be distal as compared with
the other. So for example in a well-known elementary textbook it is said that the testes of Hydra are
distally situated, 7.e. near the oral and distant from the fixed end of the animal. Michaelsen not in-
frequently queries Beddard’s use of the words, and himself employs “‘ proximal ”’ and “ distal” to signify
respectively “nearness to” and “ remoteness from, the central axis of the body.” In view of this con-
fusion I have for some time past used “ectal” and ‘“‘ental’’ to signify ‘‘nearness to the surface of the
body ” and “nearness to the internal end,” in the case of such structures as those under discussion.

1920. | J. STEPHENSON : Oligochaeta from India and E. Persia. 247

pores, but not in the male region; for ready reference they are shown in fig. 50.
From my examination of the specimens which are referred to below, I do not doubt
that when special setae are found near the spermathecal pores they are of this type;
but I have never found setae of this form in the neighbourhood of the male field,
though other more slightly modified forms are not uncommon there.
Choral, between Khandwa and Indore, Central India. 23-vi-1917. B. Prashad. One sexual
specimen and others smaller, doubtfully of the same species.
Saugor, Central Provinces. 20-vi-1917. B Prashad. A single specimen.
Bina, Central Provinces. 19-vi-1917. B. Prashad. Numerous specimens, mostly immature,
only three with sexual marks.
Teor, 8 miles from Jubbulpore, Central Provinces. 22-vi-1917. B. Prashad. A single speci-
men.
Partabgarh, S. Rajputana. Feb.1918. Col. J. Manners-Smith. Two specimens. one immature,
the other scarcely fully mature.
Poona, W. India. 3-vii-1917. B. Prashad. Several specimens, apparently mostly not fully
mature.
Wahi, on the way to Mahableshwar. W. Ghats. 4-vii-1917. B. Prashad. Six specimens,
two with sexual marks.

Some of the above specimens appeared at first to be worthy of separation as
distinct varieties, on the ground of differences between them and Michaelsen’s des-
cription,—especially as regards the copulatory setae of the spermathecal segments and
of those of the neighbourhood of the male field, joined with the difference in the at-
tachment of the spermathecal diverticulum. Buta careful comparison of the specimens
among themselves, and the examination of the co-types of the species, has shown me
that the differences are not of great importance.

The majority of the specimens encountered are not fully mature; the clitellum
is often wanting, even when all other marks of maturity are present. The length is
often much less than that of the original examples ;—lengths of 45, 56, 67 and 75 mm.
were met with, in each case the best developed of the batch being measured.

The first segment may be divided by a median longitudinal furrow behind the
prostomium.

The dorsal pores may begin in 11/12 instead of 12/13.

The papillae on or near the male field are variable. Those on xvi, or those on
xx, may be absent; and I have not seen the paired papillae on segments xvii and xix
internal to the male pores; there may however be median papillae on these segments.

Similarly with the papillae of the spermathecal region. In addition to those des-
cribed by Michaelsen, there may be a pair on segment vii similar to those on ix outside
the line of setae b, and a median papilla, it may be of considerable transverse extent,
on x.

The seminal vesicles are in ix and xii, or in xii only; in this latter case it is pos-
sible that a second pair would have developed at a later stage. The prostatic duct
is often straight, or only slightly bent; but the explanation may be the same.

The spermathecal diverticulum is in all cases attached to the ental end of the
duct, or in one specimen to the base of the ampulla.

The copulatory setae of the spermathecal region, when found, are of the form

248 Memoirs of the Indian Museum. [Vor. VII,

already mentioned. They are always longer than the measurement given by Michaelsen
(who gives ‘24 mm.),— usually about ‘5 mm., and not less than ‘4 mm.

The modified setae in the neishbourhood of the male field are not always, accord-
ing to my experience, to be found. They are much less modified than those of the
spermathecal region, and may even still retain the ordinary double curve with the
nodulus; the sculpturing consists of a number of fine semicircular lines, concave
towards the tip of the seta, or of a few faint transverse rows of closely set dots, or
of zigzag lines which indent the surface of the seta sufficiently to produce irregu-
larity of the margins (fig. 51).

Eudichogaster bengalensis, Mchlsn.
Bheraghat, Marble Rocks, 13 miles from Jubbulpore, Central Provinces. 22-vi-1917.

B. Prashad. A single specimen.

The present species has been described by Michaelsen (4). As this is the second
time it has been encountered, a few notes may be added to supplement the original
account.

The prostomium I should describe as proepilobous, with a pair of shallow grooves
dorsally on segment i; the grooves extend back over three-quarters of the segment,
but not as far as the intersegmental furrow.

The first dorsal pore I found in furrow 11/12.

My specimen showed a series of papillae in the spermathecal region,—three pairs,
on segments viii, ix, and x, oval in shape, white, including and extending beyond the
setae ab both inwards and outwards.

The last heart is in segment xii.

As the nephridial system has been shown by Michaelsen to be of importance in
this genus, and as his specimens were considerably softened and so not favourable
for examination, I give the results of my own dissection. The individual microne-
phridia are large and few; two longitudinal series are to be seen on each side of the
body in and behind the prostatic region ; 7.e. there are only two on each side of each
segment. At about one-third of the length of the animal from the posterior end
the two are still present, and similar in appearance; though the outer, which extends
on the body-wall from the site of seta c to half-way between d and the middorsal
line, is larger than the inner, which only takes up a space equal to the interval be.
A little behind this the inner becomes more opaque and rather bulkier than before,
and so more conspicuous. Towards the hinder end it maintains these characters, and
increasing in size also so as to include the interval a to c, it becomes much more con-
spicuous than the outer nephridium, which has rather diminished in size. [think there
may be a few very minute micronephridia on the body-wall in addition to the series
described above, but the condition of the specimen is none too good, though appar-
ently better than Michaelsen’s.

The penial setae are not ornamented so markedly as in the type. The spines are
extremely fine, and are arranged in short transverse rows; there is no specially pro-
jecting brush-like circle near the tip.

1920. | J. STEPHENSON : Oligochaela from India and E. Persia. 249

Eudichogaster trichochaetus, sp. nov.
Plate XI, fies. 52, 53.
Bombay, Victoria Gardens. 30-vi-1917. B. Prashad. Numerous specimens, some immature.
Bombay, near Colaba Railway Station. 1-vii-1917. B. Prashad. Seven specimens.
Bombay. under a tree near the Fort. 30-vi-1917. B. Prashad. Three specimens.
Palchar, N. of Bombay, between Bombay and Surat. 7-vii-1917. B. Prashad. A number
of specimens. Two showing sexual marks.

External Characters :—Length 32-45 mm.; diameter 1°75-2'25 mm. Colour a
yellowish grey, no difference between dorsal and ventral surfaces. Segments 103-128.

Prostomium epilobous 2/5, pointed behind, the point continued back as a groove
which divides the dorsal surface of the first segment; or in some cases the prosto-
mium itself appears to be prolonged backwards so that the point is at the furrow 1/2.

Dorsal pores begin from furrow 12/13 or 13/T4.

The ratios of the setal intervals may be expressed thus:—ab=} to 2aa=4bc
—2cd from behind the male genital region backwards; in front of the clitellum the
ratios are less regular, but ab is certainly wider, equal about to half aa; dd is nearly
half the circumference.

I have never found a recognizable clitellum.

The male field is a rectangular thickening, whitish in colour, taking up segments
xvii-xix, and extending laterally on each side to between the lines of b and ce; it
may extend slightly on to segments xvi and xx. The prostatic apertures are small
transverse slits corresponding in extent to the interval ab, on segments xvii and xix.
The seminal grooves pass longitudinally between the outer ends of the prostatic pores
of the same side, and are thus in the line 6, and some little distance internal to the
lateral border of the thickened area.

The female pores may be indicated by a pair of tiny white thickenings just in
front of, and internal to setae a on segment xiv.

The spermathecal pores were not seen externally. There is some thickening ven-
trally on segments viii and ix, indefinite, and perhaps more specially marked round
the site of the ventral setal couple.

Internal Anatomy :—Septum 4/5 is somewhat strengthened, 5/6-7/8 are thin, 8/9
is somewhat strengthened, 0/10 slightly so, the rest thin.

The two gizzards are relatively large, in segments v and vi. The calciferous glands,
three pairs, in segments x, xi, and xii, are not set off from the oesophagus. The in-
testine begins in xiv (?).

The last heart is apparently in xii.

The micronephridia are arranged in four longitudinal rows on each side of the
body; they are small twisted tubes, those of the innermost series on each side the
smallest, and situated near the next outer row.

Testes and funnels are apparently free in segments x and xi (judged from the
masses of free flocculent material in these segments). Seminal vesicles are present in
segment xii only, as considerable masses with lobate margins.

250 Memoirs of the Indian Museum. [Vor. VII,

The prostates are two pairs of relatively moderate size, twisted tubes lying in
segments xvil and xix.

The spermathecae are two pairs, in segments vill and ix, ending apparently on
the body-wall between the site of setae a and b. The ampulla is ovoid, rather
elongated and with a somewhat pointed blind end, the duct as long as the ampulla,
but not distinctly marked off, a little wider above. The diverticulum is single,
shortly finger-shaped, one-third as long as the ampulla, to the lower end of which it is
attached.

The penial setae are remarkable. In length they measure up to 2 mm.; but
notwithstanding this they and their sacs were quite invisible in the dissection, and the
prostatic duct was pulled out and mounted merely on the chance of getting something
(the setal sacs are practically always identifiable, and can usually be isolated and re-
moved separately ; but in some cases, such as the small Dichogasters, the only practic-
able way is to remove prostates and setal sacs together). The setae are very slender,
5-6» in thickness, diminishing to 4:5» near the tip. In shape they are undulating;
there is no ornamentation, but where the tip is best seen it appears bifid, with a thin
transparent web between the prongs of the fork; the base of the expanded portion
is Qu, and the length of the triangular web 18» (fig. 52).

The copulatory setae of the spermathecal region are ‘42 mm. in length, and 13,
thick in the middle. The shaft is almost straight, with a bend at the proximal end
however; the tip is slightly claw-shaped and bluntly pointed. The ornamentation
consists of short transverse ridges on the distal part of the shaft (fig. 53).

Eudichogaster prashadi, sp. nov.
Plate XI, fig. 54.

Palia, between Indore and Ujjain. 27-vi-1917. B. Prashad. Several specimens, only two
with sexual marks.

Indore. Central India. 23-vi-1917. B. Prashad. A single specimen.

Mhow Cant., Central India. 23-vi-1917. B. Prashad. A single specimen.

Khandwa, Central Provinces. 23-vi-1917. B. Prashad. Several specimens.

Bheraghat, Marble Rocks, 13 miles from Jubbulpore. 22-vi-1917. DB. Prashad. Several
specimens. ;

Teor, 8 miles from Jubbulpore, Central Provinces. 22-vi-1917. B. Prashad. Several speci-
mens, mostly immature.

Saugor, Central Provinces. 20-vi-1917. B. Prashad. Numerous specimens.

Surat, W. India. 8-vü-1917. B. Prashad. Several specimens.

Poona, W. India; on the banks of the Rivers Mula and Mutta. 3-vii-1917. B. Prashad.
Four specimens, only one showing distinct signs of maturity.

External Characters :—Length 35-67 mm., diameter 3-4°5 mm. Colour yellowish
brown, with only slight difference between dorsal and ventral surfaces; the first few
segments lighter. Segments 140-168.

Prostomium prolobous; there may be a mid-dorsal groove on the first segment,
sometimes with an irregular course.

1920. | J. STEPHENSON : Oligochaeta from India and E. Persia. 251

Dorsal pores from furrow 11/12 or 12/13.

The setal intervals vary somewhat; ab may be anything from 4 to 4 of aa,

and over the greater part ot the body=about 3 be and #cd: in front of the male
apertures be becomes rather smaller, and cd increases; the interval ab is smallest im-
mediately behind the male field. The mid-dorsal interval dd is about 2 of the
circumference. Towards the anterior end the setae are difficult to see.

I never saw a distinguishable clitellum.

The appearances of the male field are also not very definite, but such as they are,
they are pretty constant. The chief feature is the existence on each of the segments
xvii and xix of a pair of ill-defined papillae, perhaps better described as whitish thicken-
ings of the body-wall; these are generally transverse in direction, and have their cen-
tres somewhere near the line b, extending inwards and outwards to a variable ex-
tent ; their limits are rather indistinct. On segment xviii there is usually a similar
thickening which unites the outer parts of the thickenings on xvii and xix, thus mak-
ing a crescentic swelling with its concavity inwards on each side (fig. 54a).

The prostatic pores are in or just internal to the line of setae b, on segments xvii
and xix; the seminal grooves bend inwards slightly at the middle of their length, some-
what asin H. ashworthi. I saw the pores of the vasa deferentia in one specimen, on
Xvill in b, at the apex of the inward bend in the course of the seminal grooves; their
anterior and posterior lips were slightly tumid.

The female pore or pores are perhaps contained within a minute white spot mid-
ventral and anteriorly situated on xiv.

There are small transversely elongated white cushions on segments viii and ix,
in the position of the ventral setal bundle. From internal dissection the sperma-
thecae appear to discharge between the sites of setae a and b on these segments.

Internal Anatomy :—Septum 4/5 is thin; 5/6 to 9/10 are moderately strengthened,
10/11 slightly so, 11/12 still less, and the rest are thin.

The gizzards, large, rounded, and firm, are in segments v and vi; they are
separated by a soft-walled part of the oesophagus in the anterior part of segment vi.
The calciferous glands, in segments xi and xii, are shortly ovoid, and joined to the
gut by a short stalk on the inner side. The intestine begins in xv.

The last heart is in segment xii.

Behind the genital region the micronephridia are in a transverse row on each
side of each segment, and so arranged that the corresponding organs follow behind
each other in longitudinal rows; of these there are about five on each side of the
body. Towards the hinder end of the body the innermost nephridium of each trans-
verse row on each side,—that by the side of the ventral nerve cord,—increases in size
and thickness, and though still small is much the most conspicuous; the others of the
row become individually smaller and increase in number, and lose the regularity of
the arrangement in longitudinal rows.

Testes and funnels are free in segments x and xi; those in the two segments are
equal in size. Seminal vesicles are present in segments ix and xii; in one specimen

252 Memoirs of the Indian Museum. [Vor. VII,

dissected I found them only in xii, perhaps because the specimen was not fully
mature.

The prostates are two pairs, in xvii and xix, small thin convoluted tubes with a
generally transverse direction. The ducts are of the same diameter as the glandular
part, but a little more shiny in appearance; they lie transversely.

The spermathecae are two pairs, in segments viii and ix; the ampulla is an
elongated ovoid sac, and the duct cylindrical and as long as the ampulla. There is a
single diverticulum, ovoid, not apparently containing any seminal chambers, attached
by a short thick stalk to the base of the ampulla; it is bound down to the duct and.
base of the ampulla by connective tissue.

There are no penial setae.

The copulatory setae of the spermathecal region are not unlike those of E.
ashworthi ; they seem to be always present. In length they measure ‘47 mm., and are
16; thick in the middle; they are almost straight, slightly bowed towards the distal
end; the thickness does not diminish much till close to the tip, which is pointed and
rather claw-shaped. The distal fifth of the shaft is marked by a number of large
hollows scooped out of the shaft, each with a sharply defined and prominent proximal
border, crescentic with the concavity towards the tip; the distal margin of each
excavation slopes gently, and is not well marked (fig. 540).

Remarks :—It is curious that the period of sexual maturity in some of these
worms is so restricted,—that is if the presence of a clitellum is to be taken as a sign
of complete maturity. 5

The present form has much in common with E. indica (Beddard). The latter
however appears to be distinguished by the great separation of the lateral setae
(cd=2}ab), by median genital papillae behind the spermathecal region, and by possess-
ing only one pair of seminal vesicles.

Eudichogaster falcifer, sp. nov.
Plate XI, fig. 55.
Jubbulpore, Central Provinces. 22-vi-1917. B. Prashad. A number of specimens.
Bheraghat, Marble Rocks, 13 miles from Jubbulpore. 22-vi-1917. B. Prashad. A number
of specimens.
Saugor, Central Provinces. 20-vi-1917. B. Prashad. A single specimen.

External Characters :—Length 40 mm.; thickness 2 mm. Colour a nondescript
yellowish grey, no difference between dorsal and ventral surfaces, the anterior end
lighter. Segments 128.

Prostomium proepilobous.

Dorsal pores from furrow 12/13.

The setal intervals may be expressed as follows:—ab in the middle of the body
is rather greater than }bc but rather less than Zaa, and is nearly equal to cd; behind
the genital region ab is about equal to $aa and to Hbc, though aa seems just a little
larger than bc; in front of the genital region ab = à aa = $ bc = # cd; dd is about 2 of
the circumference.

1920. ] J. STEPHENSON : Oligochaeta from India and E. Persia. 253

The clitellum was indistinguishable.

The male field is represented by a very slight whitening and thickening of the
ventral surface of segments xvü to xix. The lateral parts of this area are better
marked, and constitute definite ridges, which turning inwards at their anterior and
posterior ends enclose the centre of the area as within brackets: in the longitudinal
portion of their course the ridges lie a little outside the line b.

The seminal grooves are crescentic, the convexity outwards; they begin and end
in the position of seta a on xvii and xix, and the prostatic apertures are presumably
situated at these points; the grooves by their outward curve just get outside the
line of setae b at the middle of their length.

The female pores were not visible.

There was nothing to be seen in the spermathecal region.

Internal Anatomy :—I could not distinguish any septa in front of 6/7, which is
thin; 7/8 is thin, 8/9 to 10/11 slightly thickened, the rest thin.

There are two large gizzards in segments vi and vii. The calciferous glands are

three pairs, roundly ovoid masses in segments x, xi, and xii. The intestine begins
in XV.

The last heart is in segment xii.

Funnels were identified, perhaps somewhat doubtfully, lying free in segments x
and xi. The seminal vesicles are two pairs, in ix and xii, yellowish in colour, of moder-
ate size, somewhat lobulated and rather granular-looking on the surface.

The prostates were scarcely developed.

The spermathecae are two pairs, the ampulla of each a small ovoid sac which nar-
rows ectally to become the duct; the duct may be said to be half as long and half
as wide as the ampulla. A simple finger-shaped diverticulum, half as long as the am-
pulla, arises from the junction of ampulla and duct.

The penial setae are characteristic; their length is ‘3 mm., their thickness 8-07.
The distal portion shows a gentle sickle-shaped curve, the tip being slightly bent
in the opposite direction and bluntly pointed. There may be a constriction where
the curved distal meets the straight proximal portion of the shaft. Towards the tip
are a number of indentations of the margins, which however do not form spines
standing off from the shaft (fig. 55).

Eudichogaster pusillus, sp. nov.
Plate XI, figs. 56, 57.

Saugor, Central Provinces. 20-vi-1917. B. Prashad. A single specimen, damaged some
distance behind the clitellum
External Characters :—Length 28 mm.; diameter maximum 1°5 mm. Colourgrey-
ish, not distinctive; clitellum a reddish-brown. Segments ca. IIo.
Prostomium proepilobous.
Dorsal pores not seen in front of clitellum.
The setal relations are as follows:—In the middle of the body ab = half aa =

254 Memoirs of the Indian Museum. [Vor. VII,

3be — cd or almost so,—there is very little difference between bc and cd; immedi-
ately behind the clitellum the ratios may be expressed in the same way; in front of the
elitellum bc and cd are quite equal, —i.e. the lateral setae are not paired (ab = half
aa = 3bc = 2cd).

The clitellum is well defined, swollen, smooth, and includes segments xili-xvi
ventrally, with xvii in addition laterally and dorsally ( = 4 or 5).

The prostatic pores, on xvii, are a single pair of transverse slits which take up
the interval ab.

The female pores are probably contained in a whitish area situated anteriorly on
segment xiv; this is somewhat oval, with its long axis transverse and extending
between the lines aa; it is slightly hollowed.

The spermathecal pores could not be distinguished externally.

Internal Anatomy :—Septa 7/8-13/14 are slightly thickened.

There are two relatively very large gizzards, probably in segments v and vi,
perhaps in vi and vii. There are three pairs of caleiferous glands, in segments x, Xi,
and xii; they are ovoid, and not sharply set off from the oesophagus; those in x are the
largest, those in xi the smallest. The intestine begins in xv.

The last hearts are in segment xii.

The excretory system is micronephridial.

Funnels were doubtfully identified in segments x and xi; the worm is of small
size, and had possibly passed its period of maturity. A single seminal vesicle was
found, on the right side in segment ix; none were seen in xii or elsewhere.

The prostates are one pair, short tubes in segment xvii, bent once or twice. The
duct is much narrower than the glandular part,—is indeed very fine, but widens
gradually towards its ectal end; it is opaque white in colour, not shining, almost as
long as the gland, and runs transversely inwards.

A pair of relatively very large ovaries were found in xiii; the ova were seen as
large opaque white bodies, arranged in a fern-like manner. The funnels were also of
large size. A pair of ovisacs projected backwards from septum 13/14 into segment
xiv; they contained large ova.

The single pair of spermathecae are contained in segment vii, and appear to open
in or near furrow 7/8. Their appearance is peculiar (fig. 56); each is a long narrow
twisted cylindrical tube, somewhat wider at its ectal end, where a short muscular
duct,—only a fraction of the length of the ampulla, and about one-third as wide,—
leads to the exterior. There is no diverticulum. The whole organ looks at first sight
remarkably like a nephridium.

The penial setae (fig. 57) are relatively long compared with their en ; they
measure ‘56 mm., and are only 4; thick in the middle. The shaft is almost straight,
or at any rate is not regularly curved; it tapers very gently towards the tip, which
is flattened and slightly expanded.

Remarks:—This form seems to resemble Æ. parvus (Fedarb), but is to be dis-
tinguished by the presence of penial setae and the position of the calciferous glands
and seminal vesicles.

I)
Qu
Qt

1920. | J. STEPHENSON : Oligochaeta from India and E. Persia.

Eudichogaster Kinneari, sp. nov.
(Plate XI, figs. 58, 50.)
Nasik, 100 miles N.E. of Bombay, W. India. 22-ix-1914. N. B. Kinnear. A number of
specimens.

External Characters :—The length of a fair-sized specimen is 80 mm.; the dia-
meter behind the clitellum is 3°5 mm., while the bulbous anterior end is about 5 mm.
thick at the sixth segment. Colour buff, no difference between dorsal and ventral
surfaces, the clitellum brown. Segments of a fair-sized specimen about 120; segments
iv-vi are biannular, vii and viii quadriannular, ix and subsequent segments up to the
clitellum triannular with secondary subdivisions ; triannulation again becomes marked
towards the hinder end of the body.

_ The prostomium is small, prolobous, and withdrawn into the buccal cavity; the
first segment is small, and it is divided by a mid-dorsal longitudinal groove,—a well
marked and constant character.

The first dorsal pore is at the anterior border of the clitellum, in furrow 12/13.

Behind the clitellum, and in the body generally, the setal ratios may be expressed
thus :—ab = + to + aa = to $bc — ? cd; ab is not very regular however. In front
of the clitellum de diminishes very considerably; anteriorly it becomes less than
half aa and only about 14 times ab, and only slightly greater than cd; the ventral
couples are still widely separated from each other. The mid-dorsal interval dd is
about 3 of the circumference.

The clitellum is extremely well marked; it is brown in colour, swollen, and
extends over segments xiii-xvi (= 4); it is deficient in a V-shaped interval ventrally
on xiii. Dorsal pores and indications of furrows are visible, but not setae.

The male field is rectangular, its transverse extent greater than the longitudinal
(fig. 58); it includes segments xvii-xix. The margins, especially the lateral, are

much thickened ; from the anterior and posterior two tongues project, backwards and
forwards respectively, into the central depressed space, so that this becomes H-shaped.
The floor of the H-shaped depression is deeply fissured; in it two pairs of papillae,
one pair in each of segments xvii and xix, in the four corners of the H, represent the
prostatic apertures. The seminal grooves are not always distinguishable among the
numerous fissures of this region; they are like those of H. ashworthi and E. prashadh,

)

with an inward bend at the middle of their course, thus |

The female pores appear to be situated in a broadly spindle-shaped whitish area,
which shows up markedly against the brown of the clitellum, anteriorly and mid-
ventrally on segment xiv.

The spermathecal apertures are on the anterior part of the second annulus of
segments viii and ix, in the line of a. Each is in the centre of a low squarish papilla,
which takes up the greater part of the lensth of the segment.

There are numerous other papillae both in the region of the male field and also
near the spermathecal pores. Almost constant are two pairs, with clean cut edges,

256 Memoirs of the Indian Museum. [Vor. VII,

one on the posterior part of xvi, the other taking up most of the length of xx; these
extend from about, or within, the line a to the lateral margin of the body; they are
transversely oval in shape, and slightly depressed in the middle; the anterior pair
may be narrower from front to back than the posterior. In a number of cases there
are also median papillae, circular in outline, on one or both of these segments (xvi
and xx). In the spermathecal region other papillae are usually found to the outer
side of and touching those already described on segments viii and ix; these are
circular, and include the lateral setal bundles; there may even be two such papillae
on one side. Median papillae are often found in this region also, on the anterior part
of segments x, xi and (once) xii.

Internal Anatomy :—Septum 4/5 is thin, 5/6-11/12 are moderately strengthened,
the rest thin.

The gizzards, in segments v and vi, are large, firm, and globular. The calciferous
glands are two pairs, in segments xi and xii; they are small, ovoid, well set off from
the gut, and with smooth surface, not lobed. The canal swells out to form the
intestine in xv, but is much narrowed again as it passes between the bulky prostates.

The last hearts are in segment xii. There is a pair of large oblique vessels in
xiii, but these do not join the dorsal vessel as do the hearts,—certainly not at the
usual place, the hinder border of the segment.

The micronephridia are scattered in the anterior segments. Behind segment x
they are arranged in regular transverse rows; behind the prostates there are about
six on each side of each segment; of these the more dorsal are in regular longitudinal
rows, While the two most ventrally situated are closer together and not so regular.
Towards the hinder end of the animal the innermost on each side in each segment
becomes much thickened and more opaque, and thus, though it takes up no more
space transversely, it is much more conspicuous than the rest. The other five main-
tain their number and regularity however to within a very few segments of the end.

Testes and funnels, the latter of large size, are free in segments x and xi. Semi-
nal vesicles are present in segments ix and xii; those in ix are small, those in xii
much larger, lobed, and bulging back septum 12/13.

The prostates are large, the anterior pair extending over xviii and xix, or xvii,
-xviii, and xix and even getting into xx; the posterior pair occupies xx and xxi;
thus they overlap. The glandular part consists of closely adpressed coils. The duct
is much thinner, though rather wider as it approaches its termination; it is shining,
and thrown into one or two loops. Each duct leads forwards to end in the usual
situation.

The spermathecae are two pairs, in segments viii and ix. The ampulla is ovoid,
of moderately large size, and shows slight annulation ; the duct is well marked off
from the ampulla, and is half as long and two-fifths as thick as the latter; it narrows
towards its ectal end. The diverticulum is single, and consists of a large number of
minute seminal chambers, the whole attached by a short thick stalk to the ental end
of the duct (fig. 59).

Penial setae are absent.

1920. | J. STRPHENSON : Oligochaeta from India and E. Persia. 257

The copulatory setae of the spermathecal region are in their ornamentation
exactly similar to those of Z. prashadi, and no separate figure is needed. In length
they measured from ‘73 to ‘87 mm., in thickness 254 at the middle; and the extent of
the shaft occupied by the sculpturings is about one-third of its length. It is note-
worthy that whereas in most cases it is the ventral setae of segments viii and ix
which are modified, here the lateral setae, included in the lateral papillae on these
segments, may also develop as ‘“ copulatory setae.”

Genus Dichogaster.
Dichogaster bolaui (Mchlsn.).

Bombay. June 1915. N. B. Kinnear. Several specimens.
Bassein Road, N. of Bombay, W. India. 7-vii-1917. B. Prashad. Four specimens.
Baroda, W. India. 9-vii-1917. B. Prashad. Two specimens. —
Bayana, 20 miles S.W. of Bharatpur, E. Rajputana. 15-vii-1917. B. Prashad. A single
specimen.
var. malabaricus, var. nov.
Bombay; under a tree near the Fort. 30-vi-1917. B. Prashad. A single specimen.

External Characters :—Length 31 mm.; diameter 2°25 mm. Colour buff, unpig-
mented except for a dark mid-dorsal stripe. Segments 86.

Prostomium prolobous.

A conspicuous dorsal pore is seen in furrow 5/6, and then no more till 11/12,
where there is a rudimentary one. The pores are well marked from 12/13 onwards.

In the middle of the body the setal relations are as follows:—ab = 4aa = tbe
=cd; and they are about the same in the region behind the clitellum ; in front of the
clitellum all the pairs are closer together, more distinctly ventral, and dd, which be-
hind is about + of the circumference, consequently increases.

The clitellum extends over xiii-xx (—8); dorsally it almost covers xxi in addi-
tion. It is ring-shaped over xiii, a little thinner ventrally in xiv, xv, and xvi and
thenceforward interrupted ventrally. The region is swollen, and well defined at
each end.

The seminal grooves run in the interval ab; they are straight, and bordered by
whitish thickened lips; the inner lips of the grooves are almost contiguous in the
middle line.

The spermathecal apertures were not visible. The ventral surface of segment
vili, and perhaps of ix, appears slightly thickened, and the setae rather displaced and
irregularly set.

Internal Anatomy :—Septum 4/5 is slightly strengthened; if 5/6 and 6/7 are pre-
sent, they are not discernible in this specimen (which is not in very good preservation) ;
7/8 is thin, 8/9 and 9/10 perhaps slightly thickened, the rest thin.

The gizzards are in segments vii and viii, and the alimentary tube is scarcely
constricted between them. The calciferous glands are kidney-shaped, with the hilus
turned inwards, and occupy segments xv, xvi, and xvii. The intestine begins in xviii.

The last heart is in xii.

258 Memovrs of the Indian Museum. [Vor. VII,

The testes and funnels were not identified in segment x, but they seemed to be
present in xi. There were very small racemose seminal vesicles in segment xii.

The ovaries had the usual situation; there were small ovisacs in segment xiv.

The prostates and two kinds of penial setae are as for the typical form. The sper-
mathecae however present one of the peculiar features which distinguish this worm
as a separate variety; there are two diverticula, small, sessile, attached about the
middle of the duct.

Remarks :—The two spermathecal diverticula, and the fact that the clitellum is
not saddle-shaped throughout its extent, seem to warrant the separation of this form
as a distinct variety.

Dichogaster affinis (Mchlsn.).

Baroda, W. India, in a garden. 10-vii-1917. B. Prashad. Two specimens.
Bombay, under a tree near the Fort. 30-vi-1917. B. Prashad. A single specimen.
Bombay, Elephanta Island, in a rotten tree. 30-vi-1917. B. Prashad. Four speeimens.

Dichogaster crawi, Eisen.
Pashok, 3500 ft., Darjiling Dist., E. Himalayas. 1-12-x-1917. F. H. Gravely. A single

specimen. >

Remarks :—This species has not hitherto been recorded from India; it was des-
cribed by Eisen from the Pacific Coast of N. America, and its occurrence in India
may thus be compared with that of D. bolaw subsp. palmicola (12).

Subfam. OCNERODRILINAE.
Genus Ocnerodrilus.
Ocnerodrilus (Ocnerodrilus) occidentalis, Eisen.

Bombay. June 1915. N. B. Kinnear. A number of specimens.
Kotah, Rajputana. 14-vii-1917. B. Prashad. A number of specimens.

The specimens of the batch from Kotah show some of the characters of Eisen’s
var. avizonae. I have previously on a similar occasion (9) had the opportunity of con-
firming Michaelsen’s opinion of the non-validity of this variety.

Fam. GLOSSOSCOLECIDAE
Subfam. GLOSSOSCOLECLN AE.
Genus Pontoscolex.
Pontoscolex corethrurus (Fr. Müll.).

Bombay. June 1915. N. B. Kinnear. Two specimens, immature.
Poona, W. India, Empress Gardens. 3-vii-1917. B Prashad. Seventeen specimens, some

immature.

Subfam. MICROCHAETINAE.
senus Glyphidrilus.
Glyphidrilus papillatus (Rosa):
Lucknow. 15-i-1918. G.S. Thapar A number of specimens

1920.] J. STEPHENSON : Oligochaeta from India and E. Persia. 259

The worm has been described by Rosa from Burma, from a single specimen which
was not in very good condition, and not fully mature. As I have had a considerable
number of examples, carefully preserved by Mr. Thapar, to whom my thanks are
due for these interesting specimens, I give below a fairly complete description.

External Characters :—Length 120 mm.; diameter at and in front of the clitellar
region 3 mm., behind the clitellum 2 mm. Colour flesh-colour or greyish, no distinct
pigmentation ; a slightly darker mid-dorsal stripe. The posterior half of the body is
flattened on the dorsal side, and even concave in the hinder third; the ventral sur-
face is flattened or slightly concave for the greater part of its extent. Segments 202
in the example which was counted. Segment vi is biannulate, vii is triannular, and
so are viii and ix, but these may be further subdivided; the succeeding segments as
far as xv are bi- or triannulate. The anterior end of the body is tapering; the pos-
terior end becomes gradually thinner also.

The prostomium is large, zygolobous, and triangular. Dorsal pores are absent.

The setae are paired; behind the clitellum, the section of the body being some-
what quadrangular, the couples are at the four angles; ab is a little less than half aa,
is equal to half bc, and is equal to cd; dd is a little but not much greater than aa.
The setae become very small in front of the clitellum, and also wider apart. In the
posterior part of the body the setae are still at the angles, but the intervals are rather
different; dd becomes larger and bc smaller; ab = 2aa = half be = cd = 7dd.

Under close examination the anterior segments showed rings of minute pits ;
these were on the middle annulus where the segments were multiannular; possibly
they represent sense organs.

The clitellum is rather indefinite in its extent ; beginning at segment xvi it ex-
tends back to xxvi-xxxiv. The wing-like ridges extend ventro-laterally along each
side from segment xviii to xxiii, xxiv, xxv, or the anterior part of xxvi; they arise
from the body-wall outside the line of the ventral setal couples.

The characteristic papillae are large, round, flattened or slightly depressed in the
middle, and situated on the posterior parts of the segments. They are lateral or
median; the former series occur in line with or slightly dorsal to the attachment of
the winglike folds of the body-wall, are usually paired, and may be found on any
of the segments x-xvii, as well as occasionally on xxiii, xxv, or xxvi; they are com-
monest on xili-xvii, especially xv-xvii. The median papillae are not so common as
the lateral; I have found them only on xi-xv, and xvii and xviii. Median papillae
may be absent altogether; the lateral papillae may be as few as two pairs, or two on
one side and one only on the other.

Internal Anatomy :—The first definite septum is 4/5, which is thin; 5/6 is slightly,
6/7-9/10 moderately, and a few succeeding ones slightly thickened.

The gizzard is in segments vii and viii; septum 7/8 is adherent to it at about
its middle, but there is no constriction there; it is rather small, being no wider than
the preceding segments, and is fairly soft. The intestine begins in segment xvi.

Hearts are present in segments x and xi, and, in the specimen dissected, on the
right side in ix also.

260 Memoirs of the Indian Museum. [Voz. VII,

There was much flocculent matter in segments x and xi, and much of it was tena-
cious and iridescent, seeming to indicate the presence of seminal funnels; but neither
testes nor funnels were actually identified. The seminal vesicles are four pairs, but
they are not always symmetrical in each segment; thus in ix there was a large one
on the right, a small one on the left, while in x the condition was the opposite; in
xi there was a separate round lobe on the left side in addition to the normal vesicle.
The vesicles are usually deeply lobed.

There were no prostates.

The ovaries have the usual situation. I can confirm the presence of ovisacs in
xiv; there appears to be a pair of exactly similar structures in xv.

The spermathecae, which were not present in Rosa’s specimen, are in four series
on each side, each series consisting of a transverse row of five, opening in furrows
13/14-16/17. In addition there was a single one on the right side opening in 12/13.
Each spermatheca is a small ovoid or irregularly elongated saccule, its blind end direct-
ed backwards, and the whole adherent to the body-wall. : The position of the five
organs on each side is thus :—one in each of the lines a, b, c, and d, and one between
b and ce.

Fam. LUMBRICIDAE.
Genus Helodrilus.

Helodrilus caliginosus (Sav.) var. trapezoides (Ant. Dug.).
Mount Abu, S. Rajputana. Col. J. Manners-Smith. March 1918. Very numerous specimens.
Nasratabad, Seistan, E Persia, from a garden. 25-xi-1918. N. Annandale and 8. W. Kemp
A number of specimens.
Helodrilus parvus (Eisen).

Partabgarh, S. Rajputana. Feb. 1918. Col. J. Manners-Smith. A single specimen.

REFERENCES TO LITERATURE.

1. Beddard, F.E. .. A Monograph of the Order Oligochaeta. Oxford, 1895.
Michaelsen, W. .. Neue Oligochäten und neue Fundorte altbekannter. Mt.

Mus. Hamburg, vol. XIX, 1902.

3. 5 .. The Oligochaeta of India, Nepal, Ceylon, Burma and the
Andaman Islands. Mem. Ind. Mus., vol. I, 1909.

4. > .. Die Oligochätenfauna der vorderindischceylonischen Re-
gion. Abh. aus dem Geb. der Nam Hamburg, vol.
XIX, 1910.

5. Stephenson, J. .. The Anatomy of some aquatic Oligochaeta from the
Punjab. Mem. Ind. Mus., vol. I, 1909.

6. > .. On some aquatic Oligochaete worms commensal in Soon
gilla carteri. Rec. Ind. Mus., vol. V, 1910.
7. >) -. On some aquatic Oligochaeta in the collection of the Indian

Museum. Rec. Ind. Mus., vol. VI, 1911.

1920. | J. STEPHENSON : Oligochaeta from India and E. Persia. 261

8. Stephenson, J.
2

10. 5

ET: À

12. 99

18. ;

14. +

15. Walton, L. B.

Oligochaeta, in: Zoological Results of the Abor Expedi-
tion. Rec. Ind. Mus., vol. VITI, 1914.

On a collection of Oligochaeta, mainly from Northern India.
Rec. Ind. Mus., vol. X, 1914.

On a Rule of Proportion observed in the Setae of certain
Naididae. Trans. Roy. Soc. Edin., vol. L, 1915.

On some Indian Oligochaeta, mainly from Southern India
and Ceylon. Mem. Ind. Mus., vol. VI, 1915.

On a collection of Oligochaeta belonging to the Indian
Museum. Rec. Ind. Mus., vol. XII, 1916.

On a collection of Oligochaeta from various parts of
India and Further India. Rec. Ind. Mus., vol. XTII,
1917.

Aquatic Oligochaeta of the Inlé Lake. Rec. Ind. Mus.,
vol. XIV, 1918.

Naididae of Cedar Point, Ohio. Amer. Naturalist, vol.
XL, 1906.

EXPLANATION OF PLATE IX.

1.—Nais paraguayensis; needles of the dorsal bundles; a, with bifid outer
prong; b, with bent inner prong.
2.—Nais paraguayensis var. aequalis; dorsal needle, x ca. 1200.
3.—Nais gwaliorensis; dorsal needle, x ca. 1100.
4.—The same; ventral setae; a, of segments behind v; b, of segments ii-v; X
ca. 1100.
5.—Megascolides prashadi ; male genital field.
6.—The same; spermatheca.
7.—Perionyx sansibaricus; region of male apertures.
8.-—Perionyx millardı ; spermathecae showing different characters of diverti-
cula; a, specimen from Virar; 6, from Talegaon; c, from Kalyan.
9.—Perionyx rimatus ; spermatheca.
10.—Perionyx pokhrianus ; male genital field.
11.—The same; spermatheca.
12.— Perionyx pokhrianus var. affinis; male genital field.
13.—The same; spermatheca.
14.—Perionyx alatus; male genital field.
15.—The same; spermatheca.
16.—The same; tip of penial seta, x ca. 250.
17.—Perionyx shillongensis ; spermatheca.
18.—Perionyx fossus ; spermatheca.
19.—The same; tip of penial seta, x ca. 400.

Mem. Ind. Mus., Vol VI,1920. Dane ID.

xvill

Fig.15. 9.16. Fig.17. Fig. 18. Fig 19.

J.Stephenson del. A.Chowdhary lith.
INDIAN & PERSIAN OLIGOCHAETA.

EXPLANATION. OF PLATE X.

Fic. 20.—Perionyx turaensis ; spermatheca, x 40.
„ 21. The same; tip of penial seta, x 700.
,» 22.—Perionyx pullus ; spermatheca.
,, 23.---Perionyx minimus; spermatheca as seen under the microscope.
„ 24.—-Perionyx igatpuriensis ; spermatheca.

25.—Hoplochaetella anomala; male genital area.

,, 26.—The same; diagram showing relations of prostatic ducts and vasa deferen-
tia. Gl., prostatic gland (the coils. are not intended to be shown with
exactitude); p.d., prostatic duct; v.d., the two vasa deferentia.

,» 27.—The same; spermatheca.

» 28.—The same; sketch showing relations of accessory gland in neighbourhood
of spermatheca and copulatory seta, extracted together and seen under
microscope.

,, 29.—The same; tip of copulatory seta.

» 30.—Octochaetus paliensis ; spermatheca.

» 3l.—The same; penial setae, both from the same specimen as the above; x ca.
340.

» 32.—The same; copulatory seta, from same specimen as the above; x ca. 230.

» o3.—The same; spermatheca of specimen from Poona.

» 34.—Octochaetus paliensis var. riparius ; male genital region.

» 30.—The same; copulatory seta, x ca. 350.

,, 36.—Octochaetus prashadi; spermatheca.

» 372.--The same; tip of penial seta, x ca. 175.

» 88.— Ihe same; tip of copulatory seta, x 375.

» 39.—Octochaetus montanus; spermatheca.

» 40.-—The same; penial seta, x 40.

Mem. Ind. Mus.,‚Vol.V11,1920. Blade: %..

Fig. 38 . Fig. 36.

SICH ondul lith.

J. Stephenson del.
INDIAN& PERSIAN SINE OLSEN

EXPLANATION OF PLATE XI.

. 41.—Octochaetus pallidus ; spermatheca.

42.—The same; tip of penial seta, x 600.

43.—Octochaetus ganeshae ; spermatheca.

44.—The same; tip of penial seta, x 600.

45.—The same; tip of copulatory seta, x 500.

46.—Octochaetus pachpaharensis; spermatheca; div., diverticulum (?).

47.—The same; tip of penial seta, x ca. 300.

48.—Hutyphoeus turaensis ; spermatheca seen from above, as in dissection ; the
sac is attached to the body-wall by the middle of its under surface.

49.—The same; penial seta; a, seen as a whole, x ca. 55; b, tip more highly
magnified, x ca. 180. |

50.—Eudichogaster ashworthi (co-type); copulatory seta from spermathecal
region.

51.—The same; copulatory setae of neighbourhood of prostatic apertures (not
however from segments xvii or xix); a, from Wahi near Mahableshwar ;
b, from Poona; c, from Saugor; d, from Jubbulpore.

52.—Hudichogaster trichochaetus; penial setae; a, as a whole, x 37; b, the tip,
more highly magnified, x 550.

53.—The same; end of copulatory seta; x ca. 400.

54.—Hudichogaster prashadi ; a, region of male genital apertures ; b, end of copu-
latory seta, x 500. :

55.—Hudichogaster falcifer ; distal end of penial seta, x ca. 700.

56.—Eudichogaster pusillus ; spermatheca.

57.—The same; tip of penial seta.

58.— Eudichogaster kinneari ; clitellum and male genital area (the seminal grooves
are not seen; the grooves shown are irregular fissures in the floor of the
H-shaped depression).

59.—The same; spermatheca.

Mem. Ind. Mus., Vol. VII, 1920.

Plate XI.

Fig. 46.

Fig. 45.

Fig. 59.

A .Chowdhary lith.
INDIAN & PERSIAN OLIGOCHAET A.

À R Fig. 58.
JE Stephenson del.

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RR Vol. VII, No. 4.

_ REPORT ON THE PARASITIC NEMATODES IN THE
_ COLLECTION OF THE ZOOLOGICAL SURVEY
OF INDIA.

Fun at EX
H. A. Bayuıs, M.A., D.Sc., British Museum (Natural History)
ER : AND PU Sa N à
R. DAUBNEY, M.Sc., M.R.C.V.S., Ministry of Agriculture and Fisheries.

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PUBLISHED BY THE DIRECTOR, ZOOLOGICAL SURVEY OF INDIA.
_ PRINTED a THE BAPTIST MISSION PRESS. : x

DECEMBER, 1922. |

Price Eight Rupees. *

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REPORT ON THE PARASITIC NEMATODES IN THE COLLEC-
TION OF THE ZOOLOGICAL SURVEY OF INDIA.

By H. A. Bayrıs, M.A., D.Sc., British Museum (Natural History), and
R. DAUBNEY, M.Sc., M.R.C.V.S., Ministry of Agriculture and Fisheries.

(Published by permission of the Trustees of the British Museum.)

Received for publication November 23, 1921.

CONTENTS.
| Page Page
Introduction 2e 6 .. 264 Amplicaecum varani, Sp. nov. ~.. 287
Superfamily Ascaroidea. Dujardinia helicina (Molin) so Me
Family Ascaridae. Family Heterakidae.
Subfamily Ascarinae. | Subfamily Heterakinae.
Ascaris lumbricoides, L. so OE Heterakis papillosa (Bloch) .. 289
sale CREF ce 201 Heterakis isolonche, v. Linst. .. 289
Ascaris, Sp. x -. 268 Heterakis longecaudata, v. Linst... 290
Belascaris mystax (Zeder) =. 269 Heterakis bosia, Lane Soon
Belascaris marginata (Rud.) 209 Ascaridia perspicillum (Rud.) .. 292
Toxascaris leonina (v. Linst.) so BR) Ascaridia columbae (Gmelin) .. 292
Toxascaris transfuga (Rud.) so Zul Ascaridia compar (Schrank) 22203
Ophidascaris filaria (Duj.) .. 272 Ascaridia cristata (v. Linst.) - 11209
Ophidascaris naiae (Gedoelst) .. 273 Ascaridia stroma (v. Linst.) 17994
Folydelphis sewelli, sp. nov. -- 213 Strongyluris chamaeleonis, sp. nov. 294
Polydelphis oculata (v. Linst.) .. 274 | Pseudaspidodera pavonis, gen. et
Polydelphis, sp. Se ae) otk sp. nov. es _. 997
Subfamily Anisakinae. Subfamily Subulurinae.
eee ele CTS UNDE SOUR, Subulura sarasinorum (Meyer) .. 300
champs) = ce "278 Subulura galloperdicis, sp. nov. .. 300
Porrocaecum depressum (Zeder) .. 275 Subulura, Sp. N _. 302
Porrocaecum angusticolle(Molin) .. 275

ily Oxyuridae.
Porrocaecum serpentulus (Rud.) .. 277 Far is = Ser baa ae Oe
Porrocaecum reticulatum (v. Linst.) 278 GN ree ahs te nes

Porrocaecum pristis, sp. nov. .. 280 Carre compar, Leidy -. 303
Contracaecum spiculigerum (Rud) 281 au dactylura eu) .. 9303
Contracaecum rosarium (Connal) 282 Atractis opeatura, Leidy -. 303
Contracaecum incurvum (Rud.) .. 282 Family Kathlanidae.

Contracaecum tricuspe (Gedoelst).. 284 Falcaustra testudinis, sp. nov. .. 304
Contracaecum engonium, sp. nov... 284 | Falcaustra barbi, sp. nov. 292305
Contracaecum schizothoracis, sp. Falcaustra leptocephala, sp.nov... 306

DOVE Sr BA 2285 Falcaustra stewarti, sp. nov. ae HOOT

264 Memoirs of the Indian Museum. [Vor. VII,
Page | Page
Zanclophorus annandalei, gen. et _ Superfamily Trichinelloidea.
sp. nov. Sie | Family Trichinellidae.
Zanclophorus kempi. sp. nov. 312 | Subfamily Trichurinae.
Superfamily Filarioidea. | Trichuris trichiura (L.) 330
Bam arlarudae | Trichuris suis (Schrank) 330
len lad | Uses ovis (Abildg.) 330
Filaria hare awed 214 | Capillaria columbae (Rud.) 330
Filaria abbreviata, Rud. 314 | Superfamily Dioctophymoidea.
Filaria, sp. 515 | Family Dioctophymidae.
Filaria macrophallos, Parona 315 | Hustrongylides, sp. 331
Filaria varani, sp. nov. 316 Superfamily Strongyloidea.
Setaria, sp. Ril? | Family Strongylidae.
i : : 5 Subfamily Strongylinae.
Subfamily Diplotriaeninae. ; Eu
Diplotriaena tricuspis (Fedchenko) 317 Sa Nea GE 22]
Subfamily Micropleurinae, nov. Diaphanocephalus minutus, sp.nov. 332
Micropleura vivipara, v. Linst. 317 Diaphanocephalus, sp. 334
Superfamily Spiruroidea. Diaphanocephalus, sp 334
Family Spiruridae. Family Ancylostomidae.
Subfamily Acuariinae. Subfamily Ancylostominae.
Acuaria (Acuaria) anthuris (Rud.) 319 Ancylostoma duodenale, Dubini 335
Acuaria (Echinuria) leptoptili Ancylostoma caninum (Ercolani) .. 336
(Gedoelst) 319 Ancylostoma ceylanicum, Looss 336
Subfamily Physalopterinae. Ancylostoma malayanum, Alessan-
Physaloptera alata, Rud. 321 drini en sl
Physaloptera praeputialis, v. Linst. 322 | Galoneus perniciosus (v. Linst.) 336
Physaloptera, sp. 3929 | Subfamily Necatorinae.
Don Colle, Necator americanus (Stiles) 337
Camallanus kachugae, sp. nov. 322 Family Trichostrongylidae.
Camallanides prashadi, ger. et sp. Subfamily Trichostrongylinae.
nov. 395 Haemonchus contortus (Rud.) 337
Mail Ensthostomidae Haemonchus cervinus, sp nov. 337
Subfamily Spiroxyinae. Rain Metastrongylidae:
Spironys annulata Sp. nov. 328 Supe eae
3 Rictularia, sp. 338
Subfamily Gnathostominae. Family incert.
Tanqua tiara (v. Linst.) 329 Scolecophilus lumbricicola, gen. et
Tanqua anomala (v. Linst.) 330 sp. nov. 338
Echinocephalus spinosissimus (v. Cephalobus seistanensis, sp.nov... 341
Linst.) .. 330 Monhysterides piscicola, gen. et sp.
Gnathostoma spinigerum, Owen 330 nov. 342
INTRODUCTION.

The following report deals with an extensive collection of nematodes kindly
submitted to us for determination by Dr. N. Annandale, Director of the Zoological

Survey of India.

It comprises material belonging to the Indian Museum, Calcutta,

and material, collected since August, 1916, belonging to the Zoological Survey. A

1922. | H. A. Bayzis AND R. DAUBNEY: Parasitic Nematodes. 265

large proportion of this material was collected from animals, mostly Indian, in the
Zoological Garden, Calcutta. This applies to the majority of the species for which
no locality is given.

It is difficult to judge to what extent the range of hosts of a parasite may be
affected by the presence of a number of suitable hosts in more or less close proximity
under artificial conditions in a menagerie. It has been observed that wild animals
tend to lose their original parasitic infections after a short time in captivity ; but
there are some indications in the present collection that a parasite hitherto only
known to occur in one or two hosts may, under these conditions, have been enabled
to extend its range to hosts with which it would not normally have come into
contact. As instances we may mention particularly the cases of Ascaris lumbricoides
and the various species of Heterakis, especially H. longecaudata; while it seems
probable that the species of Ancylostoma, Belascaris and Toxascaris enjoy exceptional
opportunities in this respect in a menagerie where many kinds of carnivores are kept
near to each other. The lists of hosts that we have been able to compile for these
forms seem to bear out this suggestion.

As regards the position of parasites in their hosts, there was a certain amount
of vagueness in many of the original labels. Often the label indicated the
“intestines,” but this term appears to have been applied somewhat widely to
include most of the abdominal and thoracic viscera, and on this account we have
decided, in many cases, to omit any mention of the site of election. The species of
Heterakis from birds, for example, are usually found in the caeca of their hosts, and
to mention the “ intestines’ conveys no information of any value.

The present report deals mainly with nematodes from vertebrate hosts, although
two forms found in invertebrates are described. The material submitted to us also
included a number of Mermithidae, but we have not attempted as yet to deal with
these, and have thought it advisable to publish without unnecessary delay the results
of our work on the more strictly parasitic forms. These include members of nearly
every superfamily, and, while the number of new species is not large, the collection
is valuable for the light it throws on a number of imperfectly known forms, and
for the general idea it furnishes of the parasitic nematode fauna of India.

Throughout the report the names used for the hosts, so far as Indian animals
are concerned, are for the most part those given in the Fauna of British India
(Mammalia, by W. T. Blanford, 1888—1891; Birds, Vols. I--II, by E. W. Oates,
1889—1890, and Vols. IIJ—IV, by W. T. Blanford, 1895—1898; Reptilia and
Batrachia, by G. A. Boulenger, 1890; Fishes, by F. Day, 1889). The names of
hosts (other than domestic animals) which are not indigenous in the Indian Empire
are marked with an asterisk (*).

266 Memoirs of the Indian Museum. [Vor. VII,

Superfamily ASCAROIDEA, Railliet and Henry, 1915.
Family ASCARIDAE, Cobbold, 1864.
Subfamily ASCARINAE, Travassos, 1913. (Askarinae Raill. and Henry, 1912.)
Genus Ascaris, L., 1758.
Ascaris lumbricoides, L., 1758.
The collection contains material which we refer to this species from an interest-

ing range of hosts :—
Man (European boy, Calcutta).
Orang Utan * (Simia satyrus).
Large Indian squirrel (Sciurus indicus).'
Irrawaddy squirrel (Sciurus pygerythrus).’
ÆSdquirrel-d

This species is, of course, known to be à parasite of the larger apes, as well as of
man. It has long been a matter of opinion whether the form, often called A. suum or
A. suilla, occurring in pigs, both domesticated and wild, is or is not a distinct species
from À. lumbricoides. The discovery of what appear to be full-sized specimens of
the human worm in squirrels is of great interest, especially when the relatively
small size of these animals is taken into account. We have carefully examined and
compared specimens from man, from an Indian wild pig, and from the above-men-
tioned squirrels, paying particular attention to the characters of the lips, of the male
tail, and of the eggs, and our view is that all belong to the same species. The
number and arrangement of the caudal papillae of the male have been well figured
by Schneider (1866, p. 37). Quitc characteristic is the presence of two pairs of large
double papillae behind the cloaca, and three small simple papillae, arranged in a
triangle, on either side posteriorly. There is also constantly a pair of double
papillae at some little distance from the cloaca, in the preanal series, though the
corresponding papillae of the two rows are usually very asymmetrical in position.
A curious, large, median, papilla-like structure, or cushion, just in front of the cloaca,
and the short, broad, dorso-ventrally flattened, non-alate spicules, somewhat
enlarged in the distal half, are also characters common to all the material.

As regards the lips, little need he said except that no accurate account of the
cephalic papillae appears to exist. The dorsal lip carries two large, lozenge-shaped
papillae, with double terminations, near its lateral margins, while each ventro-lateral
lip has (a) towards the ventral side a large, double papilla, and (b) towards the
opposite side two small, separate, simple papillae. In the presence of these papillae,
and in all other respects, the lips of individuals from man, pig and squirrel seem to
us to agree.

The supposed differences between A. lumbricoides and “ A. suilla ” are based chiefly on size, the
worms found in the pig being usually of slenderer build than those from man. E. Blanchard (1849)

1 Now known as Ratufa indica... See Robinson and Kloss, Rec. Ind. Mus. XV, p. 186 (1918).
2 Now known as Tomeutes pygerythrus ; loc. eit., p. 226.

1922. | H. A. Bayuis AND R. DAUBNEY : Parasitic Nematodes. 267

also mentions differences in the relative lengths of parts of the female reproductive organs, but such
differences may constantly be observed in female nematodes of different ages, of the same species and
from the same host. In our opinion, the evidences of identity furnished by the structural characters
mentioned are of much greater weight.

Recent experimental work by Stewart and others has shown that A. lumbricoides can reach a
certain stage of development in the rat, mouse, guinea-pig and rabbit, but it has not yet been found to
settle in the intestine and attain sexual maturity in these animals. The present record of the adult
worm in squirrels (assuming, as we believe to be justifiable, that the species is the same), shows that
the development may, under suitable conditions, be completed in a rodent.

Ascaris vitulorum, Goeze, 1782.
(Figs. I—3.)

The distinctive characters of this form do not seem to be at all well known, and
we are not aware of any recent description of it. Ransom (IgII) gives only the
briefest details of its anatomy, and these seem to have been mainly quoted from Neu-
mann. A few specimens taken from a calf at Siripur, Bihar, though in rather poor
condition, enable us to add a few details to the description, and to correct others.

Our specimens measure from 85 mm. (male) up to 140mm. (female) in length.
The cuticle is marked with transverse rings at intervals of 0°03-0:075 mm. Finer
striations, if present, were not seen.
The diameter of the head, at the base
of the lips, is about 05 mm.; that of the
neck, immediately behind the lips, 0°7
mm. The lips (fig. 1) are broad at the
base and narrow in front. The dorsal lip
carries a pair of large, simple papillae;
each ventro-lateral lip a large, lozenge-
shaped papilla towards the ventral side
and a small, round papilla laterally and
more anteriorly. The pulp of each lip

sends out two rounded lobes anteriorly,

and from the inner surface of each lobe 0‘17vm.

a blunt inwardly-directed process origi- Fie. 1.— Ascaris vitulorum. Dorsal lip, viewed
2 from exterior.

nates. The two processes converge a., anterior lobe of pulp; d.r., dentigerous ridge ;

slightly towards the middle line of the lip. “., internal process of anterior lobe ; p., papilla.

Dentigerous ridges, with coarse teeth,

are present. ‘There are no interlabia. The oesophagus is modified posteriorly into
a small, almost globular ‘“ventriculus,” or glandular bulb (fig. 2), measuring
about 0°35 mm. long and 0°45 mm. wide. This is not distinctly constricted off from
the muscular portion of the oesophagus, but is preceded by a slight narrowing of the
latter. The entire oesophagus measures about 4 mm. in length. The nerve-ring
surrounds it at about 0°8 mm. from the anterior extremity.

1 Since this paper was prepared, a description has been published by Boulenger (Parasitology XIV, 1, 1922, p. 87),
which agrees on the whole with ours, except that he appears to have seen more pairs of postanal papillae than we could
detect in our material.

268 Memoirs of the Indian Museum. [Vor. VII,

The tail of the male (fig. 3) measures 0°33 mm. inlength. It is not, as is sometimes
stated in text-books, without postanal papillae. On the contrary, it possesses three

Fig. 2.—Ascaris vitulorum. Pos-
terior portion of oesophagus and com-
mencement of intestine.

i., intestine; m., muscular portion
of oesophagus ; v., ventriculus.

pairs on the ventral surface, of which the most ante-
rior consists of very large, double papillae. There
appear to be about nine pairs of preanal papillae.
The spicules are stout, tubular and without alae.
They were unfortunately broken in our specimens,
and we are unable to give their length. Their dia-
meter is about 0'043 mm.

The vulva is situated at about 17 mm. from
the anterior end in a specimen mm. 125 long, thus
dividing the body in the proportion of about r: 6.
The vagina and the unpaired portion of the uterus
run back, gradually widening, with a rather sinuous
course, to a point about II mm. behind the vulva,
before giving off the two parallel uterine branches.
The coils of the ovarian tubes return towards the
anterior end as far as the level of the vulva. The
eggs are oval, and have a thick, coarsely granul-
ated shell and an unsegmented content. They
measure 0°075—0°09 X 0°06-0°07 mm.

Ascaris, sp.
A small immature female, from the intestine of

a wild pig (Sus bengalensis), near Dinapore, Bihar. It is doubtful whether this can be

assigned to A. lumbricoides.

Fic. 3.—Ascaris vitulorum. Tail of male; ventral view.

1922. | H. A. BayLıs AND R. DAUBNEY : Parasitic Nematodes. 269

Ascaris, sp. (?)
A single immature specimen, about I4 mm. in length, from Varanus salvator.
We are unable to assign this definitely to any species.

Genus Belascaris, Leiper, 1907.
Belascaris mystax (Zeder, 1800).
This species occurs in the collection from the following hosts :—
Domestic cat.
Siamese domestic cat.
Tiger (Felis tigris).
Leopard (felis pardus).
Jungle cat (Felis chaus).
Leopard cat (Felis bengalensis).
Fishing cat (Felis viverrina).

Belascaris marginata (Rud., 1802).

In addition to a single female of this species from a jackal and some doubtful
specimens, in very poor condition, from an Indian wolf (Canis pallipes), the collec-
tion contains a number of Ascarids from the stomach. and intestine of an Indian fox
(Vulpes bengalensis). After comparison with material from the domesticated dog,
we conclude that these worms from the fox belong to the same species (B. marginata).
It may be mentioned in this connection that Riley (1921) states that this species fre-
quently occurs in foxes farmed for commercial purposes in Canada and the United
States. At the same time, the Ascaris vulpes of Frölich, 1789 (=A. triquetra, Schrank,
1790), was regarded by Railliet and Henry (1911) as a distinct species of Belascaris,
although no satisfactory description of it appears to exist. The only distinctive
features of this supposed species mentioned by Railliet and Henry are the greater
development of the caudal alae, and the presence of a gutter-like depression of the
ventral surface of the tail,in the male. These, as it seems to us, are appearances
which might easily be the result of imperfect preservation or extreme contraction,
and there appears to be no other ground for accepting Belascaris vulpis as a valid
species.

It is appropriate in this place to refer to two other species recorded from more or less closely related
hosts. Belascaris masculior, Railliet and Henry, 1911, from the Fennec fox (Megalotis zerda), appears
to be a species of smaller average dimensions than B. marginata, but with somewhat longer spicules.
There is nothing else in the description by which it can be differentiated.

B. melis, Gedoelst, 1920, from the badger, attains a very large size, though its spicules are not
longer than those of B. marginata. According to Gedoelst’s account, there are only three pairs of
papillae on the terminal appendage of the tail in the male, instead of the five pairs usual in the genus,
and the number of preanal papillae appears to be unusually large.

270 Memoirs of the Indian Museum. [Vor. VII,

Genus Toxascaris, Leiper, 1907.
Toxascaris leonina (v. Linst., 1902).
This species occurred in the following hosts :—
Lion (Felis leo).
Tiger (Felis tigris).
Leopard (Felis pardus).
Ounce, or Snow leopard (felis uncia).
Fishing cat (Felis viverrina).
Leopard cat (Felis bengalensis).
Hunting leopard (Cynaelurus jubatus).
(?) Indian fox (Vulpes bengalensis).'

Toxascaris transfuga (Rud., 1819).
(Figs. 4, 5.)

The collection contains abundant Ascarid material from bears—the Himalayan
black bear (Ursus torquatus) and the sloth-bear (Melursus ursinus)—and also from the
red cat-bear (Aelurus fulgens), all of which we refer to Ascaris transfuga Rud. Aelurus
appears to be a new host for this species. Examination of the material shows clearly
that A. transfuga has all the essential characters of the genus Toxascaris, as defined
by Leiper (1907) and by Railliet and Henry (1911).

The best description of the species at present existing appears to be that of
Dujardin (1845), but he gives no figures. Schneider ((1866), pl. I, fig. 3) gives an
accurate figure of the dorsal lip, seen from the inner surface, but we are unable to
find a figure of the tail of the male. The characters of this, and of the dorsal lip
as seen from the outer surface being of considerable importance, we have prepared
figures to illustrate these points, and we propose to amplify the description somewhat.

The size attained sometimes exceeds the measurements given by Dujardin. We
have examined one female specimen (not in the present collection), from the brown
bear, which measured as much as 240 mm. in length and about 4:5 mm. in thickness.
This, however, appears to be exceptional. The anterior end of the worm (in spirit)
is usually, though not invariably, curved towards the dorsal side. The lips are
roughly semicircular in outline, and each carries two papillae, those of the dorsal lip
(fig. 4) being equal and symmetrically arranged, while those of each ventro-lateral
lip are rather unequal and asymmetrical, the papilla towards the ventral side being
large, the more lateral papilla smaller and situated slightly further forward. The pulp
of each lip sends out five processes, two in a transverse direction, near the ends of
which are the papillae, and three anteriorly. Of the anterior processes two form
large paired lobes which expand slightly and have a shallow longitudinal groove on
their inner surfaces at their distal ends (cf. Schneider’s figure). The third is the

! The last record may perhaps be due to a clerical error in the collector’s label, Vulpes having been written by a
lapsus for Felis.

1922. | H. A. BayLıs AND R. Dausney: Parasitic Nematodes. 271

median unpaired lobe, which is a supporting structure for the cuticle of the inner
surface of the lip, and appears to end in short rays which spread out beneath the cuticle.
The paired anterior lobes originate
from the inner side of the main pulp dr
of the lip somewhat behind its ante- :
rior limit, so that a kind of transverse
eroove, as described by Railliet and
Henry for the type-species, is formed
between the main mass and the ante-
rior lobes. Marginal dentigerous rid-
ges, composed of relatively very large
and rather irregular teeth, are pres-
ent.

The cervical alae are well-develop- I |
ed. The oesophagus is simple (with- olmm.

out ventriculus) and club-shaped, very Fig. rn transfuga. Dorsal lip, viewed
; rom exterior.

stout posteriorly (up to 0°95 mm.), and d.r., dentigerous ridge; m./., median lobe of pulp;

measures 4-5 mm. in length. There pl.paired anterior lobe; p., papilla; ¢., transverse
process of pulp.

are no oesophageal or intestinal divert-
icula.

The caudal end of the male (fig. 5) is curved ventrally, but the extremity is usually
recurved towards the dorsal side. The tail itself measures 0'45-0°5 mm. in length,
is bluntly conical, and ends in a short spike which has a small terminal button, and
thus resembles a terminal papilla. The postanal papillae correspond in number,
though not exactly in position, with those of T. leonina, the genotype. There are
(I) a group of four pairs near the tip of the tail, consisting of two very small ventral
and two larger, more lateral pairs. Of the latter the more posterior is the most
laterally situated, and the more anterior is the largest, of the group; (2) an isolated,
quite lateral pair; (3) a large pair with double terminations, situated at the posterior
limit of a kind of raised wall of cuticle which runs forward and round the cloaca,
bounding a horseshoe-shaped depression. The edges of this “wall” are usually
curved inwards, so that the terminations of the papillae face towards the mid-ventral
line; (4) the first of a series of upwards of thirty pairs of papillae which extend for a
considerable distance in front of the cloaca. These papillae are at first close together,
and each row tends to resolve itself into two alternating rows, but further forward the
row becomes simple and the papillae wider apart. The spicules are very short (about
0°65 mm.) and stout, and are tubular structures without alae, and covered with
small granulations.

The tail of the female is short and bluntly conical, almost rounded posteriorly, but
with a small papilla-like termination, as in the male. The vulva is situated at about
the anterior third of the total length in young females, but in large examples the
post-vulvar portion of the body appears to have grown more rapidly than the anterior
part, so that the vulva divides the body in the proportion, roughly, of 2:5. The

272 Memoirs of the Indian Museum. [Vor. VII,

narrow, convoluted vagina opens into a short (not exceeding 5 mm.) unpaired uterine
chamber, as described by Dujardin, from which the two branches of the uterus run
back parallel to each other, and nearly
straight in large specimens. The coils
of the £ovarian tubes, after running
nearly to the posterior end of the
body, return anteriorly as-far as the
level of the vulva. The eggs are
oval (not globular), and have a thick,
smooth shell measuring 0°0775-0:09 X
0°06—0'075 mm. When ready for lay-
ing, this shell appears to become
covered, as in Ascaris lumbricoides,
with an irregular external coat of a
yellowish albuminoid substance, which
perhaps gave Dujardin the impression
of a “punctulated” shell. The con-
tent of the egg is unsegmented at the
time of laying.
Genus Ophidascaris, Baylis, 1921.
Ophidascaris filaria (Duj., 1845).
This species occurred in abund-
ance in the alimentary canal of Python
molurus on nineteen occasions. We

Fra. 5.—Toxascaris transfuga. Tailof male; latero- have also to record the presence of

ventral view. immature forms of various ages in the
s., right spicule ; w., wall of depression surrounding
cloaca.

lung of Python molurus and P. reticul-
atus. In two cases the same animal
harboured adults in the intestine and larvae in the lung at the same time. The
immature worms in the lung measured from about 9 to 60 mm. in length, but in
the largest of them the lips had not yet acquired their definitive structure (except in
one case where the label stated that the material came from the lung, but the accuracy
of this statement may be doubted, as the uteri of the females already contained
Ova).

From the occurrence of the young forms in the lung of the python, it appears probable that the
larvae have a course of migration within the body of the host, like that of the larvae of Ascaris
lumbricoides, before finally establishing themselves in the alimentary canal.!

! It has recently been shown by Ortlepp (JJ. of Trop. Med. and Hyg., XXV, p. 97) that the embryos of Polydelphis
attennata are capable of partial development in the mouse, and that they migrate through the lungs as in the case of
Ascaris lumbricoides. Our note was written before the appearance of Ortlepp’s paper.

1922.] H. A. Bayrıs AND R. Dausney: Parasitic Nematodes. 273

Ophidascaris naiae (Gedoelst, 1916).

We refer tentatively to this species forms found in the intestine of a cobra (Naja
tripudians) and in the stomach and intestine of two kraits (Bungarus fasciatus).
Their determination is open to question, all being young and not yet full-sized,
although two of the females contain ova. Apart from measurements, they seem to
agree fairly well with Gedoelst’s (1916) rather brief description.

Genus Polydelphis, Duj., 1845.
Polydelphis sewelli, sp. nov.
(Figs. 6, 7.)

Host: Montpellier snake (Coelopeltis monspessulana*). Locality: Khan- Yunis,
Palestine, May, 1917.

Some Ascarids collected from the “‘abdomen and lung’ of the above-named
snake by Major R. B. 8. Sewell prove to belong to the genus Polydelphis, and to
the section of that genus in which the uterus
has six branches.! The worms are relatively
slender and of almost uniform thickness
throughout. The male measures about 66
mm. in length and 1:2 mm. in thickness, the
female 73 mm. and I’4 mm. respectively. The
cuticular striations are exceedingly fine. The
diameter of the head is 0°3-0°34 mm. The
lips (fig. 6) are somewhat hexagonal in outline,
and slightly broader than long. Their anterior o-1mm.
and lateral borders are somewhat emarginate. Fi. 6.—Polydelphis sewelli. Dorsal lip of
Marginal dentigerous ridges are present, and ua en nz utior.

; a.l., anterior lobe of pulp; d.r., denti-
the pulp of the lip has two well-developed serous ridge; p., papilla.
antler-like anterior lobes, each having two main
divisions. The dorsal lip carries two simple lateral papillae, each ventro-lateral lip
one large ventral and one very small lateral papilla. The oesophagus is simple,
somewhat enlarged posteriorly, and measures about 5 mm. in length. There is, at
least in some individuals, an intestinal caecum, which may reach a length of about
o°6 mm., or may be quite rudimentary and not more than 0125 mm. long. A pair
of very small cervical papillae is present at about I’4 mm. from the anterior end.
The nerve-ring is situated at 0°7-0'85 mm. from the same point.

The tail, in both sexes, is very short and rounded, and terminates in a small
spike about 0°05 mm. long.

In the male, the tail (fig. 7) measures 0:25 mm. in length. The spicules are
short, subequal and broadly alate. The right spicule measures 1°35 mm, the left
I’4 mm., and the dorso-ventral diameter of each is about 0°06 mm. The preanal

1 See Baylis (1921).

274 Memoirs of the Indian Museum. [Vor. VII,

papillae are arranged in a single fairly regular and close series of about 43 on either
side. There is also one papilla, more laterally situated, at about the same level as
the fourth of the series, on each side. There are six pairs of postanal papillae, of
which the first, or most posterior, papilla on each side is relatively large and dorsally
placed; the second is very small and lateral; the third, fourth and fifth form a
triangle, two of them being ventral and one more lateral; and the sixth is a large,
double papilla near the cloaca and separated by a considerable space from the rest.
The tail of the female is 0°3 mm. long. The vulva is situated at about 34 mm.
from the posterior end, dividing the body in the proportion of about 19 : 17. The

Fic. 7.—Polydelphis sewelli. Tail of male; lateral view.

narrow, irregularly coiled, muscular vagina leads into a common uterine chamber
about 1°6 mm. long, which gives off the six parallel uterine branches posteriorly.
These run back to a point about 15 mm. from the posterior end, where they pass into
the ovarian tubes. The coils of the latter extend back to 10 mm. from the posterior
end, and then run forward to the level of the vulva. The ova are roundish-oval or
nearly spherical, and have a thick, finely granulated shell, measuring 0'09-0°I X
0°075—-0'087 mm.
Polydelphis oculata (v. Linst., 1899) (?).

Two females from the stomach of a python from Assam (? Python molurus or P.
reticulatus) may belong to this species, but are in too poor a condition to be deter-
mined with certainty.

Polydelphis, sp.
[near to P. hexametra (Gedoelst, 1916). |

Two female specimens, one in very poor condition, were taken on separate occa-
sions from examples of the common chameleon of India (Chamaeleon calcaratus).
They belong to the section of the genus in which the number of uterine branches is

1922. | H. A. Bayzis AND R. DAUBNEY : Parasitic Nematodes. 275

six, but it is impossible to determine whether the species is identical with the Ascaris
hexametra of Gedoelst (1916), from an African chameleon.

Subfamily ANISAKIN 4, Railliet and Henry, 1912, emend. Baylis, 1920.
Genus Porrocaecum, Railliet and Henry, 1912.
Porrocaecum crassum (Deslongchamps, 1824).

One female specimen of this species was collected from a duck, at Bombay.

Porrocaecum depressum (Zeder, 1800).
(Fig. 8.)

Examples of this species occurred in the intestine of the cinereous vulture (Vul-
tur monachus). The characters of the dorsal lip and of the male tail have been
figured by Schneider (1866). We append
a new figure of the dorsal lip for purposes
of comparison with the next species, P.
angusticolle, since the differences between
these two species are so slight as to require
emphasis.

Porrocaecum angusticolle (Molin, 1860).
(HiSS NO Os)

The material consists of a few com-
plete specimens and some fragments from
a kite (Milvus govinda). =

This species is nearly related to the ;
foregoing. The female was well described by 4 ie

Fig. 8.— Porrocaecum depressum. Dorsal lip of
von Drasche (1883), who figured the dorsal female, viewed from exterior.
lip. The male has, up to the present, not d.r., dentixerous ridge; m.l, median lobe of
been described. The principal character Pulp; ?» Heure
which serves to distinguish the species from
P. depressum is the shape of the pulp of the dorsal lip. In P. angusticolle the dorsal
lip is almost hexagonal in outline. The main mass of the pulp resolves itself into
two principal lobes, rounded anteriorly and joined by a saddle. Springing from the
inner surfaces of these lobes are two processes which become visible anteriorly as two
projecting plates, flattened and expanded distally. Towards the base of the lip there
is on either side a cuticular band (fig. 9, c.) which stands out somewhat prominently.
There is the usual pair of papillae on the dorsal lip, while a dentigerous ridge may be
traced round the greater portion of the lip near its edge. Small triangular interlabia
are present. Figs. 8 and 9 show the difference between this arrangement and that of
P. depressum. In the case of the latter each of the main lobes divides anteriorly into
two more or less finger-like processes, as described by Schneider, while internally
there is a large, median lobe, rounded anteriorly, which is distinctly visible where it
projects beyond the saddle joining the two main lobes.

276 Memoirs of the Indian Museum. [Voz. VII,

The male measures up to 55 mm. in length and I'ı mm. in thickness: the female
90 mm. and I'5 mm. respectively. The cuticle has transverse striations about 17,

eee)
0-1M™M.

Fig. 9.—Porrocaecum angusticolle. Dorsal lip of
female, viewed from exterior.

c., ‘cuticular band”’; d.r., dentigerous ridge: p.
papilla.

apart. Anteriorly the body is tapered
for a considerable distance, forming a
long, slender neck. The head is small,
its diameter being 0'245 to 0'26 mm.
There is a slight constriction at the
junetion of the head and neck. The
oesophagus is 4°8 mm. long, including
the short, oblong ventriculus, which
measures 0°6 mm. in length. The ante-
rior caecal prolongation of the intestine
measures from 2°7 to 3:0 mm. in length.
The nerve-ring is situated at about 0°85
mm. from the anterior end. At about
1'7 mm. from the head there is a pair of
large, sessile cervical papillae. The ex-
cretory pore opens, as usual, just behind
the base of the lips in the median ventral
line.

The tail of the male (fig. 10) is conical and measures 0°39 mm. inlength. About
half-way between the cloaca and the tip of the tail there is a distinct constriction.
There are no caudal alae. The postanal papillae are all sessile. There are about

FIG. 10.—Porrocaecum angusticolle.

Tail of male; ventral view.

1922. ] H. A. BayLıs AND R. DAUBNEY : Parasitic Nematodes. 211

twenty pairs of preanal papillae, the most posterior of which is situated just anteriorly
to the corners of the cloacal opening. The postanal papillae consist of five pairs; a
large, ventral pair of double papillae just behind the cloaca, and the remaining four
pairs on the posterior half of the tail, 2.e., behind the constriction mentioned above.
Two of these latter pairs are distinctly ventral and two ventro-lateral. The spicules
are equal and simple (not alate). They measure 0°95 mm. in length.

The tail of the female is blunter than that of the male, and measures 0°7-mm. in
length. The caudal papillae are situated at 0-2 mm. from the tip. The vulva is
situated in the anterior half of the body, dividing the latter in the proportion of 3 : 5.
The eggs measure 0°085-0°093 mm. X 0'058-0'074 mm.

In addition to the difference in the pulp of the dorsal lip, there are certain other
points in which this species diverges from P. depressum. The mature female of
P. depressum is much shorter in proportion to its thickness than the female of
P. angusticolle, and the vulva in P. depressum is situated further back, towards the
middle of the body, the proportion in which it divides the body being about 5 : 6.

Porrocaecum serpentulus (Rud., 1809).
(re nur)
Examples of this species occurred in the common crane (Grus communis) and in
the demoiselle crane (Anthropoides virgo). The characters of the head, and more
especially of the dorsal lip, have been described and figured by von Linstow ((1899),

Fie. 11.—Porrocaecum serpentulus. Tail of male; lateral view.

p. 7; pl. 1, fig. 9). The bifurcation of the anterior lobes of the pulp of the lip
appears to be less pronounced than is-indicated in his figure. Dentigerous ridges are
not, as he states, absent. In describing the tail of the male the same author

278 Memoirs of the Indian Museum. [Vor. VII,

mentions four pairs of papillae on the finger-shaped terminal appendage, two ventral
and two “dorsal.” We find, in addition to two ventral and two subdorsal pairs,
a fifth pair, lateral in position (fig. 11).! The one other postanal papilla on either
side, mentioned by von Linstow, is on the thicker portion of the body, and is a
double papilla, facing posteriorly. There is a regular series of about 15 preanal
papillae on either side, as stated by von Linstow. The spicules are equal in length
(1:25 mm. in a moderately large specimen). Each is composed of a tubular shaft
having a transversely striated appearance and gradually increasing in diameter to-

wards its base, and two very broad membranous alae.
This species, when taken from cranes, has a relatively short and stout build, some females attaining
a diameter of about 4mm. Some specimens from Ardea cinerea in the British Museum are so slender in
proportion to their length, as compared with those from cranes, that it seemed probable that they
belonged to a different species. We have found no important difference, however, in the structure of
the head or of the male tail, and therefore conclude that the forms in cranes and in herons are all
P. serpentulus.
Porrocaecum reticulatum (v. Linst., 1899).
(Fig. 12).

Syn. Ascaris reticulata, v. Linstow (1899), p. 7; pl. I, fig. 11.

Ascaris ardeae, Smith, Fox and White (1908), p. 287; pl. VI, figs. 1-7.

(nec Ascaris ardeae, Frölich, 1802; A. ardeae, Diesing, 1851; A. ardearum, Rud., 1819.)

Material from the intestines of the Eastern purple heron (Ardea manillensis),

the night heron (Nycticorax griseus) and an egret (species not mentioned), all from
the Caleutta Zoological Garden, appears to us to be referable to Ascaris reticulata.
All our specimens, however, are rather small compared with the measurements given
by the authors cited, and although the sexual characters are developed and the
females already contain ova, it is probable that they had not yet attained their full
size. On comparison of the descriptions and figures given by von Linstow and by
the American authors, we feel no doubt as to the identity of A. reticulata and
A. ardeae, Smith, Fox and White. In both cases, however, important points appear to
have been overlooked by the observers, as our own material shows. Von Linstow
states that interlabia and dentigerous ridges are absent, while omitting any descrip-
tion of the structure of the oesophagus and the anterior part of the intestine. The
American authors, on the other hand, observed the well-developed interlabia and the
presence of dentigerous ridges, but state that the oesophagus has no ‘bulb’, and
make no mention of an intestinal caecum. From our own observations it is clear
that the characters of the head have been accurately described by Smith, Fox and
White. It is just possible to understand how the interlabia were overlooked by von
Linstow, since in a cleared specimen in certain positions they are almost entirely
hidden by the lips, and their delicate outlines become very elusive. It is less easy
to explain how the American authors failed to see the extremely well-developed
intestinal caecum, which runs forward beside the oesophagus for a considerable

! Our figure is taken from a specimen from Grus australasiana, in the British Museum, and not from Indian Museum

material.

1922. ] H. A. BayLis AND R. DAUBNEY : Parasitic Nematodes. 279

portion of its length, and is quite a conspicuous organ. ‘he rather short, oblong
ventriculus is less conspicuous, but is not difficult to observe in a well-cleared speci-
men. All these characters, taken together, leave no doubt as to the generic position
of the species.

The characters of the caudal end of the male appear to have been adequately
described and figured by Smith, Fox and White, (von Linstow’s material consisted of
females only)—with the remarkable omission of any mention of an accessory piece, in

0:3 MAL.

Fig. 12.—Porrocaecum reticulatum. ‘Tail of male; lateral view.
a.p., accessory piece ; s., right spicule.

addition to the two spicules. The presence of such a structure in an Ascarid is highly
remarkable, yet in the material at our disposal every male possesses a conspicuous
accessory piece (fig. 12, a.p.) composed of clear, yellowish-brown chitin. This organ
appears smooth, whereas the spicules (which are of a darker colour and are simple,
tubular structures, without alae), have a rough, granular appearance. As regards
the caudal papille of the male, we are in agreement with the description given by
the American authors. There are two very small pairs on the finger-like caudal
appendage; one large postanal pair just before the constriction of the tail; and an
anterior row in which we have counted five on either side, commencing with a pair
at the level of the cloaca.

The almost spherical eggs, in our specimens, are slightly smaller than the
measurements given by the former describers, but this is possibly due to the im-
maturity of the females. The egg-shell is pitted externally, as described by the Ameri-

280 Memoirs of the Indian Museum. [Vor. VII,

can writers, the walls between the pits doubtless forming the “network of ridges”’
referred to by von Linstow.

This species is readily distinguished from P. serpentulus by the absence of cepha-
lic alae, and by the simple character of the spicules and the presence of an accessory
piece in the male.

Porrocaecum pristis, sp. nov.
(Figs. 13-15.)

Host: Saw-fish (Pristis perotteti). Position: intestine. Locality: Ulubaria, R.
Hughli.

The male measures up to 26°6 mm. in length and 0°74 mm. in thickness; the
female 34'2 mm. and 1:06 mm. respectively. The diameter of the head is 0°I6-0'18
mm. The cuticle has transverse striations 8°7-Ior apart. The lips (fig. 13) are
small, and pass into the neck without any constriction at their bases. Each has a

narrow, bilobed, anterior process, carry-

dl ing two small cuticular projections on its

ERNST xg inner surface. ‘The dorsal lip is shorter

SI N: than the ventro-lateral lips, and has two
large papillae. Each ventro-lateral lip
has one large, lozenge-shaped ventral
papilla and a much smaller lateral pa-
pilla, situated a little more anteriorly.
Dentigerous ridges are present, at least
on the anterior processes of the lips.
The teeth are very small. Interlabia are
absent. The oesophagus has a straight
posterior glandular portion, or ventricul-

0: y, D
: ee ous, of oblong shape. The distance from
Fia. 13.—Porrocaecum pristis. Head of female ; :
dorsal view. the head end to the posterior end of the
d.l., dorsal lip; p.. right papilla of same. ventriculus is 2°3-2°7 mm. The ventri-

culus is 0°6 mm. long in the male,
0°75-0'78 mm. in the female. The anterior caecal prolongation of the intestine
measures 0'95-I'3 mm. in length. There is a pair of prominent, rounded, cervical
papillae at 0°65-0°67 mm. from the anterior end. The nerve-ring is situated at
0'46-0°53 mm. from the same point. The excretory system terminates, as usual in
the genus, in a long unicellular gland with a very narrow duct leading to the excre-
tory pore, which is situated just between and behind the ventro-lateral lips.

The tail of the male (figs. 14, 15) is conical, slightly curved ventrally, and
measures 0°38 mm. in length. There are slight caudal alae, and most of the papillae
have rather long, rib-like pulps. There are about 40 pairs of preanal papillae, and
in addition to these there is one median sessile papilla on the anterior lip of the
cloaca. One pair of papillae, apparently belonging to the preanal series, is situated
just at the corners of the cloacal opening. There are seven pairs of postanal papil-

1922.] H. A. Bayzis AND R. DAUBNEY : Parasitic Nematodes. 281

lae, of which the second and third from the tip of the tail are laterally, the rest ven-
trally situated. The fifth papilla on either side is larger than the rest, and has double
terminations. The spicules are equal, simple and without alae. They measure only
0° mm. in length.

The tail of the female is bluntly conical and measures 0°44 mm. in length. The
caudal papillae are situated at 0'162 mm. from the tip. The vulva is somewhat be-

Fie. 14.—Porrocaecum pristis. Tail of Fig. 15.—-Porrocaecum pristis. Tail of male;
male; lateral view. ventral view.

hind the anterior third of the body—at 12°3 mm. from the anterior end in a specimen
34:2 mm. long. The muscular vagina, which runs posteriorly, is very short (about
0'7 mm.), expanding in its posterior half to a diameter of 0‘19 mm. This swollen
portion is packed with ova. Then follows a wide uterine reservoir, about 2 mm. long,
which gives off posteriorly the two uterine branches. These run almost straight to-
wards the posterior end. The posterior limit of the coils of the ovarian tubes is
about I’5 mm. from the posterior end. The ova are spherical, with a thin shell,
measuring 0‘0475 mm. in diameter. The content of the egg is unsegmented when
ready for laying.

Ascaris circularis v. Linst. is recorded as a parasite of Pristis antiquorum in the Cameroon. Von
Linstow (1907), in his description of it, mentions the presence of an intestinal caecum, and it is not
improbable that the species also belongs to the genus Porrocaecum. But the figure of the dorsal lip
(l. c., pl. 6, fig. 1) is sufficient to differentiate it from P. pristis.

Genus Contracaecum, Railliet and Henry, NORA,

Contracaecum spiculigerum (Rud., 1809).
Hosts:
Little cormorant (Phalacrocorax javanicus).

Indian shag (Phalacrocorax fuscicollis).

282 Memoirs of the Indian Museum. [Vor. VII,

Contracaecum rosarium (Connal, 1912).
(Fig. 16.)

The collection contains worms taken on two occasions from the night-heron
(Nycticorax griseus). The material consists in one case of two immature males, the
tails of which are eroded and useless for purposes of identification, and in the other
case of three immature females and one rather damaged, immature male. We have

O-L ITUTTe.

Fig. 16.—Contracaecum rosarium. Tail of male; ventral view.

assigned these to Contracaecum rosarium. There is nothing in Connal’s (1912) de-
scription to indicate a difference between his species and C. microcephalum (Rud.),
except that there are three pairs of postanal papillae in the male. With the material
at our disposal it is not possible to redescribe the species. The head generally and
the dorsal lip in particular are indistinguishable from those of C. microcephalum.
However, the tail of the male in the second set shows quite clearly that there are
more than three pairs of postanal papillae, and that the number and arrangement of
these papillae (which, in the only specimen available, are unfortunately somewhat
asymmetrical), will serve to differentiate this species from C. microcephalum. There
are nine pairs of postanal papillae. Those of the pair at the tip are stalked and
nipple-like, while the remainder are flattened. The fifth pair have double termina-
tions. |
Our best thanks are due to Dr. L. Gedoelst, of Brussels, for kindly obtaining for us the loan of
the type-material of Kathleena arcuata, Gedoelst, 1916. the property of the Congo Museum at Tervueren.

One of us (Baylis, 1920 a) had already suggested that this form was probably identical with Contra-
caecum microcephalum (Rud.), and our examination of the material confirms this view.

Contracaecum incurvum (Rud., 1819) (?).
Syn. Ascaris incurva, Rud.
(GE 7er)
Two male individuals of an Ascarid from the stomach of the peacock fish

(Histiophorus gladius) are probably referable to this species, though they are small
and perhaps not fully mature. The characters of the oesophagus and of the head

1922.] H. A. Bayzis AND R. DAUBNEY : Parasitic Nematodes. 283

show them to belong to the genus Contracaecum. The head corresponds fairly well
with the figures given by Schneider ((1866), pl. II, figs. rr @ and 6) and by Linton
((Egor), pl. IV, figs. 29, 30), while the
presence of a long intestinal caecum has
been noted by Dujardin (1845) and by
Linton (l. c.), although the oesophageal
appendix was not observed, and Dujar-
din placed the species among the forms
now considered to belong to the genus
Porrocaecum.

The characters of the caudal end in
the male do not appear to have been
accurately described. Stossich ((1902),
pl. III, fig. 1) shows only two pairs
of postanal papillae, and a peculiarly Fia. 17.—Contracaecum incurvum. Head of male ;
shaped tail. Linton’s figure ((1901), pl. rs Dre

; a., cervical ala; g., groove in cuticle; 2., interla-
IV, fig. 32) is scarcely adequate for deter- bium; p., papilla.
mination. For these reasons it may be
worth while to give some details and figures of the anatomy of the present specimens.

The larger of the two specimens measures 34°3 mm. in length and 0°55 mm.
in thickness. The diameter of the head is about 0'I9 mm. The distance from
the head end to the posterior end of the oesophagus (including the small spherical
ventriculus) is about 3 mm. The ventriculus is 0°16 mm. in diameter. The oeso-
phageal appendix is relatively very long (3 mm.), and the intestinal caecum runs
forward to a point 1:05 mm. from the head end. The cervical alae originate just
dorsally to the lateral interlabia, and extend back to a point about 2 mm. from the
anterior end. They are about 0:05 mm. wide at the widest part. The body is rela-
tively slender and of almost uniform thickness throughout. The cuticular striations
are coarse (up to 070125 mm. apart), and form small saw-teeth in optical section.
The lips have sinuous margins anteriorly and laterally, and are produced into broad
cuticular flanges at the posterior corners. There are deep grooves running round the
bases of the lips from the interlabia, similar to those characteristic of the genus
-Ophidascaris. The interlabia are rather short and compressed between the lips.
The ventro-lateral lips are somewhat asymmetrical in shape, though hardly so much
so as is indicated by Schneider’s figure ((1866), pl. II, fig. 11 6), the ventral angles
being considerably produced. The dorsal lip bears a pair of rather small, rounded
papillae; the ventro-lateral lips one each, towards the ventral side. There is a pair
of conspicuous, but sessile, cervical papillae situated dorsally to the cervical alae and at
0°65 mm. from the anterior end. The nerve-ring is at 0°55 mm., and the excretory
pore at 068 mm., from the anterior end.

The tail of the male (fig. 18) is o‘2 mm. long, sharply tapering, curved ventrally
and drawn out at the tip into a slender spike. There are no caudal alae. The ven-
tral surface of the caudal region, from about 0°7 mm. in front of the cloaca forward

284 Memoirs of the Indian Museum. [Vor. VII,

for about 2 mm., has the cuticle raised into pronounced longitudinal ridges, inter-
rupted by transverse grooves at intervals of 0°03 mm. The spicules are equal in
length (4° mm.) and have broad alae,
except for a short distance at the tip.
The caudal papillae are all rather small
and sessile. There are about 15 pairs
of preanal papillae, those near the clo-
aca small and close together, the more
anterior gradually becoming larger
and wider apart. There is also a pair
of double adanal papillae, and four
postanal pairs, of which two are ven-
tral and two lateral.

The host in which the adult form of this

species has been commonly recorded is the
sword-fish, Xiphias gladius,! and the worm ap-

pears to be very widely distributed and to
Fic. 18.—Contracaecum incurvum. Tail of male; later- attain a large size. Linton (1901) has also
al view. | recorded immature stages, probably of this
species, from several other fishes In the pre-
sent collection there are included some encapsuled larvae from the mesentery of Nandus marmora-
tus and Wallago attu, and from the body-cavity and peritoneum of another (unnamed) fish, which show
the same oesophageal structure as the specimens from Histiophorus, and are perhaps to be referred to
the same species The largest of these larvae is about 33 mm. long. The oesophagus does not yet
exceed 3 mm. in length, and the oesophageal appendix 0:5 mm. The intestinal caecum is already of
considerable length. The lips are not yet formed, so that it is impossible to confirm the determination
by a study of their stucture.

Contracaecum tricuspe (Gedoelst, 1916).

This species was described by Gedoelst from an African heron. We have to
record its appearance at Calcutta in the Indian darter or snake-bird (Plotus melano-
gaster).

Contracaecum engonium, sp. nov.
(Figs. I9, 20.)

A single male specimen was collected from the black stork (Ciconia nigra). It
measures 13 mm. in length and 0°57 mm. in maximum thickness. The head
measures 6°19 mm. in diameter, and is constricted off from the body. The interlabia
are simple and undivided at the tip. The dorsal lip (fig. 19) is rounded anteriorly
and carries a pair of double papillae. The pulp roughly follows the shape of the lip,
but is indented on its anterior edge. Each lip is provided with a pair of flattened
processes springing from the internal surface and projecting anteriorly like two small
horns at the shoulders of the lip. The muscular portion of the oesophagus measures

| Histiophorus gladius, though probably related, not distantly, to Xiphias, is not the same fish, and appears to
be a new host.

1922.] H. A. BayLıs AND R. DAUBNEY: Parasitic Nematodes. 285

2°75 mm. in length and 0'I4 mm in thickness. The intestinal caecum is broad, and
reaches to within 0°69 mm. of the head-end. There is a short ventriculus, measuring
about 0‘14 mm. in length and about as
broad as long. From this is given off a
posterior caecum which is o‘7 mm. in
length and 0‘15 mm. in thickness.

The spicules are equal, long and slen-
der, measuring 1°8 mm. in length and
0'022 mm. in breadth. They consist of
a cylindrical shaft with narrow lateral
alae. The cloaca is situated at 0'125
mm. from the tip of the tail (fig. 20), ane mn engonium. Head of male:
which is abruptly attenuated to a coni- i., interlabium.
cal point. There are ten pairs of post-
anal papillae, of which the first, fourth, fifth and seventh are latero-ventral. The
remaining six pairs are lateral and pedunculate, and fall into two groups: a group

Fia. 20.—Contracaecum engonium. Tail of male ; ventral view.

of two pairs (the second and third) close to the tip, and a group of four pairs extend-
ing from about the middle of the tail almost to the cloaca.

Contracaecum schizothoracis, sp. nov.
(IMG, Bit, 22)

Host: Schizothorax zarudnyi.* Position: intestine. Locality: Hamun-i-Hel-
mand, Seistan, Eastern Persia.

This is a relatively short and stout species, tapering to a considerable degree at
each end. The male is 16°75 mm. long and o°8 mm. thick; the female 20°2 mm. and
1°0 mm. respectively. The cuticular striations are 4-5, apart. The diameter of
the head is 0°2-0'22 mm. The lips (fig. 21) are small, with a deep indentation in the
middle of the anterior margin, cuticular flanges at the sides, and a projection on the
inner surface at each anterior angle. The dorsal lip bears two large, lozenge-shaped
papillae, the ventro-lateral lips one each towards the ventral side. The interlabia

286 Memoirs of the Indian Museum. [Vou. VII,

(fig. 21, 2.) are almost as long as the lips, and are bifurcate at the tip. There are no
cervical alae. The distance from the head-end to the posterior end of the oesophagus
(including the small, almost globular ventriculus) is 22-25 mm. The ventriculus
measures 0°18 mm. in length and 0'24 mm. in width. The oesophageal appendix is

o-1mm.
Fia. 21.—Contracaecum schizothoracis. Head of female; dorsal view.
i., interlabium.

about 0°6-0'7 mm. long, and the intestinal caecum runs forward to a point about 0°5
mm. from the head-end. The prominent cervical papille and the nerve-ring are
situated at 0'3-0'4 mm. from the anterior end. The position of the excretory pore
was not made out.

Fra. 22.—Contracaecum schizothcracis. Tail of male ; lateral view.

The tail in both sexes is short and bluntly conical. In the male, it is 0°12 mm.
long and has no alae. The spicules are very long (at least 5°5 mm.) and provided
with broad alae. Their length could not be accurately measured owing to the whole
of the extruded portion being thrown into spiral coils. Their dorso-ventral diameter
is about 0'035 mm. There is a regular series of about 23 pairs of small preanal

1922.] H. A. BayLıs AND R. DAUBNEY: Parasitic Nematodes. 287

papilla. The postanal papille are arranged in five pairs, of which the most posterior
is lateral, the rest ventral. The two anterior pairs are at almost the same transverse
level, just opposite to the posterior lip of the cloaca.

The tail of the female is 0°23 mm. in length. ‘No caudal papillae were seen.
The vulva is situated a little behind the anterior fifth of the body (at 4°3 mm. from
the anterior end). The simple muscular vagina, after a preliminary coil anteriorly,
pursues a very irregular course posteriorly to about 15 mm. from the vulva before
the origin of the uterine branches. These are about 8 mm. in length, their posterior
ends serving as receptacula seminis. The ovarian tubes appear to double upon them-
selves some distance in front of the anus, and return towards the anterior end. The
ova are nearly spherical, with a thick shell measuring 0'0575-0'0725 mm. in
diameter.

Genus Amplicaecum, Baylis, 1920.
Amplicaecum varani, sp. nov.
(Figs. 23, 24.)

A few specimens of an Ascarid which appears to belong to the genus Amplicaecum
were collected on one occasion from the intestine of Varanus salvator in the Zoologi-
cal Garden. The only Ascarid hitherto recorded in Varanus,! so far as we are able
to discover, is Ophidascaris filaria, which
is, however, usually found in pythons.
This is a much larger species, and could
not be confused with the present form.

There were several adult males, but
unfortunately only one fully mature
female. The measurements in the fol-
lowing description were taken from this
female and the three largest males.

The male measures 22°2-24'9 mm. in
length, the female 24°75 mm. The great-
est thickness is 0°73 mm. in the male, 0°8

SE=
L_otmm |
f ; Fig. 23.—Amplicaecum varani. Head of female ;
mm. in the female. The diameter of dorsal view.

the head is 0'25-0'29 mm. The cuticular dr., dentigerous ridge; g., grove in cuticle; ¢.,

iati interlabium ; p., papilla.
striations are fine (about 0'005 mm. ""erabtum; P» Pap

apart). The lips (fig. 23) are nearly square in shape, and have a deep indentation
on the inner surface at the anterior margin. The interlabia are very small and almost
hidden by thelips. From the interlabia well-marked semicircular grooves in the cuticle
run round the bases of the lips, nearly meeting in the median line of each lip. These
grooves have upstanding membranous cuticular borders posteriorly. The dorsal lip has
two moderately large papillæ. Fach ventro-lateral lip has one large, lozenge-shaped
papilla towards the ventral side, and one very small papilla laterally. The dentiger-

1 See Baylis (1921).

288 Memoirs of the Indian Museum. [Vor. VII,

ous ridges are weli-developed and marginal. The oesophagus is without bulb or
ventriculus, and measures 3°5-4°7 mm. in length. A well-developed, but rather
narrow, intestinal caecum, 0'9-I’O mm.
long, is present, running forward beside
the oesophagus. The very small cervical
papillae are situated at about 0°9 mm.
from the anterior end, the nerve-ring at
0'7-0'74 mm., and the excretory pore at
0° mm., from the same point.

In the male, the conical tail (fig. 24)

is only o'16 mm. long, and there are no

caudal alae. The two equal spicules are
remarkably short (0°5 mm.) and are sim-
ple, cylindrical, slightly tapering rods.
There are some 32 pairs of preanal papil-
lae, those nearest to the cloaca being very

small, the more anterior much larger.
; In addition to these there is one small,
Fic. 24.—Amplicaecum varani. Tail of male; 2 : À : 5

arena “ew. - sessile, median papilla on the anterior lip
c., cloacal aperture. of the cloaca. The postanal papille are
_ arranged in five pairs, of which the first

and third from the tip of the tail are ventral, the rest lateral, in position.
In the female, the tail is conically pointed and 0°32 mm. long. There is a pair
of caudal papillae at 0°065 mm. from the tip. The vulva is situated at 6:5 mm.
from the anterior end of the body, 7.e., a little behind the anterior quarter. There is
a long muscular vagina, following a very irregular course in a generally posterior
direction, but with occasional forward loops. The two uterine branches are wide and
thin-walled, and run backward with a rather sinuous course. The coils of the ovaries
occupy the posterior region of the body, as far back as about 1°5 mm. from the tip
of the tail. The eggs are oval, with a rather thin shell, measuring 0:0675-0°075 X

0°05 mm.
Genus Dujardinia, Gedoelst, 1916.
Dujardinia helicina (Molin, 1860).'

We refer to this species two immature females from the stomach of Crocodilus
porosus trom Port Canning, Gangetic Delta. The specific determination is possibly

1 Gedoelst (1916) describes a form, from an African crocodile, which he identifies with A. helicina, Molin, erecting for
it anew genus Dwardinia. Skrjabin (1916) also describes what he believes to be A. helicina, Molin, from an African
crocodile, and proposes for it the new genus Trispiculascaris. Travassos (1920) considers both Gedoelst’s and Skrjabin’s
species distinct from A. helicina, Molin, and renames Gedoelst’s form Dujardinia dujardini, and that of Skrjabin Tris-
piculascaris trispiculascaris. As, however, this author gives no morphological reasons for his views, we are unable to
discuss them. Examination of African material existing in the British Museum, and already regarded as Dujardinia
helicina (Molin) of Gedoelst, shows that the oesophageal and intestinal structure described by Gedoelst is present.
This is definitely stated by Skrjabin to be absent in his material. On the other hand, an accessory piece similar to that

1922. | H. A. BayLıs AND R. Dausney: Parasitic Nematodes. 289

open to doubt. Ascaris helicina was originally described from Crocodilus acutus' in
America, but has also been recorded from Africa in Crocodilus niloticus more than
once, though not hitherto from an Indian crocodile.

Larvae of Anisakinae.

Immature Ascarids of various sizes (the longest measuring about 18 mm.)
occurred under the peritoneum of the fish Pelamys chiliensis. They have a long
ventriculus, and may be the larvae either of an Anisakis or of a Porrocaecum. If
the latter, the intestinal caecum has not yet been developed.

Family HETERAKIDAE, Railliet and Henry, 1914.
Subfamily HETHRAKINAE, Railliet and Henry, 1912.
Genus Heterakis, Duj., 1845.
Heterakis papillosa (Bloch, 1782).
Syn. À vesicularis (Frölich, 1791).
The collection contains examples of this species from the following hosts:

Ring-necked pheasant (Phasianus torquatus*).
Common hill-partridge (Arboricola torqueola).

Heterakis isolonche, v. Linst., 1906.
Hosts:
Crimson horned pheasant (Tragopan satyra).
Monal (Lophophorus impeyanus).
Blood pheasant ([thagenes cruentus).

An accurate description of this species has been given by Lucet and Henry
(I9II). The remarkable cuticular “ papillae’? usually present in the neighbourhood
of the vulva of the females are not constant in number or position, and some indivi-
duals have none. Moreover, they correspond very closely in diameter with the inter-
nal diameter of the preanal sucker of the male. We are therefore inclined to believe
that they are actually caused by the action of the sucker of the male in attempting
to copulate. It may be that the cuticle of this part of the ventral surface of the
female is soft and readily drawn into the sucker. In any case, these raised “hold-

described by Skrjabin is also present. The spicules are of great relative length and very slender. Gedoelst gives the
length of the spicules as 280u, and does not mention an accessory piece. 280u is slightly less than the length of the
accessory piece in our material, and we suggest that this structure has been mistaken for the spicules. . We are therefore
inclined to believe that our material and Gedoelst’s belong to the same species, but that the form described by Skrjabin
is distinct, unless the important structure of the alimentary canal has been overlooked. 8 The structure of the oesopha-
gus and the presence or absence of diverticula of the alimentary canal are characters which we regard as of consider-
able systematic value among the Ascaridae; but, as has been shown above (see Porrocaecum reticulatum) an accessory
piece may occur in an isolated species of a genus in which it is normally absent, and we do not feel that it is of equally
great systematic importance.

We wish here to acknowledge our indebtedness to Prof. R. T. Leiper for kindly assisting us to consult Skrjabin’s
paper.

1 More correctly, C. americanus.

290 Memoirs of the Indian Museum. [Vor. VII,

fasts’”’ would be very effective in assisting close union if, as we suggest, they are em-
braced by the sucker.

Heterakis longecaudata, v. Linst., 1879.

This species, which is very closely related to the genotype, H. papillosa, appears
to have been recorded up to the present only in its type-host, M egacephalon maleo,
a bird of the Megapodiid family, found in Celebes. It is not improbable, however,
that its exceedingly close resemblance to H. papillosa has led to its being often con-
fused with that species when found in other hosts. In the Zoological Garden, Calcutta,
it occurred in the following birds :—

Monal (Lophophorus impeyanus).

Crimson horned pheasant (Tragopan satyra).
Swamp-partridge (Francolinus gularis).

Red spur-fowl (Galloperdix spadicea).

In the two first-mentioned hosts it appears to occur not uncommonly, though in
small numbers, if we may judge from the small amount of material available. In
the horned pheasant it sometimes occurred together with H. bosia or H. isolonche, or
both. In the monäl both longecaudata and isolonche occur, but we have not found
them together.

It may be useful to amplify somewhat the short original description given by von
Linstow (1879). The measurements here given are based on examples from the
monal.

The length of the male is from 7:9 to 9'I mm., that of the female 7-9 to 9°6 mm.
The maximum thickness, measured dorso-ventrally, is 0'3-0'4 mm. The cuticular
striation is so fine as to be scarcely visible in some specimens. The anterior end of
the worm is usually curved towards the dorsal side. The three lips are simple and
very similar to those of H. papillosa. The diameter of the head at the base of the
lips is 0'068 mm. There are well-developed lateral alae, commencing at a short dis-
tance from the head, and running throughout the greater part of the length of the
body. At 0°55 mm. from the anterior end a pair of small cervical papillae project
into the alae. The oesophagus, measured from the anterior end of the worm to the
back of the bulb, is r-o-1'r mm. long. It commences with a small anterior section,
or “pharynx,” narrower than the oesophagus proper and 0-08-0709 mm. long. Pos-
teriorly the oesophagus passes gradually into a large, pear-shaped bulb, o‘18-0:22 mm.
in diameter, and containing a well-developed valvular apparatus. The nerve-ring is
situated at 0°3 mm., and the excretory pore at 0°45-0°5 mm., frum the anterior end.

In the male, the caudal end is provided with very broad alae. The tail proper
(2.e., the postanal portion) measures 0°45-0°5 mm. in length, and tapers beyond the
alae to a fine filament. The preanal sucker measures 0‘08-0‘09 mm. in diameter, and
is situated at o-I-0'15 mm. from the cloaca. Both spicules are alate, the alae of the
short left spicule being very broad, those of the long right spicule much narrower.
The shorter spicule has a double bend near the tip. like that of H. papillosa, while the

1922. | H. A. BayLıs AND R. DAUBNEY : Parasitic Nematodes. 291

tip of the longer spicule is simple and straight. The lengths of the spicules are
2°38 mm. and 0°75 mm. respectively in a large specimen. The twelve pairs of caudal
papillae are arranged in the manner indicated by von Linstow, with the exception
that of the two pairs nearest to the posterior end one is situated laterally, the other
(slightly larger) ventrally, and both are at an equal distance from the tip of the tail.
In number and arrangement there is therefore no difference from H. papillosa.

In the female, the tail is straight and gradually tapering, and measures I’I-I'2
mm. in length. There is a conspicuous pair of caudal papillae at about 0°68 mm.
from the posterior end. The vulva is situated very slightly behind the middle of the
body (not in front of it, as stated by von Linstow), viz., at 3°7-4°75 mm. from the
posterior end. The vagina is long, and pursues a complicated course almost precisely
similar to that of H. papillosa. The terminal portion (ovejector) runs posteriorly
from the opening. The tube then makes a sharp turn anteriorly, then a curve to the
right and dorsally, then bends posteriorly, and from this point runs straight back to
a point about 1°5 mm. from theanus. Here it doubles upon itself, and at about 1 mm.
behind the vulva gives off the two apparently opposed uteri. The greater part of the
coils of both ovaries lie in the anterior portion of the body, between the vulva and
the posterior end of the oesophagus. The ova are somewhat oblong, with a thick
shell, which is slightly dimpled at each pole.
They measure about 0°075 X 0'0425 mm.

Perhaps the most reliable character by
which this form can be distinguished from A.
papillosa is the larger size of the preanal sucker
in the male—(outside diameter 0°06—0'07 mm. in
papillosa, 0:08-0°09 mm. in longecaudata). The
spicules are also a little longer, the right spicule
being longer than that of any other species
recorded in Galliform birds. The preanal sucker
is, as a rule, situated somewhat nearer to the
cloacal aperture than in H. papillosa, and the
caudal alae of the male are broader. On placing
examples of the two species side by side, the
males are fairly readily separable, but it would
be difficult to find characters by which the
females could be easily distinguished.

Heterakis bosia, Lane, 1914.

This interesting form, the male cf which is Ri io
easily distinguished from that of other species 16. 25- Ascaridia perspicillum. Tail of
= ; ; male; ventral view.
by the peculiar shape ot its left spicule, was found
frequently in the crimson horned pheasant (Tragopan satyra). Although evidently
common, it does not appear to give rise to heavy infections. It occurred once in as-
sociation with H. longecaudata only, and once with both this species and H. isolonche.

292 Memoirs of the Indian Museum. [Vor. VII,

Genus Ascaridia, Duj.. 1845.
Ascaridia perspicillum (Rud., 1803.)

(Fig. 25.)
Hosts:

Common fowl.
Blood pheasant (/thagenes cruentus).
In view of the scarcity of figures of the male tail we furnish a new one
(fig. 25).
Ascaridia columbae (Gmelin, 1790).
(Figs. 26-28.)
Syn. Heterakis maculosa (Rud., 1802), Schneider, 1866.

Specimens which we assign to this species were collected from the Bengal green
pigeon (Crocopus phoenicopterus) on three occasions, and from Phlogoenas luzonica and
other pigeons. Some difficulty was at first experienced in definitely determining these
specimens, owing to the variations not infrequently encountered in the position and

28
Fies. 26, 27, 28.—Ascaridia columbae. Tails of three males, in ventral view, to show variation in
papillae.

number of the caudal papillae of the male. Actually the typical number of papillae
is 14 pairs, which is the number given by von Linstow (1001 a), but the figure of this
author is rather too diagrammatic to show clearly their arrangement.

There are five pairs of distinctly postana] papillae, the third pair of which is
rather ventrally placed, while the remainder are lateral. There is an adanal group of

1922.] H. A. Bayris AND R. DAUBNEY : Parasitic Nematodes. 293

four pairs, one pair of which is large and laterally placed, the remaining three pairs
being small and arranged in a triangle on the ventral surface. There is a series
of three pairs of preanal papillae on the ventral surface between the cloaca and
the posterior margin of the sucker. Near the level of the anterior margin of
the sucker is another pair more laterally placed, and finally there is a pair placed
anteriorly to the sucker. This last pair may be duplicated (fig. 28). Another varia-
tion may be furnished by the presence of an additional pair of papillae in the row
between the sucker and the anus (fig. 26). Not infrequently also, the most posterior
pair of postanal papillae appears to be absent (fig. 28). The spicules are equal and
measure from I'2 to 1°35 mm. in length.

An additional character of A. columbae which has, so far as we are aware, hither-
to escaped notice, is the presence of 26 to 30 pairs of cervical papillae extending back-
wards from near the posterior end of the cephalic alae, the first two or three pairs
being situated in the alae.

The species varies greatly in size, the males in our material measuring from
60 to 70 mm. in length and about 1:1 mm. in thickness; the females from 70 to 95 mm.
and up to 2°5 mm. respectively.

Ascaridia compar (Schrank, 1790).
(Fig. 29.)

This species has been recorded in Caccabis saxatilis,
Coturnix dactylisonans, Coturnix communis, Ortyx virginianus,
Perdix cinerea, Tetrao urogallus, T. lagopus, T. tetrix, Gallus
gallinaceus, Gallus domesticus, Numida meleagris, and Colinus
virginianus.

We have now to record its occurrence in the Chakor
(Caccabis chucar).

The material agrees in all essential features with the
descriptions given by v. Linstow (1899) and by Miiller (1897),
though the figures of the tail of the male given by both these
authors are not quite accurate. Miiller, indeed, describes the
post-anal papillae correctly, but his figure fails to indicate
clearly the number and arrangement of the small papillae near Me 20 orodin Gore
the tip of the tail. It may therefore not be out of place to par. Tail of male; ventral
give in the present paper a new figure (fig. 29). ee

Ascaridia cristata (von Linstow, 1001).
(Fig. 30.)

This species was described by von Linstow from material taken from Balearica
regulorum. We have to record its occurrence in the West African crowned crane
(Balearica pavonina*) and in the sarus (Grus antigone). We propose to amplify some-
what the original description.

294 Memoirs of the Indian Museum. [Vor. VII,

The females measure from 38 to 40 mm. in length and ı'I to 1:2 mm. in thick-
ness; the males about 35 mm. and I'l mm. respectively. The head measures from
0°27 to 0°28 mm. in diameter. The dorsal
lip is shorter and broader than the two
ventro-lateral lips, and carries two papillae.
The oesophagus measures from 2°I to 2°3
mm. in length, and is encircled by the nerve-
ring at a distance of 0:46 mm. from the
head. The excretory pore opens at about
0:69 to 0°77 mm. from the anterior end.
A series of 27 pairs of cervical papillae,
similar to those described in A. columbae
above, extends from a point about 0-9 mm.
from the head backwards for a distance of
6‘o to 65 mm. The anterior pairs are
placed just dorsally to the cervical alae, and
the distance between successive pairs varies
from 0°15 to 0’ 3 mm. The tail of the male
is furnished with 13 pairs of papillae, of
which 7 are postanal and 6 preanal. Von
Linstow (IgoI b) described 7 postanal pairs
and 2 preanal, while Gedoelst (1916) in

ee pte a ee assigning specimens to this species describes
ventral view. 3 pairs of preanal papillae. The arrange-
ment of the papillae in our material is shown
in fig. 30. Gedoelst also mentions that the sucker does not possess the “ unpaired
papilla’ on its posterior border. In the specimens we have examined this structure
was easily detected in some and in others apparently absent. The spicules are long
and slender and measure 0°95 mm. in length and 0'042 mm. in width. They are
alate. The anus is situated at about 0°62 mm. from the tip of the tail.

The vulva of the female is situated about 20 mm. from the anterior end, and is
slightly salient. There is a short transverse vagina. The ova measure 0'085 X 0°058
mm. The tail measures 0°7 mm. in length.

Ascaridia stroma (von Linstow, 1899).

Von Linstow described this species from Grus paradisea. The present collection
furnishes us with specimens from the common crane (Grus communis) and the sarus
(Grus antigone).

Genus Strongyluris, A. Müller, 1894.
Strongyluris chamaeleonis, sp. nov.
(Figs. 31-33.)
Host: Chamaeleon vulgaris * (Zoological Garden, Calcutta).
This is a small species, measuring 6°3 mm. in length in the male, 8-4-8'75 mm. in

1922. ] H. A. Bayrıs AND R. DauBney : Parasitic Nematodes. 295

the female. The maximum thickness is 05-07 mm. The lateral fields are broad,
and of the type characteristic of the genus, consisting of a single row of some 70
large, granular cells with clear, rounded nuclei. There are no lateral alae. No cer-
vical papillae have been detected, nor do the longitudinal rows of small papillae on
the body, which occur in some species, appear to be
present in either sex. The cuticular striation is
exceedingly fine. The diameter of the head is about
0‘06 mm. There are three distinct lips, of somewhat
elongate shape, each terminating anteriorly in a flat-
tened lobe consisting only of cuticle, which, seen
in profile (fig. 31), gives the lip the appearance of
ending in a kind of curved tooth or spine. Each
lip bears a relatively large papilla on the outer
surface of its basal portion. The oesophagus, as
has been observed in some other members of the
genus, is marked off into a narrow anterior portion,
or pharynx, the lumen of which describes a peculiar
ventral bend posteriorly; and a wide posterior
portion, the oesophagus proper, ending posteriorly
in a well-developed bulb. The distance from the
anterior extremity of the lips to the posterior end of
the oesophageal bulb is about I’I mm. in the male,
1:45 mm. in the female. Of this the pharynx oc- Fig. 31.—Strongyluris chamaeleonis.

cupies 0°18-0-22 mm. The bulb is almost spherical, Anterior end of female; lateral view.
d.l., dorsal lip; n.r., nerve-ring ;
ph., pharynx.

UNUT-0

measuring 0°2-0'25 mm. in both antero-posterior
and transverse directions. The nerve-ring is situated
at 0°37-0°39 mm., and the excretory pore at 0°6-0°85 mm., from the anterior end.

The caudal end of the male (figs. 32, 33) is obliquely truncate, terminating in a
small, conical spike. Anteriorly to this there are broad alae, forming an almost
circular bursa-like expansion. Near the anterior limits of the alae there is a rounded
sucker with chitinous ring, measuring 0°09 mm. in outside diameter, and having its
aperture somewhat posteriorly directed. There is a little depression in the posterior
edge of the chitinous ring, as in Heterakis. This has been described as a papilla in
some species. The two equal spicules, which measure I’I mm. in length and 0°0275
mm. in maximum thickness, are covered externally with rather coarse granulations,
and taper gradually from their bases to slender points. No chitinized accessory piece
appears to be present. There are apparently nine pairs of caudal papillae, of which
seven project more or less laterally into the alae, while two are situated ventrally
behind the cloacal aperture. As only one male was available, and in this specimen
the spicules were extruded, it is not certain whether any further ventral papillae may
have been hidden from view by them. Of the laterally-placed papillae, two relatively
small pairs are close to the tail-spike, the most posterior being directed more ventrally,
the second more dorsally. The next two pairs are very close together and rather

296 Memoirs of the Indian Museum. [Vor. VII,

Fia. 32 —Strongyluris chamaeleonis. Posterior
end of male; lateral view.

slender. The remaining three pairs form a
groupon either side of the sucker, decreasing
in size from behind forwards. The most
posterior of these three pairs is very mas-
sive. The length of the tail (2.e., from the
cloaca to the posterior extremity) is about
o'13 mm., of which the terminal spike
measures 0°06 mm.

In the female the short, conical tail
measures 0°3 mm. in length, and bears a
pair of small papillae at 0°14 mm. from the
tip. The vulva is situated at 3°0-3°3 mm.
from the posterior end. The vagina is long,
slender, and pursues a rather tortuous
course, the general direction of which is
posterior from the vulva. The branches of
the uterus are parallel, running at first
posteriorly to within a short distance of the
anus, then returning towards the anterior
end. The coils of the ovaries are situated in
the anterior half of the body. The eggs are
oval, with a thick shell, slightly flattened
externally and thickened internally at each
pole. They measure about 0'0875 X 0°055

mm. When ready for laying the content of the egg is still unsegmented and coarsely

eranular.

Of the species referred to the genus Strongylu-

ris, two, S. sonsinoi (v. Linst., 1894) and S. elegans
(Gendre,. 1909), occur in chamaeleons, the former
in the same chamaeleon as the present species. S.
sonsinot, however, differs widely from our form,
and from all others except S. campanula (v. Linst.,
1899), in the elongate and conical shape of the tail
in the male and in the possession of small, sessile,
caudal papillae instead of the typical elongate,
ray-like papillae. The present species is more
closely related to S. elegans, but differs from it in
its smaller size and the much longer spicules of
the male, besides other details; and we are equally
unable to identify it with any of the known species
parasitic in lizards.

As regards the systematic position of Strongyluris, Seurat
(1917), in opposition to the view taken by most authors,

Fie. 33.—Strongyluris chamaeleonis.
Posterior end of male; ventral view.

1922. ] H. A. Bavzis AND R. DAuBnEy: Parasitic Nematodes. 297

regards it as related rather to the Oxyuridae than to the Heterakidae. His contention is based chiefly
on the characters of the lateral fields and of the body-muscles. The other points mentioned (presence
of lateral alae on the body, absence of caudal alae in the male), are clearly not universal characters of
Strongyluris as at present constituted. On the other hand, the structure of the preanal sucker, which
is exactly similar to that of Heterakis, is a character probably quite as important as the lateral fields :
while the arrangement of the musculature does not appear in all cases to be a reliable guide to clas-
sification. Travassos, in a recent paper (1920 [?] ) has suggested placing Strongyluris in a new subfamily,
Spinicaudinae, of the Heterakidae. Railliet and Henry (1914), regarded it as a subgenus of Heterakıs.
We prefer to treat it as a genus, with close relationships to Heterakis, and reserve judgment on the
question of including it in a separate subfamily. The following tabular arrangement of the species
shows that there are two well-marked groups within the genus as hitherto constituted, these groups
being characterized chiefly by the presence or absence of caudal alae in the male.
A. Tail of male without alae.
a. ‘Tail long, straight and tapering. An accessory piece present.
S. sonsinoi (v. Linst., 1894).

S. campanula (v. Linst., 1899).
b. Tail obliquely truncate ventrally, but with an elongate

terminal cone. An accessory piece present.

S. icosiensis, Seurat, 1917.
B. Tail of male with bursa-like alae and obliquely truncate.
Accessory piece absent.

S. brevicaudata, Miller, 1894.
S. paronai (Stossich, 1902).
S. elegans (Gendre, 1909).

S. chamaeleonis, sp. nov.

S. ornata (v. Linst., 1897) and S. streptoesophageus, Conral, 1912, are probably synonymous
with S. brevicaudata.

It seems justifiable to restrict Strongyluris to the forms (B) which agree with its genotype,
S. brevicaudata, in the characters mentioned; while we propose to erect a new genus, Sonsinia, to

include the non-alate forms, with S. sonsinoi as genotype. S. icosiensis appears to occupy a somewhat

intermediate position, but for the present may be referred to Sonsinia.

Genus Pseudaspidodera, nov.
Pseudaspidodera pavonis, sp. nov.
(Figs. 34-37.)

Hosts: Burmese peatowl (Pavo muticus) and ‘ white peafowl’’ (Pavo cristatus).

This is a small worm, the male measuring about 6 mm. in length, the female 7
mm. The greatest thickness is about 0°25 mm. in the male, 0:3 mm. in the female.
The cuticular striations, if present, are too fine to measure. The head (fig. 34) is
ornamented with “cordons’’ resembling those of Aspidodera, opening in pairs at the
interlabial spaces, and consisting of tubular grooves running below the surface of the
cuticle, with a narrow external opening along their length. The members of each
pair of cordons diverge at once and, after running back for a short distance, turn
forward, each on to the outer surface of one of the three lips, where, instead of joining
the corresponding member of the next pair, as in Aspidodera, it ends separately. The
diameter of the head at the posterior limit of the cordons is about 01 mm. Narrow

298 Memoirs of the Indian Museum. [Vou. VIi,

lateral alae run down the body from a little in front of the nerve-ring nearly as far

as the tail. The oesophagus is muscular throughout, 1°4-1°48 mm. long in the male,

I'5-I°6 mm. in the female. It is divided a little behind the head into a very short

anterior portion, and a long posterior portion which ends in a well-developed pyriform

bulb. At the division of these two portions

Po ae there appears to be some kind of valvular ap-

= paratus. The bulb measures 0‘25-0°26 mm.

in length and 0'I7-0'I9 mm. in diameter

transversely, and contains the usual valves.

The nerve-ring is situated at 0°4-0°46 mm.,

and the excretory pore at 0°6-0'65 mm., from
the anterior extremity.

In the male, the tail (figs. 35, 36) which
is 0'38-0'43 mm. long, is provided, for rather
less than the anterior half of its length, with
wide alar expansions, into which some of the
caudal papillae project. The remainder of the
tail is simple and slender, ending in a fine,
tapering point. There is a circular preanal
sucker, 0°I2—0'I13 mm. in diameter, with well-

Fic. 34.—Pseudaspidodera pavonis, Head of developed chitinous wall, situated at 0°15-0°17
femaie; lateral view. mm. in front of the cloaca. The greatest
En cordon ; c’., optical section of same; d.l., diameter (antero-posterior) of the opening of

orsal lip ; p., papilla. Ä

the sucker is about 07°07 mm. The two
spicules are very unequal and dissimilar. The right spicule is slender and simple,
measuring 0°78 mm. in length. The left is provided with broad alae at the sides, has
a barbed tip, and is only 0°45 mm. long. There is no accessory piece. There are
twelve pairs of caudal papillae, the arrangement of which can be understood most
readily by reference to the figures. Three pairs, of which the middle pair is more
ventrally situated and slightly larger than the others, form a group just in front of the
filamentous portion of the tail. The fourth pair is solitary, projecting laterally into
the alae. There is an adanal cluster of papillae consisting of four more or less lateral
pairs with long stalks, and two small, sessile, ventral pairs, one in front of and one
behind the cloaca. Of the four lateral pairs the most posterior is the stoutest, and
projects laterally. The next is more ventrally directed. The next is again lateral ;
while the most anterior of the group projects ventrally. There are two very slender
and long-stalked papillae on either side of the sucker.

In the female the tail is long and straight, tapering to a slender point. It
measures 1°0-1°02 mm. At about the middle of its length there is a very minute
pair of caudal papillae. The vulva is situated behind the middle of the body, at
about 3 mm. from the posterior end. It leads into a vagina (fig. 37) which is convoluted
in a characteristic manner—running forward at first, as a strongly muscular ovejector,
it curls first in a semicircle so as to return towards the body-wall on the ventral side.

MUL 7-0

1922.] H. A. Bayrıs AND R. DAUBNEY: Parasitic Nematodes. 299

Then, taking a turn to the right, and dorsally again, it doubles back upon itself. On
reaching a point just behind the level of the vulva the character of the walls changes,

MU £0

03mm.

Fic. 36.—Pseudaspidodera pavo-
nis. Posterior end of male ; ventral

Fig. 35.—Pseudaspidodera pavonıs. view.

Posterior end of male; Jateral view.
L., left spicule; r., right spicule.

l., left spicule (protruded): r.,
right spicule.

the circular coat of muscles being much less strongly developed. The tube runs back
from this point quite straight to a distance of about 0°8 mm. behind the vulva. Here

it doubles upon itself again, and at
about 0°15 mm. behind the vulva gives
off the two opposed uteri. As in
Heterakis, the two oviducts, doubling
upon themselves in the anterior and
posterior halves of the body respect-
ively, return and cross each other so
that the coils of the ovary belonging
to the anterior uterus are disposed in
the posterior half of the body, and
those of the other ovary in the anteri-
or half. The ova are relatively large,
of somewhat oblong shape, with a thin

olmm. Cc
Fic. 37.—Pseudaspidodera pavonis. Vulva and
vagina of female, in lateral view.
c. plug of cementin vulva. The arrow points in the
direction of the head.

300 Memoirs of the Indian Museum. [Vou. VII,

shell measuring about 0°07 x 0°04 mm., and usually showing a slight internal thickening
at one pole. One end of the shell, as seen in ulero, is occasionally drawn out almost
to a point. The content of the egg is unsegmented at the time of laying.

The characters of this form are such that it appears to form a link between the genera Heterakis
and Aspidodera. It has cephalic ‘ cordons ” similar to, but rather less highly-developed than, those of
Aspidodera ; while it possesses long, pedunculate caudal papillae in the male, like those of Heterakis in
shape and arrangement, and unlike the more sessile papillae of Aspidodera. The markedly dissimilar
spicules and the absence of an accessory piece are also characters ot Heterakis rather than of Aspido-
dera.

Two species of Heterakis have been recorded in peafowl—d. papillosa (Bloch) and H. hamulus, v.
Linst., 1906. The former is, of course, a well-known species and the genotype of Heterakis. Although
the description of H. hamulus is rather brief, it appears sufficient to prevent the identification of the
present form with that species.

Subfamily SUBULURINAE, Travassos, IQI4.
Genus Subulura, Molin, 1860.
Subulura sarasinorum (Meyer, 1896).

This species occurred in the intestine of a slender loris (Loris gracilis) ! in the
Calcutta Zoological Garden. .

Subulura galloperdicis, sp. nov.
(Fig. 38.)

This species was collected from the intestine of the Red spur-fowl (Galloperdix
spadicea).

The female measures 11:5 to 12°5 mm. in length and about 0°4 mm. in thickness;
the male 9°5 to 10 mm. and 0°3 mm. respectively. The head is small, measuring
about 0:08 mm. in diameter. There are narrow cephalic alae which extend to a dis-
tance of about I mm. from the anterior end. The buccal cavity is 0°06 mm. deep,

about 0'023 mm. wide at the anterior end, and 0‘031 mm. at the posterior end.
There are three triangular teeth at the base of the buccal cavity, two sub-dorsal, and
one ventral and median. The height of the teeth is about 0'013 mm. The nerve-
ring is situated at 0°27 mm. from the head, whilst the excretory pore opens on the
ventral surface at 0°45 mm. from the head. The oesophagus consists of a long mus-
cular portion with the usual prebulbar swelling, and a large bulb containing the grind-
ing apparatus. The anterior portion is I’5 mm. in length, and the prebulbar swelling
0°14 mm. in thickness. The bulb is roughly spherical and has a diameter of o'2 mm.
The vulva is situated in the anterior half of the body, dividing the latter in the ratio
of 3: 4. There is a short transverse vagina from which large, well-developed ovejec-
tors run anteriorly and posteriorly. The ovaries commence at the anterior and pos-
terior bends of the uteri, and terminate in the vicinity of the vulva. The bend of
the anterior genital tube is at about 0°42 mm. from the oesophageal bulb, while that
of the posterior is at about 0°22 mm. from the tip of the tail. The ova measure

| Now called Loris lydekkerianus.

1922. ] H. A. Bayuıs AND R. DausBney: Parasitic Nematodes. 301

0°065 X 0'035 mm. and contain fully-formed embryos. The anus is situated at 1°1 mm.
from the tip of the tail.

The tail of the male (fig. 38) measures 0°21 mm. in length and is drawn out at
the tip into a fine point. There are eleven pairs of papillae, four pairs of which are
preanal, two adanal, and five postanal. The sucker is situated at about 0°65 mm. in
front of the anus, and is spindle-shaped. Of the preanal papillae a latero-ventral pair
is situated on the lateral border of the sucker and towards its anterior margin:
variation in the position of this pair is not uncommon. The remaining three pairs
are placed between the sucker and the anus. The anterior of these is laterally placed
at about 0°15 mm. behind the sucker. The next two pairs are ventral, one being
about 0°07 mm. behind the preceding pair, and the other just anterior to the adanal

SKS
> INN =
ca SS

Fie. 38.—Subulura galloperdicis. Posterior end of male : lateral view.
a.p., accessory piece.

papillae. The two pairs of adanal papillae are placed on the anterior border of the
cloaca, one lateral to the other. Of the postanal papillae, the third pair is quite
lateral, the remaining four pairs ventral. The first and second pairs are small and
close to the tip of the tail. The fifth pair is immediately posterior to the anus, and
the fourth pair is a little less than half-way between the second and fifth pairs. The
third pair, which is lateral, is situated about midway between the anus and the tip
of the tail.

The spicules are equal, long and slender. They measure from 0:76 to 0'8 mm.
in length, and are tapered to a fine but rounded point. They consist of a cylindrical
axis, which measures about 0'0II mm. in diameter, and two alae. Their totai width
is about 0702 mm. The edges of the alae are very finelv serrated. There is an

302 Memoirs of the Indian Museum. [Vor. VII,

accessory piece (fig. 38, a.p.), which is slender and curved. It measures about
0°18 mm. in length and has a spur at about o'06 mm. from its anterior end.

The male genital tube is much coiled and reaches to within 0:7 mm. of the
oesophageal bulb.

The number and disposition of the caudal papillae in the male, the length of the
spicules and accessory piece and the position of the vulva in the female are salient
characters which serve to distinguish Subulura galloperdicis from the other members
of this genus occurring in galliform birds.

There are five members of this group which possess eleven pairs of papillae, viz., S. curvata (v.
Linst., 1883), S. strongylina (Rud. 1819), S. olympioi, Barreto, 1919,! S. halli, Barreto, 1918 ! and
S. seurati, Barreto, 1919.!

S. curvata has unequal spicules measuring 1-2 and 0°9 mm. respectively, and the papillae are made
up as follows :—2 pairs preanal, 2 adanal, and 7 postanal. S. strongylina possesses a chitinous tail
appendage. The papillae consist of 3 preanal, 2 adanal, and 6 postanal pairs. The spicules measure
0'899 mm. in length. S. olympioi has 3 pairs of preanal, two pairs of adanal and six pairs of postanal
papillae. S. halli has five pairs of preanal papillae; and the spicules measure 1:5 mm. in length. In

S. seurati the spicules are unequal and there are 5 pairs of preanal papillae, while the vulva is situated
at the junction of the anterior and middle thirds of the body.

Subulura, sp.

The collection contains two females of a species of Subulura taken from the
Button-Quail (Turnix, sp.) One of the specimens is badly damaged. The complete
specimen measures 14 mm. in length and 0°41 mm. in thickness. The head measures
about 0'08 mm. in diameter. The buccal capsule is 0'035 mm. deep and 0:62 mm.
wide. The usual teeth are present.

Narrow cephalic alae extend as far as the beginning of the prebulbar oesophageal
swelling. The total length of the oesophagus is r°I mm., while the diameter of the
bulb, which is roughly spherical, is 0-15 mm. The nerve-ring encircles the oesphagus
at o'r mm. from the anterior end. The excretory pore opens at 0°4 mm. from the
head in the median ventral line. The anus is situated at 0°32 mm. from the tip of the
tail, which is acutely pointed. The vulva is situated at 6:1 mm. from the anterior
end. The posteriorly-directed vagina is just discernible. For the rest the body of
the worm is completely filled with eggs, reaching anteriorly up to within 0:07 mm. of
the anterior end, and posteriorly to within o‘1 mm. of the tip of the tail. The eggs
are in various stages of development, some containing fully-formed embryos. They
are thin-shelled and measure 0°085 X 0:056 mm.

It is not possible to assign these specimens to a definite species.

Family OXYURIDAE, Cobbold, 1864.
Genus Oxyuris, Rud., 1803.
Oxyuris anthropopitheci, Gedoelst, 1916 (?).
Host: Black-headed lemur * (probably Lemur brunneus).

1 In Barreto’s (1919) monograph of the subfamily Subulurinae, these species are given as ‘‘ S. olympioi, Barreto,
1918,” * S. halli, Barreto, 1917” and “‘ S. seurati, Barreto, 1917.” The names, however, do not appear to have been
published previously to 1919, although the work was in preparation in 1917.

1922 | H. A. BayLıs AND R. DAuBNEY : Parasitic Nematodes. 303

No Oxyuris appears to have been recorded in true lemurs, though two forms,
O. corollatus, Schneider, 1866 and O. coronata, v. Linst., 1903, are recorded in Galeo-
pithecus. It is impossible to identify the present material with either of these, both
of which have characteristic spinous structures on or near the head. Of the forms
found in apes and monkeys, the nearest appears to be O. anthropopitheci, from the
chimpanzee. Both Gedoelst’s material and our own has unfortunately consisted only
of females, so that the characters available for determination are scanty, and we
have been compelled to rely chiefly on measurements. While, therefore, we find a
fairly close agreement between our material and Gedoelst’s (1916) description, the
determination, especially in view of the difference of hosts, can only be regarded as
tentative.

Oxyuris compar, Leidy, 1856 (?).

A single female, from the intestine of a domestic cat in Calcutta, is doubtfully

referred to this little-known and apparently rare species.

Genus Atractis, Duj., 1845.
Atractis dactylura (Rud., 1819). .

Examples of this species, all young females, occurred in association with
Zanclophorus kempi (see p. 312) in the intestine of Testudo elongata at Baradighi,
Jalpaiguri, Bengal.

Atractis opeatura, Leidy, 1891.
Syn. A. cruciata, v. Linst., 1902.

This form occurred in large numbers in the intestine of an iguana* (species not
mentioned) in the Zoological Garden, Calcutta. Unfortunately, the specimens are in
rather poor condition and net quite mature. They show no important differences
from the descriptions furnished by von Linstow ((I90I a) and (1902)) and by Railliet
and Henry (1912), except that there is an additional pair of small caudal papillae,
adanal in position, in the male.

Note.—Travassos (1920 [?]) proposes a family Atractidae, which he considers to belong to the super-
family Rhabdiasoidea (= Angiostomoidea). The included genera are Atractis, Ozolaimus, Rondonia.
Labiduris, Crossocephalus, Macracis, Cobboldina and Cyrtosomum. These are, for the most part, little.
known forms, and in the present unsatisfactory state of our knowledge of the Oxyuridae as a family, and
in the absence of a definition of the family Atractidae, we prefer to adopt a conservative attitude as to
the position of Atractis. The whole question of the relationships of the Angiostomoidea (or Rhabdia-
soidea) is much involved at present; but it may be remarked, in passing. that the definition of the
Angiostomoidea is based mainly on the fact that its species have two heterogenetic generations, and
the parasitic phase is without males. This is not known to be the case with the forms included in
Travassos’ proposed family, though in Alractis dactylura an alternation of generations of a different
kind is said to occur (Macé, 1887). Here the females of the parasitic phase are viviparous, and the
generation to which they give rise is said to consist entirely of oviparous females.

Family KATHLANIDAE, Travassos, 1918.
Genus Falcaustra, Lane, 1915.

The collection includes examples of four species of this genus, all of which appear to
be new. The following known species have been assigned to the genusup to the present :

304 Memoirs of the Indian Museum. MMor-avallr

F. falcata (v. Linst., 1906), from Geoemyda trijuga,

F. lambdiensis, Seurat, 1918, from Clemmys leprosa,

F. siamensis, Baylis, 1920, from Hieremys annandalei ;
while we may also refer to it

F. [Oxysoma] kachugae (Stewart, 1914), from Kachuga lineata.

The hosts of all these species are tortoises. Of the new species to be described,
two are from tortoises and two from a fish Some of them show considerable diver-
gence from the typical characters, and will necessitate some modification of the
generic diagnosis. We do not, however, feel justified in splitting up the genus at
present, as it would be hard to find a clear and satisfactory dividing-line between
the typical and the atypical species.

Taking first the forms that depart least from the characters of the genotype, we
have the following :—

Falcaustra testudinis, sp. nov.
(Fig. 39.)

Host: Testudo elongata. Locality: Assam (Tura, foot of Garo Hills).

This is the smallest species hitherto met with. The male measures 10°2-10°4 mm.
in length and 0°6 mm. in thickness; the female 9'2-11°5 mm. and 0°6-0'75 mm.
respectively. The diameter of the head is 0‘15-0°:16 mm. This is followed by a
slightly narrower neck. The buccal cavity measures about 0°05 mm. in length and
0°03 mm. in diameter. The distinct
anterior division of the oesophagus, to
which we shall hereafter refer as the
‘“pharynx, ” is 0‘13-0°14 mm. long. The
entire oesophagus, from the extremity
of the head to the back of the bulb,
measures 17-2'Imm. The bulb consists
of two swellings separated by a narrow
neck ; its length is 0°4-0°45 mm. and the
diameter of the larger (posterior) swelling
0°26-0°27 mm. The prominent cervical
papillae are situated at 1‘03-1:1 mm., the
nerve-ring at O‘44-0‘45 mm., and the
excretory pore at 1:22-1:26 mm., from

er the anterior end.

a. Se testudinis.— Posterior end of In the male the tail is 0:81 mm. long,
a.p., accessory piece. and there is no preanal sucker-like organ.
The caudal papillae (fig. 39) consist of
the typical eleven pairs and one unpaired preanal papilla. Nos. 1 and 2 are close
together, ventral; No. 3 lateral; No. 4 isolated, ventral; No. 5 ventral; No. 6 lateral,
at about the same level as No. 5; Nos. 7 and 8 at the sides of the cloaca and close
together. The spicules measure 0:8 mm. in length and 0:07 mm. in greatest width.

The chitinized portion of the accessory piece is 0'I5-0'17 mm. long.

1922. | H. A. Bavyzis AND R. Dausney: Parasitic Nematodes. 305

In the female the tail is about 1 mm. long, and the caudal papillae are at 0°5 mm.
from the tip. The vulva is situated at 30-47 mm. from the posterior end. The
vagina is long, running forward for a distance of 2°3
mm. before giving off the uterine branches. The

eggs measure 0°125—-0°137 X 0°075-0'087 mm. \
Falcaustra barbi, sp. nov. iS es
(Figs. 40, 41.) AN
Host: Mahseer (Barbus tor). Locality: Torsa N =
River, Falakata, Eastern Bengal. vo
This species measures 15'2-16°5 mm. x 0°65-0°7 = \\
mm. in the male; 15'5-19°6 mm. X0°65-I°0 mm. in S N =
the female. The cuticular striations, if present, are > SI =
excessively fine. The almost globular head has a => \)
diameter of 0'2-0'22 mm., and is followed by a a DIN

distinct neck. The buccal cavity measures about

0°07 mm. in length, the pharynx o'r mm. The re, AD eme barbi Pose
distance from the anterior end to the end of the terior end of male; lateral view.
oesophagus, including the bulb, is 2'5-2'8 mm. ea musica uckerlibe
The anterior swelling of the bulb is oval in shape

and sharply constricted off from both the preceding portion of the oesophagus and the
rest of the bulb. The dimensions of the bulb are 0°5-0'59 mm. in length and 0'34-

Fie, 41.—Falcaustra barbi. Tail of male; lateral view.
a.p., accessory piece.

306 Memoirs of the Indian Museum. [Vou. VII,

0°37 mm. in diameter. The small, but prominent, cervical papillae are situated at
1°2-1°4 mm., the nerve-ring at 0'4-0'5 mm., and the excretory pore at I’55-I'9 mm.,
from the anterior end.

The tail of the male is about 0°6 mm. long. There is a single preanal, sucker-
like, fan-shaped aggregation of muscles, situated in front of the long series of oblique
caudal muscles (fig. 40). The caudal papillae (fig. 41) are very small and inconspicu-
ous. There are ten pairs and an unpaired preanal papilla. Of the postanal papillae
Nos. 1 and 2 are close together and ventral; No. 3 lateral. There are three more
pairs close together and ventral, and one lateral, just behind the cloaca. The spicules
are 1'I3 mm. long, and 0'I mm. wide dorso-ventrally at the widest part, which is near
the root. The accessory piece is well-chitinized, and measures 0°2 mm. in length.

The tail of the female is 0°65-0°8 mm. long, and carries a pair of inconspicuous
papillae at 0°35 mm. from the tip. The vulva is situated at 6°25-7°3 mm. from the
posterior end. The vaginaisshort (about Imm.) and nearly straight. The eggs are
roundish oval, and measure about 0°075 X0°05 mm.

The intestine of every specimen examined contained large numbers of diatoms.

Falcaustra leptocephala, sp. nov.
(Fig. 42.)

Host: Mahseer (Barbus tor). Locality: Torsa River, Falakata, Eastern Bengal.
This species occurred together with the preceding, in large numbers, in the same
fish. It is easily distinguished from #.
barbi by its very narrow head and the
absence of a neck. It is a large, stout
form, especially as regards the females.
These appear an opaque white in spirit,
owing to the large numbers of eggs in the
uterus. The males, in spirit, remain
semi-transparent. In both sexes the
intestine shows through the body-wall
as a blackish line, and this may be partly
due to the fact that its contents, as in
the case of F. barbi, consist very largely
of diatoms.

length and 1°3 mm. in thickness; the
Fic. 42.— Falcaustra leptocephala. Tail of male: rene me Senn: and ARCS re
lateral view. respectively. The cuticular striations
are about 0'002 mm. apart. The diame-

ter of the head is o‘1-0°12 mm. The buccal cavity is about 0:05-0'06 mm. long,
the pharynx 0°14-0'17 mm. The whole oesophagus, from the head end to the back
of the bulb, measures up to 3°55 mm. The bulb is flask-shaped, having no sharp

The male measures up to 19 mm. in.

«
FA

1922. | H. A. BayLıs AND R. DAUBNEY : Parasitic Nematodes. 307

constriction between the two swellings. Its length is 0°6-0°67 mm., and the diameter
of the posterior swelling 0°45-0'47 mm. The prominent, almost bristle-like, cervical
papillae are situated at 1'3-I'4 mm., the nerve-ring at 0°45-0°5 mm., and the
excretory pore at 2'0-2'I5 mm., from the anterior end.

The tail of the male (fig. 42) measures 0°7-0°85 mm. in length. There is no pre-
anal sucker-like organ. The number and arrangement of the papillae are the same as
in the preceding species (F. barbr) —there being ten pairs and a median preanal papilla.
The spicules are about I mm. long and 0-09 mm. wide. There appears to be no chi-
tinized accessory piece. |

In the female the tailis 1°:1-1°3 mm. long. The caudal papillae are very incon-
spicuous, and are situated at 0-6 mm. from the tip. The vulva is at about II mm.
from the posterior end. The vagina is narrow, and apparently short, but its
course is difficult to trace owing to the dense masses of ova in the uterus. It
runs forward and dorsally from the vulva, keeping close to the body-wall. The
branches of the uterus are wide, nearly filling the body-cavity. The ova are very
much more numerous and considerably smaller than in most of the other species,
and of a much more spherical shape. They measure 0:‘075 X0‘055 mm., and their
contents appear to be unsegmented at the time of laying.

Falcaustra stewarti, sp. nov.
(Figs. 43, 44.)
Hosts and localities :
Kachuga smithii ; Ferozpore, Punjab.
Hardella thurgi ; Siripur, Saran, Bihar.

This is a species of moderate size, differing from the typical forms in having a
larger number of caudal papillae in the male, and in other features. In the material
from Kachuga smithii, which we take as
typical, the length of the male is 17-198 DIR
mm., that of the female 19-22°6 mm. =
The greatest thickness is 0°6-0°7 mm. in
the male; 0°65-0°75 mm. in the female.
The cuticular striation is exceedingly fine.
The head has a diameter of 0'Ig-0'21

mm., and is followed immediately by a S

A GEG

slightly narrower neck. The buccal x Za
: i

cavity is very shallow, measuring only
0°06 mm. in length. A distinct pharynx
is present, oo mm. long. The total

length of the oesophagus is 2°I-2°5 mm. en GE Ot Hoel
The bulb has no marked constriction, and ., cuticular ring; p., forked pulp of papillae;
measures 0°5-0'55 mm. in length and ph., pharynx.

0°3-0°32 mm. in diameter. The small,

but very prominent, cervical papillae are at 1°3-1°37 mm., the nerve-ring at 0°5-'0°6

308 Memoirs of the Indian Museum. [Vor. VII,

mm., and the excretory pore at 1-6-1°65 mm. (male), or 175-1'85 mm. (female), from
the anterior end.

The tail of the male (fig. 44) is 1°4-1°7 mm. long, and tapers to a slender. point.
There is no preanal sucker-like organ, but the oblique caudal muscles are well-deve-
loped. There are 16-18 pairs of caudal papillae and one median, unpaired papilla.
the latter and three pairs being, as usual, preanal. Of the postanal papillae two
pairs are lateral, the rest ventral. Oc-
casionally one or two of the anterior
ventral pairs become adanal in position.
The members of the more posterior pairs
sometimes become displaced anteriorly
or posteriorly, so as to disturb the sym-
metry of the paired arrangement. The
spicules are short (0°5 mm.) and have a
maximum width of 0:09 mm. A vaguely-
defined mass of imperfectly chitinized
tissue represents the accessory piece, and
a fan-shaped bundle of muscles extends
from it to the dorsal body- wall.

The tail of the femaleislong (2°25-2°6
mm.), straight and tapering, and ends in
a fine point. The caudal papillae are
situated at about 1°6 mm. from the tip.
The vulva opens at 7°75-10°3 mm. from

Fie. 44.—Falcaustra stewarti. Tail of male; ins Rosvenen end. ne Tag ne IS Son
ame slew, 1'5 mm. long. The ova measure about
0'I5X0'IOo5 mm., have a shell 5” thick,

and contain an embryo curled upon itself into a U-shape when ready for laying.

The material from Hardella thurgi so closely resembles that from Kachuga in
almost all respects that we do not feel justified in erecting another species for it.
There are, however, certain differences, of which the most conspicuous are the greater
relative and absolute width of the head and the much coarser striation of the cuticle.
The striae in the Hardella material are about 10: apart, whereas in the Kachuga
material they are so fine that accurate measurement is scarcely possible Apart
from these points the Hardella material is slightly larger in almost all dimensions
than that from Kachuga, and has a stouter general appearance. The measurements
may be most conveniently given in tabular form, for comparison with those of the
Kachuga material :—

[6% 2
mm. mm.
Length Se of Si Ai: .. 18:6-2C°4 20:2-22:1
Thickness (max.) oe 50 = .. 0°8-0°9 0:8-1:05
Diameter of head Do ss er .. 0°28-0°3 0:3-0°35

Length of tail So 8 Bb en 1:3-1:9 2:5-2:7

1922. ] H. A. Bayuıs AND R. DAUBNEY : Parasitic Nematodes. 309

af ®
mm. mm.
Distance from ant. end to end of oesoph. (incl. bulb) .. 2°6-2°8 2°7-2'8
Fe „ nerve-ring Ae 32 0.56 0:6
a ., excretory pore a ao 1:9 2:0
Length of pharynx > *: Le LUS 0:1
Oesophageal bulb, length or ah ee OD 0 Sm 2.056
ns „ greatest diameter a me 0:35 0:4
Spicules, length J Fe de .. 0°54-0°56 oe
Vulva. distance from posteriorend .. 2 = de 3:5-8°8
Caudal papillae, 2 , distance from tip of tail ES a cee 2:0
Ova, measurements aR Br Bis EEE 0:15 x 0:0875

Falcaustra kachugae, according to Stewart’s (1014) description, seems to differ
notably from F. stewarti in its much smaller dimensions; but as the type-material
consisted only ofa single female, which may not have been mature, the question of
identity must be left open.

In view of the severai new species just described, it is necessary to revise our
conception of the generic diagnosis of Falcaustra. Diagnoses have been attempted
by Seurat (1918) and by Baylis (1920 b), but both require some alteration. The
following is an attempt at a fresh generic characterization.

Falcaustra, Lane, 1015.

Ascaroidea: Kathlanidae:! Meromyarian. Body usually stout, tapering at each end. Lateral
fields wide. No lateral alae. Mouth with three lips, each bearing two outer and two inner papillae;
the pulp of each outer papilla sends a branch to one of the inner papillae, and is thus Y-shaped.
Buccal cavity short, surrounded by a continuous ring ? of thickened cuticle. Muscular oesophagus
divided into a short anterior portion, or pharynx, and a long posterior portion, the latter ending ina
well-marked bulb which is constricted in the middle so as to take the form of two more or less distinct
swellings connected by a narrower neck. The oesophagus, with the exception of the bulb, is usually
considerably coloured with a reddish-brown pigment, and there are generally special masses of this
pigment in the region of the nerve-ring. ‘Excretory pore towards the posterior end of the oesophagus.
Tail in both sexes tapering and pointed. Caudal end of male without alae, and provided with ten or
more pairs of papillae (of which three pairs are constantly preanal), and an unpaired. median preclo-
acal papilla. Of the postanal papillae two pairs are constantly lateral. Preanal caudal muscles well-
developed, sometimes aggregated into one or several fan-shaped groups to form sucker-like organs.
Spicules equal, sickle-shaped, broad dorso-ventrally and compressed laterally, each having the appear-
ance of a spicule within a spicule. An accessory piece usually present. sometimes imperfectly chitinized
or even absent. Vulva towards posterior third of body. Vagina runs forward and gives off two
opposed uteri, each of which doubles upon itself in a number of longitudinally-disposed U-shaped
loops in the anterior or posterior region of the body respectively. Each ovary forms a loop in the

1 Travassos (1918) established this family to include the genera Kathlania, Tonaudia. Falcaustra and Florencioia.
It is the family Pseudo-heterakidae, Travassos, 1917, renamed and reconstituted. No family diagnosis, however, seems
to have been attempted. In our opinion the genus Cruzia, Travassos, 1917, should also be included in the family, and
not referred to a separate family Cruzidae, as Travassos has proposed. We have also to add a further new genus,
closely allied to Falcaustra (see below, p. 310).

2 Seurat (1918) in his diagnosis of the genus, speaks of the buccal cavity being ‘‘ encadrée dans sa région moyenne
par trois plaques chitineuses.’’ If such a structure exists in F. lambdiensis, the form studied by him, it would appear
to approach our genus Zanclophorus (see below, p. 310), though the rest of its characters seem to be those of Falcaustra.

310 Memoirs of the Indian Museum. [Vor. VII,

anterior region of the body. that belonging to the anterior uterus being confined to this region, while
the posterior ovary eventually runs back to terminate in the hinder region. Ova usually large and
thick-shelled, of oval shape, and laid at different stages of development in different species.

Hab. ‘ntestine of Chelonia and freshwater fishes.

Genotype: F. falcata (v. Linst., 1906), from Geoemyda trijuga.

Owing to the great similarity in structure between most of the species of Falcau-
stra, specific determination depends very largely upon measurements. The following
admittedly artificial attempt at a key to the species is based almost entirely on
male characters, and therefore omits one species (F. kachugae) of which only the
female is known.

A. Inhabiting tortoises.

I. Several preanal sucker-like organs present in male.
a. Pairs of caudal papillae 10; spicules about 0-9 mm.
long 2% we siamensis.
b. Pairs of caudal papillae 11; spicules about 1°3 mm.
long ar ap ae .. lambdiensis.
IT. Preanal sucker-like organs absent.

a. Pairs of caudai papillae 10 a .. falcata.
b. Pairs of caudal papillae 11 Sis .. testudinis.
c. Pairs of caudal papillae 16-18 Me .. stewart.

B. Inhabiting fishes.
I. A preanal sucker-like organ present in male. Head wider

than neck 26 IR re a.) anor
II. Preanal sucker-like organ absent. Head narrower than
neck sip ee - .. leptocephala.

Genus Zanclophorus, nov.

The collection contains two interesting species which are clearly very closely
related to Falcaustra, but differ from it, in our opinion, sufficiently to necessitate the
formation of a new genus, which may be defined as follows :— .

Kathlanidae : closely resembling Falcaustra in general appearance. Head somewhat narrower than
neck, surrounded by a slight cuticular collar at the base. Three large, flattened lips. each carrying a
pair of rather prominent papillae, and bordered internally by cuticular fringes. A long and wide buccal
cavity present, with a cuticular lining. In place of the continuous cuticular ring which surrounds the
buccal cavity in Falcaustra, there are three separate cuticular supports, in the form of double horse-
shoes, at the corners of the mouth. There is no distinct pharynx, but the structure of the oesophagus
is otherwise the same as in Falcaustra, and it is coloured in the same way with reddish pigment, of which
there is a special mass in the neighbourhood of the nerve-ring. Bulb pear-shaped, with narrow middle
region. Cervical papillae small and sessile, some distance behind the nerve-ring. Excretory pore
towards hinder end of oesophagus. Caudal end of male without alae, but with a single, well-developed,
muscular, preanal sucker (not a mere fan-like arrangement of muscles). Spicules similar to those of
Falcaustra, but relatively much longer. A large, but not completely chitinized, accessory piece
present. Female genital organs as in Falcaustra.

Hab. Stomach and intestine of Chelonia.

Genotype: Z. annandalei, sp. nov., from Testudo travancorica.

1922.] H. A. BayLıs AND R. DauBNEY: Parasitic Nematodes. 311

Zanclophorus annandalei, sp. nov.
(Figs. 45-48.)
Host: Testudo travancorıca. Position: stomach. Locality: Cochin State

Forests, Western Ghats.

O*1 MM.

Fie. 45.—Zanclophorus annandalei. Head of female; dorsal view.
b., lining of buccal cavity; c., one of the three cuticular supports; p., papilla.

The male measures 15°5-15°9 mm. in length, the female 15'0-17'4 mm. The
maximum thickness is 0°85-1' mm. The cuticular striations are about 2 * apart.

The head (figs. 45, 46) has a diameter of
0°2-0'23 mm. The buccal cavity
measures 0'I4-0'I5 mm. in length and
0°09 mm. in greatest diameter. The
distance from the anterior end to the end
of the oesophagus, including the bulb, is
26-285 mm. The bulb measures 0°6-
0°65 mm. in length and 0°4-0'44 mm.
in diameter. The cervical papillae are at
I°5-1°74 mm., the nerve-ring at 0°55-0°6
mm., and the excretory pore at 2°2 mm.,
from the anterior end.

In the male, the tail (figs. 47, 48) is

0°45-0'5mm.long. The sucker is situated
at about 1°5 mm. in front of the cloaca.
The caudal papillae are arranged in ten
pairs and one median precloacal papilla.
Ofthese, four pairs are postanal (2 ventral
and 2 lateral), the rest preanal, consisting
of three pairs close together near the

Fie. 46. —Zanclophorus annandalei. Lips, viewed
en face.

c., one of the three cuticular supports; d.l.
dorsal lip; f., cuticular fringe of lip; p., papilla.

312 Memoirs of the Indian Museum. [Voz. VII,

cloaca, and three more pairs, more widely separated, between these and the sucker.
The spicules measure 2°2—2°3 mm. in length, and have a maximum width of 0°058 mm.
The large accessory piece (figs. 47, 48, a.p.), which is only partially chitinized, is
deeply cleft in front.

3
=

Fie. 47.—Zanclophorus annandalei. Posterior Fic. 48.—Zanclophorus annandalei. Tail of
end of male; lateral view. male; ventral view.
a.p., accessory piece: s., sucker. ap... accessory piece.

In the female, the tail measures 0°7-0°75 mm, in length. The caudal papillae
were not seen. The vulva opens at 5'0-5°5 mm. from the posterior end. The vagina
runs forward for about 2 mm. before giving off the two directly opposed uteri. Very
few of the females contained ova, and it may be doubted whether those seen were
quite fully-formed. They measured about 0'125 x 0'075 mm.

Zanclophorus kempi, sp. nov.

(Fig. 49.)

Host: Testudo elongata. Position: intestine. Localities: Baradighi, Jalpaiguri,
Bengal; and near Tura, foot of Garo Hills, Assam.

The length of the male is 10'9-12'8 mm., and its thickness 1‘0-1°1 mm. The
corresponding measurements for the female are 134-158 mm. and 1‘2-1‘4mm. The
cuticular striations are exceedingly fine. The diameter of the head is 0°22-0'24 mm.
The buccal cavity measures 0°13 mm. in length and 0'09-0'I mm. in greatest

1922.] H. A. BayLis AND R. DAUBNEY : Parasitic Nematodes. 313

diameter. The distance from the head end to the posterior end of the oesophageal
bulb is 2°3-2°5 mm. The bulb measures 0°45-0°6 mm. in length and 0'4-0'48 mm. in
diameter. The cervical papillae are at 1°5-1°6 mm., the nerve-ring at 0°52-0°58 mm.,
and the excretory pore at 1°7-1°95 mm., from the anterior end.

The tail of the male (fig. 49) is 0'45-0'55 mm. long. The sucker, which is deep
and strongly muscular, is situated at about I'I mm. in front of the cloaca. There
are nine pairs of caudal papillae and the usual median precloacal papilla. The
arrangement is the same as in Z. annandalei, except that one of the three pairs im-
mediately in front of the cloaca is absent. The two pairs present in this position
are small, and the three more anterior pairs much larger. The spicules are relatively
large, measuring 2°9 mm. in length and o'r mm. in width. The accessory piece is
similar to that of Z. annandale:.

Fie. 49.—Zanclophorus kempr. Posterior end of male; lateral view.
a.p., accessory piece; s., sucker.

The tail of the female is bluntly conical, and measures 0°55-0'8 mm. in length.
There is a pair of caudal papillae at 0°27 mm. from the tip. The vulva is situated
at 4’0-4°8 mm. from the posterior end. The vagina is simple and narrow, about
2 mm. long, and runs forward as usual from the vulva. The eggs are oblong-oval in
shape and measure 0°125-0°137 mm. X 0'075-0'085 mm. The content is unsegmented.

It is always interesting to discover fresh cases of close relationship betwen the
parasites of closely-related hosts, as showing how parallel evolution may be taking
place in both hosts and parasites simultaneously. The present case is a very good
example, neither the hosts nor their parasites having as yet developed sufficiently
divergent characters to obscure their extremely close relationships.

Dr. Annandale has been good enough to supply us with the following interesting
note on the hosts of our new genus :—

314 Memoirs of the Indian Museum. [Vor. VII,

‘* Testudo elongata and 7’. travancorica belong to a small group of species which are very closely allied
but remarkably isolated geographically. This group consists of four species: 7. travancorica, from
the southern part of the Malabar Zone of Peninsular India; 7. parallelus, of which only a single specimen
is known, from Chota Nagpur, in the middle of Peninsular India ; 7’. elongata, with the widest range
in the group, extending from Jalpaiguri in the extreme north-east of the plains of Bengal, through
Assam, Burma and the northern part of the Malay Peninsula to Siam and Cambodia ; and 7’. forstenii, from
the island of Celebes. Discontinuous as the range of the group appears to be, the close structural simi-
larity and the remarkable resemblance in facies indicate that the range was once continuous. Moreover,
the existence of a single specimen from the interior of Peninsular India, captured many years ago, sup-
ports this view and suggests that rare annectant forms may linger on as yet undiscovered in inacces-
sible districts, perhaps of very limited area.”

Superfamily FILARIOIDEA, Weinland, 1858.
Family FILARIIDAE, Claus, 1885.
Subfamily FILARIINAE, Stiles, 1907.
Genus “ Filaria ”, sens. lat.

Filaria haje, Wedl, 1862 (?).

Young Filariid worms, perhaps belonging to this species, occurred twice in
the intestine of the cobra (Naja tripudians) and twice in that of the banded krait
(Bungarus fasciatus). All of these are immature forms, about 6-8 mm. in length, and
without characters which would enable them to be assigned definitely to any well-
established genus. They have a relatively long posterior glandular portion of the
oesophagus. Wedl’s (1862) description is very brief, and does not enable his species
to be recognized with certainty. His specimens were found either free or encapsuled
in the thoracic cavity, outside the lung. It is not indicated whether the present
material occurred free in the lumen of the intestine or not.

Filaria abbreviata, Rud., 1819 (?).
(Fig. 50.)

The collection contains two female specimens of a Filariid from the orbit of
Saxicola, sp. They are possibly referable to the above-named species, of which no
full description appears to exist. Our material appears to agree fairly well with the
account of F. abbreviata given by Molin (1858), and for this reason we tentatively
refer it to this species.

The larger specimen is about 24 mm. long and 0°57 mm. in thickness. The
anterior end of the body is abruptly attenuated and sharply truncated. The cuticle
is smooth, and we are unable to detect the longitudinal rows of deciduous spines to
which Molin refers. The mouth opens into a small buccal cavity which is about
0'032 mm. deep and 0°02 mm. in diameter. We are unable to distinguish any teeth
at the base of the buccal cavity, but its wall is thrown into folds presenting an ap-
pearance which might easily have been mistaken for teeth. The oesophagus consists
of two parts. The anterior portion is 0°25 mm. in length and distinctly more slender
than the posterior portion. The latter measures 0°75 mm. in length and about o-12

1922.] H. A. Bayrıs AND R. DAUBNEY : Parasitic Nematodes. 315

mm. in thickness. It is very slightly enlarged posteriorly. The nerve-ring surrounds
the oesophagus at about o'I3 mm. from the anterior end, while the excretory pore
opens at about 0'2 mm. from the head. The
posterior extremity is rounded and not notice-
ably attenuated. The anus is small and
subterminal. The vulva is situated close to
the head, at about 0°45 mm. from the anterior
end. There is a short transverse vagina,
directed slightly backwards, from which two
ovejectors are given off. The uteriand ovaries
are both posterior, but a forwardly-directed
loop in the ovejector of one of them indicates
that this represents an anterior uterus. The
eggs in the uterus measure about 0°024 X 0'017
mm., and are in various stages of segmentation.

Henry and O’Zoux (1909) include F. abbreviata in
a list of species of the subgenus Diploiriaena. Walter
(1866) gives a description and figure of a worm from
Motacilla alba, which he regarded as F. abbreviata, but
these make it quite clear that his materia] belonged
to a species of Diplotriaena. He also describes and
figures as a Filariid under the name of F. (attenuata 2),
from Corvus corone, Garrulus glandarius, :Saxicola
rubicola and Falco tinnunculus, a form which is clearly a
Diplotriaena ; aud it seems not improbable that the
material in both of these instances was referable to D.
tricuspis, which is recorued from a very similar range sail of females lai ew

f hosts. It appears to us questi à
HE PESTE ml puonable Wits ne us b., buccal cavity; e., excretory pore; nr.
original F. abbreviata was in reality a Diplotriaena. nerve-ring ; v., vulva.

Fic. 50.—Filaria abbreviata (?). Anterior

“Filaria,” sp.

A single specimen of a different species from those mentioned under F. haje
occurred in the intestine of a cobra. It is an immature form measuring about 20
mm. in length. The anterior end is broad and blunt, the posterior end more taper-
ing. The oesophagus is relatively shorter than in “ F. haje.”’

Filaria macrophallos, Parona, 1880.
(Fig. 51.)

Hosts: Varanus salvator, V. flavescens, V. nebulosus, ‘ Bengal monitor”,
Varanus, sp. Position: from the labels it is not quite clear whether the habitat is
the actual cavities of the lungs, or the thoracic cavity. In three cases the lungs are
mentioned, in one no position is specified, and in two the “intestines”’ are mentioned.

Great difficulty was evidently experienced in collecting whole specimens of this
worm. We have only succeeded in finding one whole female among the material at
our disposal, the rest consisting of much-tangled fragments,

316 Memovrs of the Indian Museum. [Vor. VII,

The complete specimen came from an unnamed species of Varanus. It measures

about 250 mm. in length and I’4 mm. in thickness.

like depression.
prominent, chitinoid teeth, projecting forward,
one on either side of themouth. A little further
back on each side there are three small, sessile,
cephalic papillae.
short, narrow, anterior, muscular portion and a
very long and wide, posterior, glandular portion.
The length of the former is 0°6 mm.; of the
latter, 30 mm.
glandular portion occupies the whole width of
the body-cavity.
anterior portion somewhat behind its middle.

The cuticle is marked through-
out with prominent, raised, transverse wrinkles
at irregular intervals, suggesting spines when
seen in optical section. The head, which has
a diameter about 0°45 mm., is squarish in front,
the mouth often lying at the base of a funnel-

There are two small, but

The oesophagus has a very

At its commencement the

The nerve-ring surrounds the

Fre. 52.—Filarta varani. Tailot male; Neither cervical papillae nor an excretory pore

ventral view.

à were seen.
s., Spicule.

The anus is almost terminal. The tail-end is bluntiy rounded, and carries a pair
of papillae at its extremity. The vulva opens at 1'I5 mm. from the anterior end.
The muscular vagina may run straight back, or may be much convoluted. The ova
(fig. 51) are of very characteristic shape, somewhat resembling a barrel, with an annu-
lar thickening near each pole. They measure 0°05 X 0°03 mm., have thick shells, and

contain coiled embryos when ready for laying.

This species, according to Henry and O’Zoux (1909), should be referred to the genus Diplotriaena,

but with this view we are unable to agree.

Filaria varani, sp. nov.

(Pig. 52.)

A single male individual from Varanus flavescens does not agree with the male
of F. macrophallos, and must be regarded provisionally as belonging to a new species.

It measures about 108 mm. in length and 0°7 mm. in
thickness. The characters of the anterior end are
similar to those of F. macrophallos. The diameter of
the head is 0°23 mm. The anterior portion of the
oesophagus is 0°35 mm. in length, and the posterior

portion 16 mm.

The tail (fig. 52) is 0'I6 mm. long, and there are
well-developed caudal alae, which are continuous round

de®

Fie. 51.—Filaria macrophallos.
Ova.

1922. ] H. A. BayLıs AND R. DAUBNEY: Parasitic Nematodes. 317

the posterior extremity. Only one spicule' appears to be present, and this, from its
position, seems to be that of the right side. It is a very broad structure, of charac-
teristic shape (fig. 52, s.), and measures 0°6 mm. in length and 0:07 in width. There
are seven pairs of caudal papillae, of which four are preanal. These and the most
anterior postanal pair have long peduncles. Their arrangement is best indicated by
means of the figure. The posterior lip of the cloacal aperture is tumid.

Genus Setaria, Viborg, 1795.
Setaria, sp. (?).
Three larvae, taken from “ inner surface of cartilage’’ of a Javan mouse deer, or
chevrotain (probably Tragulus javanicus).
The worms are of a Filariid type, but are too immature to show recognizable
generic characters.

Subfamily DIPLOTRIAENINAE, Skrjabin, 1916.
Genus Diplotriaena, Railliet and Henry, in Henry and O’Zoux, 1900.
Diplotriaena tricuspis (Fedchenko, 1874).

The collection includes one male specimen belonging to this species, from Blan-
ford’s laughing-thrush (Trochalopterum meridionale).

Subfamily MICROPLEURINAE, nov.
Genus Micropleura, v. Linst., 1906.
Micropleura vivipara, v. Linst., 1906.

(Figs. 53, 54.)

Host: Gharial (Gavialis gangeticus). Position: Liver.

For the purpose of confirming the determination of this material, we were fortu-
nate in being able to obtain from the Indian Museum five examples from the type
series. These, as well as the new material, unfortunately proved to be all females,
but we are in a position to add a few details to the description furnished by von
Linstow (I906 a).

The dimensions of the female may be slightly larger than those given by that
author, reaching about 43 mm. in length and 1 mm. in thickness. The cuticle, al-
though without striations, is not perfectiy smooth, as stated by von Linstow. There
are distributed rather irregularly about its surface, especially on the hinder portion
of the worm, little longitudinal series of from 2 to 7 very minute, raised, papilla-
like structures. There is an appearance of a remarkable structure lining the body-
cavity—..e. within the musculature. This “structure”
refringent network, strongly suggestive of a series of longitudinal tubules connected
by smaller tubules running transversely. It seems probable, however, that this ap-
pearance is an artifact—it may perhaps be the result of the fluid contained in the

takes the form of a highly

1 In F. macrophallos there are two unequal spicules. Our specimen does not appear to be in any way damaged.

318 Memoirs of the Indian Museum. [Vor. VII,

body-cavity having coagulated on fixation and formed a membrane which is thrown
into folds in the peculiar pattern indicated.

The mouth is, as v. Linstow states, without lips. There are six small cephalic
papillae, two lateral and four sublateral. The lateral papillae are slightly larger than
the others. The diameter of the head at
the level of the papillae is 0°15 mm. The
oesophagus is distinctly divided into an
anterior, narrow, muscular portion and a
much longer and wider, posterior, glandular
portion. The muscular portion measures
only 0°75-0°9 mm. in length, while the total
length of the oesophagus is 3°25-3°5 mm.
The muscular portion itself is slightly gra-
nular in appearance for nearly the posterior
half of its length. It is surrounded by the
nerve-ring at 0'67-08 mm. from the
anterior end.

Contrary to the statement of v. Linstow,
a minute excretory pore is present on the
ventral surface a little behind the junction
f the two portions of the oesophagus (at
I'0O-I'I mm. from the anterior end), and
connected with this there is a structure
apparently representing the excretory
“bridge,” into which faint indications of
ducts can be seen coming from both anterior
Fig. 53.—Micropleura vivipara. Anterior end and posterior directions. These ducts,
ne Reste however, have not been traced along the
e., excretory pore; n.r., nerve ring oes. 1?., :
oes.1®., two portions of anterior division of oeso- lateral fields, nor has any connection been
phagus ; oes.2., posterior division of oesophagus; observed between them and the peculiar
ov., oviduct.
“network ” already referred to.

The vulva is very hard to see in mature females, even when perfectly cleared,
owing to the dense mass of embryos contained in the uterus. It is situated slightly
in front of the middle of the body. The vagina consists of a very narrow, non-mus-
cular duct running through the body-wall in a postero-dorsal direction from the open-
ing, and a very short, somewhat muscular portion returning towards the head and con-
nected with the uterus. The whole of the vagina is not more than 0-4 mm. in length.
The two branches of the uterus are directly opposed, and form one continuous straight
tube joining the ovaries, which are situated at opposite ends of the body-cavity. This
tube fills the whole width of the body-cavity with the exception of the space occupied
by the very narrow intestine, which runs in close contact with the body-wall. The
ovaries are exceedingly short in proportion to the length of the worm. They are
usually reflexed, but occasionally continued in a straight line with the uterus. They

uU G-0

1922.] H. A. Bayzis AND R. DAUBNEY : Parasitic Nematodes. 319

are connected by narrow ducts with the respective ends of the latter. The develop-
ment of the embryos appears to be very rapid, the uterus being entirely filled, from
end to end, with young apparently fully-formed
and not enclosed in membranes. These em-
bryos have, as v. Linstow observes, a cuticle
marked with conspicuous transverse striations,
a blunt head and a long, tapering tail.

The tail of the adult femaie (fig. 54) is
rounded at the tip, and measures 0'2-0'35 mm.
in length. At o'I-o'I4 mm. from the tip, and
somewhat towards the ventral side, there is a
pair of large, prominent caudal papillae. =,

This genus, with the uteri directly opposed, Tas Be à
the vulva placed far back from the head, the nn a en
short ovaries, and the spicules of the male (ac- p., caudal papilla.
cording to v. Linstow) of equal length, does
not appear to fit very well into any existing subfamily of Filariidae. It seems
justifiable, therefore, to regard it as the type of a new subfamily, Micropleurinae.

A
=

Superfamily SPIRUROIDEA, Railliet and Henry, 1075.
Family SPIRURIDAE, Örley, 1885.
Subfamily ACUARIINAE, Railliet, Henry and Sisoff, 1912.
Genus Acuaria, Bremser, 1811.
Acuaria (Acuaria) anthuris (Rud., 1819).

One female, which we assign to this species, was collected from the Red-billed
Blue magpie (Urocissa occipitalis).

Acuaria (Echinuria) leptoptili (Gedoelst, 1916).
(Figs. 55, 50.) |

This form was collected from the Adjutant (Leptoptilus dubius), at Calcutta.
The species was described by Gedoelst (1916) trom females only. The type host was
Leptoptilus crumenifer.

The females in the present collection measure from 13 to 15 mm. in length, and
about 0°36 mm. in thickness; the males 11 to 11°5 mm. and 0'234 mm. respectively.
The cuticle has fine transverse striations about 4 „apart. At the anterior end it is
ornamented with “cordons.” These cordons actually consist of a continuous band
folded so that two folds lie dorsally and two ventrally. The transverse portions of
the band are arranged so that they run across the Jateral lines posteriorly and across
the dorsal and ventral surfaces anteriorly, The cordons are 0:02 mm. broad and they
extend backwards to a point about 1°:1 mm. from the anterior end. The posterior
transverse portions show a slight forward bend as they cross the lateral lines. The
disposition of the longitudinal portions is not markedly asymmetrical or inclined to-

320 Memoirs of the Indian Museum. Vor. VII,

wards the ventral surface, as described by Seurat (1919), for Echinuria, Soloviev,

1012, and there are no cuticular spines. The cervical papillae are prominent and

slender and are situated a little behind the lateral bends

of the cordons.

oe Hae The head is small and provided with two simple lips,
TBH € and is not constricted off from the body. The oesophagus

consists of the usual three portions, a pharynx, a muscular

portion, and a glandular portion.

ar These measure 0°31,
0'4I, and 2°3 mm. respectively. The nerve-ring surrounds
the oesophagus at about 0:27 mm. from the anterior

extremity, while the excretory pore opens at about 0°54
De mm. from the head. The vulva opens at about 0:17 mm.

RU pop De

Sy: A from the tip of the tail. Its anterior lip is modified into a
Ny] IB À 6 a ae 2 .
| : 2 prominent bulia. From the vulva there is a long, straight,
S 2 muscular-walled vagina which runs forward for about I’4
À mm. and then turns transversely to join the uterus. The
Ä species may be considered to be monodelphiec, the posterior
IE e . Ö
ES set of organs being represented merely by a blind sac-like
WY: ee) . re :
Ry N) uterus which runs back to the vicinity of the vulva. The
A: ‘
IN: . . .
ke | anterior uterus runs forward to within 1°75 mm. of the an-
06538 ss terior end of the body, the ovary commeneing at this point
SS LE 2 5 2 à
AN Ba and running backwards. In a mature specimen practically
A. AA
ES ee

Bike A the whole of the body-cavity is occupied by the uterus,
Fig. 55.—Acuaria (Echinu- which is packed with eggs containing fully-developed em-

ria) leptoptili. Anterior end N Fant lhe ;

6 females nel bryos. The ova measure 0°028 X 0'0I8 mm.

c., cordon; e., excretory Situated at 0:05 mm. from the tip of the tail.
pore; n.r., nerve-ring oes. 1.,

nee The tail of the male forms several turns of a spiral.

oes.2., 0es.3., three divisions of ;

oesophagus. Membranous alae are present, extending for about 0°7 mm.
trom the tip of the tail.

The anus is

The cloaca opens at 0'II5 mm.

There are nine pairs of supporting papillae in the
alae; 4 preanal pairs, about equidistant from each other and close to the cloaca, and

5 postanal. The postanal papillae are arranged as follows: four pairs in a group,
occupying the posterior half of the tail, and one pair about midway between this
group and the anus. The papillae are all stalked and slender, the preanal ones being
much longer than the postanal. The spicules are unequal and dissimilar. The right
spicule is long and slender. It is gently curved, and if partly extruded the distal
portion may be generously incurved but it is not twisted. It measures 0°65—0°675 mm.
The left spicule (fig. 56), is short and very much curved. It lies roughly at right
angles to the long axis of the right spicule, measures 0°19-0°21 mm. in length, and
is twisted and flanged. The root of the spicule is expanded. Its edges are faintly

serrated in the part towards the tip. The testicular coil runs forward to within 3°5
mm. from the head.

from the tip of the tail.

A genus Echinuria (genotype H. jugadornata) was erected by Soloviev in 1912, the month of publi-

1922. | H. A. Bayuis AND R. DAUBNEY : Parasitic Nematodes. 321

cation being September. In December of the same year Railliet, Henry and Sisoff published a note on
the relationships of the members of the genus Acuaria, Bremser, 1811, and erected a new sub-family
Acuariinae containing the genera Acuaria (type), Bremser, 1811,
Cosmocephalus, Mol., 1858, Histiocephalus, Dies., 1851, and Strep-
tocara, Raill., Hen. and Sis., 1912. The members of the genus
Acuaria were separated as far as possible into five sub-genera, vız.,
Acuaria, Cheilospirura, Diesing, 1860, Dispharynx, Raill., Hen. and
Sis., 1912, Synhimantus, Raill., Hen. and Sis., 1912, and Hamannia,
Raill. Hen. and Sis., 1912.

The sub-genus Hamannia is recognised by Seurat (1919) as a
synonym of Echinuria, Soloviev, 1912. While he retains full generic
rank for the latter, for his new genus of 1918, Chevreuxia, and for
Rusguniella, a genus erected in the same paper, he assigns species
to the following sub-genera of Acuaria: Acuaria, Dispharynx and

o1mm.

Synhimantus.
The emended diagnosis of the genus Acuaria, Bremser, 1811,
given by Railliet, Henry and Sisoff, runs as follows: Acuariinae
Fig. 56.—Acuaria (Echinuria)
leptoptili. Left spicule; lateral
view.

without vesicular swelling at the anterior end, but bearing four
cutaneous cordons in the form of gutters or bands salient from
or countersunk in the cuticle, these cordons extending sometimes
directly backwards, more often returning forwards or even united two by two across the lateral surfaces.
Parasites of the oesophagus, ventriculus or gizzard. Type, Acuaria anthuris (Rud., 1819).

Accepting this definition, then, we can see no reason why Echinuria, Soloviev, 1912, Rusguniella,
Seurat, 1919, or Seuratia, Skrjabin, 1916, should be given full generic rank.

_ The suggested arrangement, then, is as follows: Acuarnnar, Raill, Hen. and Sis., 1912:
Genotype, Acuaria, Bremser, 1811. Other genera: Cosmocephalus, Mol., 1858 ; Histiocephalus, Dies..
1851 ; Streptocara, Raill., Hen. and Sis., 1912; Chevreuxia, Seurat, 1918.

Genus Acuaria, Bremser, 1811. Sub-genera : Acuaria, Bremser, 1811, emend. Raill., Hen. and Sis.,
1912; Cheilospirura, Dies., 1860, emend. Raill., Hen. and Sis., 1912; Dispharynx, Raill., Hen. and Sis.,
1912; Synhimantus, Raill., Hen. and Sis., 1912; Echinuria, Soloviev, 1912; Rusguniella, Seurat, 1919;
Seuratia, Skrjabin, 1916.

Sub-genus Hehinuria, Soloviev, 1912 (Synonym Hamannia, Raill., Hen. and Sis., 1912). Diagnosis
after Raill. Hen., and Sis. (1912) :— Acvaria with cuticular cordons non-recurrent but anastomosed in
pairs across the lateral lines. Body sometimes spiny. Males with spicules unequal and unlike. Four
or five pairs of postanal papillae.

Habitat : ventriculus or gizzard.

Type, Acuaria (Echinuria) jugadornata, Soloviev, 1912.

Other species: A. uncinata (Rud., 1819); A. contorta (Molin, 1858); A. longeornata (Mol., 1860);
A. calcarata (Mol., 1860); A. spinifera (Schneider, 1866); A. squamata (v. Linst., 1883) ; A. phoeni-
copteri (Seurat, 1916), A. leploptili, Gedoelst, 1916.

Subfamily PHYSALOPTERINAE, Stossich, 1898 (fide Stiles and Hassall).
Genus Physaloptera, Rud., 1810.
Physaloptera alata, Rud., 1819.

One immature female specimen was collected from Montagu’s harrier (Circus

cineraceus).

322 Memoirs of the Indian Museum. [Vor. VII,

Physaloptera praeputialis, v. Linst., 1889.
Syn. Chlamydonema felineum, Hegt, 1910.

Specimens of this curious form occurred in the following hosts :—

Fishing cat (Felis viverrina).
Jungle cat (Felis chaus).
Leopard cat (Felis bengalensis).
Leopard (felis pardus).

We are of the opinion that the form described by Hegt (1910) and discussed by
Nierstrasz (1910) is identical with von Linstow’s species, and that there is no
justification for the erection of a separate genus Chlamydonema. The species has all
the characters of a true Physaloptera, though complicated or obscured to a varying
extent by the development of a cuticular caudal sheath,

Physaloptera, sp.

Host: Indian fox (Vulpes bengalensis). Position: stomach.

Apparently the only species of Physaloptera recorded in foxes is P. cesticillata,
Sonsino, 1889, from the Fennec fox (Megalotis zerda) of Northern Africa. The
present material consists only of a single female specimen, and, in the absence of a
male, it is impossible to identify it with Sonsino’s species. In the characters of the
lips, and in some other respects, our specimen appears to approach very closely to
P. digitata, Schneider, 1866. This species, however, has only been recorded from the
puma (Felis concolor) in Brazil, and without examining further material it would be
unwise to suggest the identity of this with a form from the Indian fox.

Physaloptera, sp.
A number of immature forms, measuring about 5-7 mm. in length, were found
in the intestine of a “Raket Bausi”’ (Coluber, sp.). In the apparent absence of the
inner teeth of the lips they resemble P. colubri (Rud., 1819), according to von Drasche’s

(1883) account of that form ; but since there are no sexually mature specimens the
determination remains uncertain.

Family CAMALLANIDAE, Railliet and Henry, 1915.
Genus Camallanus, Railliet and Henry, 1915.
Camallanus kachugae, sp. nov.
(Figs. 57-59.)

Host: Kachuga smithii. Locality: Ferozpore, Punjab.

This species has the typical characters of the genus, which appear to vary little
except in small details in different species. The worms, as indicated by the collector’s
label, had the usual reddish colour during life. The total length is 10°9-14°5 mm. in
the male and 20°$—22'0 mm. in the female. The male has a maximum thickness of
0°3-0°37 mm., the female of 0-45-0'5 mm. The cuticular striations are fine, the
interval not exceeding 5 ;. The dorso-ventral diameter of the head, measured at the

1922.] H. A. BayLıs AND R. DAUBNEY : Parasitic Nematodes. 323

anterior angles, is 0°I3-0'I5 mm. in the male, 0°17 mm. in the female. The chitinous
buccal valves are slightly broader than long, their length (not including the posterior
ring) being 0°II—o'I3 mm. and their width

about 0'14-0'I6 mm. The number of D
longitudinal ridges on each valveis either
eight or ten, the latter number being seen
only in large specimens. The posterior
ring of the buccal apparatus has a dia-
meter of o‘:mm. The dorsal and ventral
““tridents ” are well developed, and the
middle prong measures 0'08-0°I mm. in
length. The head bears three papillae
on each side near the extremity. The
oesophagus shows the usual division into
an anterior, clear, muscular portion and
a posterior, more opaque, glandular por-
tion. The former is distinctly club-
shaped, and (measuring from the ex-
tremity of the head) 054-066 mm. in
length. The total length of the two
portions (from the head-end) is 1°18-1°55 Fie. 57.—Camallanus kachugae. Head of female ;
mm. The minute, bristle-like cervical lateral view.

papillae are situated at 0°5-0°55 mm., the a Be n.r., nerve-ring; p.,p., papil-
nerve-ring at 0'2-0'23 mm., and the CR

excretory pore at about 0'5 mm., from the anterior end. The intestine is very narrow,
considerably more so than the oesophagus.

In the male the tail measures about 0°21 mm. in length. The alar region is
somewhat thicker than the preceding portion of the body. The ventral region
between the alae is probably capable of being depressed by the action of the well-
developed caudal] muscles, so as to produce the effect of a sucker. The number and
general arrangement of the caudal papillae are the same as those described for certain
other species (C. microcephalus, C. americanus). Unfortunately, although a number of
species of Camallanus have been described, the majority of the descriptions are very
incomplete, and it is impossible to determine whether the number of papillae is constant
throughout the genus. It seems probable, however, that the seven pairs of rib-like
preanal papillae, projecting into the alae, will be found to be constant. The present
species has, in addition to these (fig. 58), six pairs of postanal papillae and two small
pairs of adanal papillae, as in the two other species cited. Their arrangement agrees
more closely with that of C. microcephalus than that of C. americanus, in that the first,
or most posterior, pair occupies an isolated position close to the tip of the tail. The
second pair is also isolated. Pairs 3 to 5 form a group, close together. All these are
lateral in position, while the sixth pair is more ventrally situated, just behind the cloaca.
The two small adanal pairs are not indicated in the figure. The two spicules are very

o-1 mm.

324 Memoirs of the Indian Museum. [Vor. VII,

unequal. The right spicule is fairly stout and measures about 0°97 mm. in length.
It has no barb or prong, such as is found near the tip in several species, but the tip

er N

cf!

seen,
IE
nu

NaN
QE
ONE
poison N
N

7%
v MIT.
(ii
Ra
à

Fig. 58.—Camallanus kachugae. Posterior end

of male; lateral view.
i., intestine; /., left spicule; r.,
s.d., sperm-duct.

right spicule ;

appears to be simple and finely pointed.

The left spicule is very slender and delicate,

and measures only about 0°43 mm. in

length.

In the female the tail is about 0°3 mm.

long, finger-shaped, rather blunt, and

slightly bifid at the tip. There is a pair of
caudal papillae at 0°17 mm. from the tip,
situated in little dimples in the cuticle:

The vulva is in front of the middle of the
body, at 9 1-10°4 mm. from the anterior
end. It has exceedingly prominent lips, the
anterior lip being considerably larger than
the posterior, and overlapping it. Each
of the lips (fig. 59) consists of a cuticular
swelling with granular contents, and has
the appearance of being divided internally
into a number of compartments by parti-
tions originating from the cuticle. The
muscular vagina is very narrow, running
back almost in a straight line for about 2
mm. At its inner end it gradually widens
into the short unpaired portion of the
uterus, which continues to run posteriorly.
The two uterine branches are directly op-
posed. The posterior branch, as usual,
has no ovary, but ends blindly at a short
distance in front of the anus. This blind

branch becomes filled with embryos derived from the other branch, in mature females.

The worm is, as usual, viviparous.

This form is extremely closely related to the European C. microcephalus (Duj., 1845). On comparison,
however, with specimens which we believe to belong to that species, we find that, apart from size,
which is too variable to be of much importance, C. kachugae differs from them in the following points :—

(1) The ‘“tridents” of the buccal apparatus are of slightly different shape, their outer prongs
being longer and more flattened and expanded at the free end in C. microcephalus than in C. kachugae.
The colour of the whole buccal apparatus, and especially of the ring and tridents, is much darker in

C. microcephalus.

(2) The longer (right) spicule of the male is simple in ©. kachugae, pronged in C. microcephalus.
In the former it measures 0:97 mm. in length, in the latter 0°85 mm. f

(3) The lips of the vulva in ©. kachugae are much more strongly developed than in €. microcephalus.
In the latter the posterior lip is often completely hidden by the anterior lip.

As regards other species already described from tortoises, C. americanus, Magath, 1919, has a barbed

1922. | H. A. Bavyris AND R. DAUBNEY : Parasitic Nematodes. 325

or pronged right spicule in the male. C. dumerilis (Perrier, 1871), as also the C. dumerilii, Perrier of v.
Linstow, 1897 and 1909, and consequently the C. confusus of Railliet and Henry (1915), are probably
all synonymous with ©. microcephalus. CO. tris-
pinosus (Leidy, 1851) and C. undulatus, Railliet

me he ee ee ee ee

and Henry, 1915 (= Cucullanus viviparus, v.
Linst., 1906, renamed), are so briefly described
that it is doubtful whether the species can be
identified. C. roseus (Leidy, 1851) probably be-
longs to another genus.

Magath (1919) is of the opinion that the C_
microcephalus of Seurat (1915), from Clemmys
leprosa, is a different species from C. microce-
phalus (Duj.), and proposes for it a new name, ©.
seurati. He considers it questionable whether

UUUL $°0

any form can be identified from Dujardin’s
description, and yet appears to find in the two
descriptions sufficient grounds for concluding
that the species are distinct. Such discrepancies
as there are in these descriptions, however, are
in matters of measurement, and it is clear that :
RN female; lateral view.

these are subject to great variation. We feel, 2 : : é

2 : À : The arrow points in the direction of the head.
therefore, that Magath’s view is somewhat ntestine and uterus omitted.
premature, and regard Seurat’s determination as
correct, unless a detailed comparison with material from Æmys orbicularis should prove the contrary.

Fria. 59.—Camallanus kachugae. Vulvar region of

Genus Camallanides, nov.
Camallanides prashadi, sp. nov.
(Figs. 60-63.)

Host: Banded krait (Bungarus fasciatus).

The buccal apparatus in this species differs somewhat from that typical for the
genus Camallanus. The paired valves, on superficial examination, appear to be
represented by four separate masses of brown chitin. Actually, these are joined
together in lateral pairs by relatively thin plates on their inner surfaces, so that they
are really pronounced external thickenings of the usual buccal valves. The usual
rib-like structures, terminating in tooth-like projections anteriorly, are present on
each inner plate to the number of about 14. The ‘‘tridents” are represented by
simple chitinoid rods of rather irregular shape and of a yellow colour. These are
connected at their bases with a dorsal and a ventral chitinoid body, lying opposite to
the edges of the buccal valves.

The male measures 5°8-6°6 mm. in length and 0'21-0'25 mm. in thickness. The
female is more than twice as large (14°2-17'7 mm. long and 0‘4-0°47 mm. in maximum
thickness). The cuticular striations are very fine and indistinct, at intervals of about
3-4. The diameter of the head, measured dorso-ventrally at the anterior corners,
is 0°08-0'09 mm. in the male, 0°12—0'13 mm. in the female. There appear to be three
pairs of cephalic papillae. The measurements of the buccal valves are: length,

326 Memoirs of the Indian Museum.

[Vor. VII,

0°06 mm. in male, 0'09-0'I mm. in female; width (dorso-ventral), 0°075 mm. in male,
o'r mm. in female. Thelength of the rods representing the tridents is about 0°06 mm.

Fra. 60.—Camallanides prashadi. Head of female ;

dorsal view.
b., buccal valve; p., papilla; ¢., “‘ trident.”

in the male, 0°07-0°I mm. in the female.
The posterior ring of the buccal appara-
tus has a diameter of 0'033 mm. in the
male and 0:045 mm. in the female. The
oesophagus has the same structure as
in Camallanus, the anterior muscular
portion measuring (from the extremity
of the head) 0:38 mm. in length in the
male, 0°47-0°5 mm. in the female. The
distance from the head-end to the poste-
rior end of the glandular portion is 0°8
mm. in the male, 1’02-1'08 mm. in the
female. The nerve-ring is situated at
0°'I5 mm. in the male, 0'IQ mm. in the
female, from the anterior end; the
excretory pore at 0°25 mm. (male), 0°31
mm. (female); and the minute, bristle-
like, cervical papillae at 0-28 mm. (male),
0°35 mm. (female).

In the male, the caudal region has

the general structure characteristic of Camallanus, with well-developed alae and caudal
muscles. The tail is very short (a little over 0:06 mm.), sharply pointed, and usually

curled ventrally, with its tip hidden by
the alae. The caudal papillae are
elongate and rib-like, diminishing in
size towards the tip of the tail. As in
Camallanus, there are seven pairs of
preanal papillae, projecting into the
alae. Thereare two large pairs curving
inwards towards the ventral surface at
the sides of the cloaca, and five more
pairs of lateral postanal papillae. The
three anterior pairs of these are rela-
tively large and close together. The
spicules are markedly unequal in size
and character. The right spicule is
stout and provided with alae for the
greater part of its length. It is about
0°24 mm. long. Its tip is curled into
a hook, but there is no barb. The left
spicule is without alae, slender and

Fig. 61 —Camallanides prashadi. Head of female;
lateral view.

Lettering as in fig. 60.

1922.] H. A. Bayris AND R. DAUBNEY : Parasitic Nematodes. 327

tapering, and measures only about 0'I4 mm. in length. There is a yellow chitinoid
accessory piece, more or less triangular in shape and measuring about 0'025 mm. in

length.

In the female the tail is relatively
long (0°4-0°6 mm.), and gradually taper-
ing, with the exception of a slight thicken-
ing just before the conical tip. No caudal
papillae were seen. The general arrange-
ment of the genital organs is that seen in
Camallanus, but the prominent lips of the
vulva are here modified into a tubular
appendage (fig. 63,) somewhat flattened
dorso-ventrally and projecting freely
from the body-wall in a posterior direction
to a distance of o'3-0'4 mm. This ap-
pendage originates at a point a little
behind the anterior third of the length

of the worm (at 5'°9-6'I mm. from the.

head-end). The vulvar aperture is situat-
ed on the ventral surface of this struc-
ture, near its extremity. The muscular
vagina runs from the opening to the base
of the appendage, and then turns back

just within the ventral body-wall, running

straight back for about 2 mm. before opening into the uterus.

Fic. 62.—Camallanides prashadi. Posterior end

of male; lateral view.

a.p., accessory piece; L., left spicule; r., right

spicule.

The posterior branch

of the uterus is, as in Camallanus, without an ovary. The worm is viviparous, both

AUUL 8-0

[]
Um

Der SEE
I Tose
COC AAT

Mini

[TT]

Fie. 63.—Camallanides prashadi. Vulvar region

of female; lateral view.

i., intestine; v., vulva. Uterus, full of embryos,
omitted. The arrow points in the direction of the

head.

branches of the uterus, in mature females,
being filled from end to end with free
embryos.

We have considered it necessary to
make this form the type of a new genus,
on account of the following important
characters in which it differs from Camal-
lanus : —

(1) The structure of the chitinous
buccal valves, each of which
has two large thickenings,
giving the appearance of
two separate masses oi
chitin.

(2) The reduction of the “ tri-
dents” to simple, rod-like
structures.

328 Memoirs of the Indian Museum. [Vor. VII,

(3) The alate condition of the right spicule, and the presence of an accessory
piece, in the male.
(4) The presence of a tubular appendage, carrying the vulva, in the female.

A nematode has been described under the name Camallanus bungari from Bungarus candidus,
in Java, by MacCallum (1918). The description of this form is not easy to understand, but it
appears impossible to identify it with the species just described, the most striking difference being
that, whereas in the present form the vulva is situated on a very conspicuous outgrowth from the
body-wall, in C. bungari it appears to have had no prominent lips, and to have been so inconspicuous
that some doubt remained as to its position. At the same time it may be observed that the accompany-
ing figure in MacCallum’s paper (l.c., fig. 65) gives quite a different impression from the description,
and shows an arrangement of the uterus and vagina hitherto unknown in the genus Camallanus.

Family GNATHOSTOMIDAE, Railliet, 1895, emend. Baylis and Lane, 1920.
Subfamily SPIROXYINAE, Baylis and Lane, 1920.
Genus Spiroxys, Schneider, 1866.
Spiroxys annulata, sp. nov.
(Figs. 64, 65.)

Host: Chitra indica.’ Position: stomach. Locality: Budha Stream, Ludhiana,
Punjab.

This species, which was collected by Dr. Baini Prashad, approaches closely to
the only other Indian species of Spiroxys at present known (S. gangetica, Baylis and
Lane, 1920). In size, and also in the dimensions of many of its organs, it is interme-
diate between that species and the genotype, S. contorta (Rud.).

The lips, when seen in a lateral view, have a very similar shape to those of
S. gangetica, the dorsal and ventral lobes
being placed almost at right angles to the
middle lobe. In a dorsal or ventral view,
however, the thickness of the lips is rela-
tively much less than in S. gangetica, and
they are seen to be much more wedge-
shaped (fig. 64), so that the head has not the
Same square appearance in such a view.
The six pointed teeth on each lip (two on
each lobe), characteristic of S. gangetica,
are absent in the present species. The
cuticular thickening on the inner surface
of the middle lobe is well developed, but
‚olmm. does not form a distinct tooth at its apex.

Fig. 64.—Spiroxys annulata. Head of female : The total length is from 20 to 25°4mm.
dorsal view.

in the male, and 30 to 34 mm. in the female.
The maximum thickness is 0°5-0:6 mm. in the male, 0'8-0'85 mm. in the female.
The diameter of the lips, measured dorso-ventrally, is 0'I6 mm,, and their length
0:00 mm. The transverse striations of the cuticle are exceptionally coarse (up to

1922.] H. A. Bavzs AND R. DAUBNEY : Parasitic Nematodes. 329

0'025 mm. apart), forming a series of rings with prominent posterior edges, so that in
optical section the outline of the body has a rather saw-like appearance.

There is a well-marked cuticular collar (seen also in S. gangetica) just behind
the base of the lips (fig. 65, ¢).

The tail is unusually short (about 0:2
mm.) in both sexes. ‘I'he length of the oeso-
phagus (measured from the anterior extremity
of the lips) is 2°8-3°5 mm. Except for a short
portion at the anterior end, which is purely
muscular, its walls contain many pocket-like
glands arranged in several linear series. The
cervical papillae, which are prominent and
resemble small, backwardly-directed spines,
are situated at a little over I mm. from the
anterior end. ‘The nerve-ring is at 0'62—0°65
mm., and the excretory pore at 0'8 mm., from

the anterior end.

The inflation of the caudal alae in the
male is very pronounced. As usual in the
genus, they are joined anteriorly by an inflated
cushion of cuticle which passes over the
ventral surface in front of the cloaca. There is also a sucker-like preanal depres-
sion, produced by the well-developed caudal muscles, within the bursa-like area
thus marked out. There is nothing worthy of special notice in the caudal papillae,
which are present to the usual number (11 pairs) and arranged as in the other
members of the genus. The lateral papillae are, as usual deeply buried in the
inflated alae. The small pre- and postcloacal pairs are sessile. The spicules are
nearly equal in length (from 1 6 to 2:3 mm.), slender, cylindrical and finely pointed.

The tail of the female is sharply conical, with the tip bent ventrally. The
caudal papillae are at 0°17 mm. from the tip. The vulva is situated slightly behind
the middle of the body, at 13°8-15°5 mm. from the posterior extremity. The muscu-
lar vagina is very narrow, running forward from the vulva. The ova have a very
thin, membranous shell, measuring about 0°075 x 0°06 mm.

oJ] mm.

Fig. 65.—Spiroxys annulata. Head of fe-
male; lateral view.
c., cuticular collar.

Subfamily GNATHOSTOMINAE, Baylis and Lane, 1920.
Genus Tanqua, R. Blanchard, 1904.
Tanqua tiara (v. Linst., 1879).
This common form occurred in great abundance in monitors of the following
species :—
Varanus salvator,
V. flavescens,

V. nebulosus,
V. bengalensis.

330 Memoirs of the Indian Museum. [Vor. VII,

Tanqua anomala (v. Linst., 1904).

Hosts:
“Mud snake,”
Banded krait (Bungarus fasciatus),
“Green snake.”’

The specimens from the last-mentioned host were in bad condition, and their
determination is questionable.

Genus Echinocephalus, Molin, 1858.
Echinocephalus spinosissimus (v. Linst., 1905).

A few mature, though rather small, specimens occurred in the spiral valve region
of the intestine of Trygon (Hypolophus) sephen from the Chilka Lake on the Kast
coast of India. The description of this species by Baylis and Lane (1920) necessarily
omitted the measurements of the ova. The average size of the eggs in the present
material is 0:05 X 0°0375 mm. The host is a new one.

Genus Gnathostoma, Owen, 1836.
Gnathostoma spinigerum, Owen, 1836.
Specimens occurred in a fishing cat (felis viverrina) and in a leopard (Felis
pardus).
Superfamily TRICHINELLOIDEA, Hall, 1916.
Family TRICHINELLIDAE, Stiles and Crane, 1910.
Subfamily TRICHURINAE, Ransom, 1911.

Genus Trichuris, Roederer, 1761.
Trichuris trichiura (L., 1771).
(Syn. Trichocephalus dispar auctt.)

Host: Gibbon (Hylobates, sp.).

Trichuris suis (Schrank, 1788).
(Syn. Trichocephalus crenatus auctt.).

Host: Wild pig (Sus bengalensis), near Dinapore, Bihar.

Trichuris ovis (Abildg., 1795).
(Syn. Trichocephalus affinis auctt.)
Host: “ Antelope’’—probably the Indian antelope or black buck (Antilope
cervicapra).
Genus Capillaria, Zeder, 1800.
Capillaria columbae (Rud., 1819).
The collection contains a few specimens of this species, which were taken om
the intestine of a pigeon in company with numbers of Ascaridia columbae. The
worm has recently been redescribed by Irwin-Smith (1920) in Australia.

1922. | H. A. BayLis AND R. DAUBNEY : Parasitic Nematodes. 331

Superfamily DIOCTOPHYMOIDEA, Railliet 1910 ( fide Travassos, 1920 [?])
Family DIOCTOPHYMIDAE, Railliet, 1915.
Genus Eustrongylides, Jägerskiöld, 1909.
Eustrongylides, sp. (?)

Two larvae taken from a prawn at Karachi by Dr. Baini Prashad, February 11,
1915.

These two specimens, which we refer tentatively to the genus, measure respec-
tively 12°5 mm. and 13°25 mm. in length and 0:26 mm. and 0°33 mm. in maximum
thickness. The head is somewhat swollen and almost globular in shape, with a
maximum diameter of 0'2I-0'22 mm. The mouth is situated in a large depression,
on the border of which there are six small papillae. The second ring of papillae
characteristic of the adults of Hustrongylides has not been detected. The oesophagus
is of enormous relative length, measuring 7°9 mm. and 8°25 mm. in the two specimens,
or about two-thirds of the total length. It consists of a short, narrow, muscular,
anterior portion, measuring 0'43-0'45 mm. from the head-end, and a very long and
considerably wider, glandular, posterior portion. The tail is short (0‘13-0°15 mm.)
and bluntly conical, with a little cuticular button at the extremity.

The life-history of the genus Zustrongylides and its allies is at present obscure, but it has been
suggested that certain immature forms found in fishes are the larvae of Eustrongylides ignotus

Jägerskiöld, a species occurring in various fish-eating birds. So far as we are aware, no form found in
an invertebrate has hitherto been assigned to the genus.

Superfamily STRONGYLOIDEA, Weinland, 1858.
Family STRONGYLIDAE, Baird, 1853, s.s. Lane, 1917.
Subfamily DELETROCEPHALINAE, Raill., 1916.
Genus Diaphanocephalus, Diesing, 1851.

Diaphanocephalus willeyi (Linst., 1904) Railliet and Henry, 1909.
(Fig. 66.)
Syn. Kalicephalus willeyi Linst., 1904.

Examples of this species collected from the banded krait (Bungarus fasciatus)
form part of the collection. The species was described from Coluber helena and
Vipera russellii by von Linstow (1904) and has since been reported from T'yphlops
braminus by the same author (1906 b.) and from Bungarus fasciatus (1908). We have
followed Railliet and Henry (1909) in regarding the genus Kalicephalus, Molin, 1861,
as part of the genus Diaphanocephalus, Diesing, 1851.

From the generic diagnosis of Kalicephalus given by Molin (1861) it is apparent
that this genus was separated from Diaphanocephalus on account of (a) the absence
of the dorsal hump just anterior to the male bursa, and (b) the presence of a
papilliform outgrowth carrying the vulva in the female. This combination of
characters is not constant, as a reference to the descriptions of D. willeyz and D. minutus

332 Memoirs of the Indian Museum. [Vor. VII,

will readily show. There is no appreciable difference in the structure of the mouth-
parts, upon which it is usual to base new genera in this family.

The figure of D. willeyi is given in order to show the rudimentary “ leaf-crown,”’
or corona radiata, surrounding the entrance to the buccal capsule. The presence of

WU Jo

Fıs. 66.—Diaphanocephalus willeyi. Head of female; lateral view.

c.r., corona radiata; d., duct of dorsal oesophageal gland; s., wall of buccal capsule (optical
section).

this structure in a member of the genus Diaphanocephalus indicates a close proximity
to the members of the subfamily Strongylinae.

The genus Diaphanocephalus is here referred tentatively to Railliet’s subfamily
Deletrocephalinae, which, if we exclude the members of Skrjabin’s genus Kiluluma,
would appear to constitute a natural group.

Diaphanocephalus minutus, sp. nov.
(Figs. 67-09.)

Host: Cobra (Naja tripudians).

This is the smallest species so far encountered in this genus. The males measure
from 4°9 to 5°0 mm. in length and 0°2 mm. in thickness ; the females from 5°1 mm. to
53 mm. and 0‘21 mm. respectively. The dorso-ventral diameter of the head is 0°15
to 0°:16 mm. The head is laterally compressed. There is a distinct constriction behind
it. The mouth is furnished with three pairs of small papillae which are actually the
terminations of the three pairs of longitudinal parenchymatous bands characteristic
of the genus. These bands are lateral and spring from a collar which surrounds the
base of the buccal capsule. The collar is in the form of a horseshoe with the open
ends coming practically into apposition in the dorsal middle line. The buccal capsule
is about 0:2 mm. deep and is thick-walled. When viewed in optical section in the

1922.] H. A. BayLıs AND R. DAUBNEY : Parasitic Nematodes, 333

dorso-ventral position itis roughly oblong. The
central portion of the cavity extends some little
way into the oesophagus. The duct of the dorsal
oesophageal gland projects for more than half
the length of the buccal capsule. It is supported
by the wall of the capsule, which is considerably
thickened (figs. 67, 68) dorsally and ventrally
at its base s. Viewed laterally, the buccal
capsule appears roughly triangular, with the
apex of the triangle directed backwards. The
oesophagus is short and club-shaped, measuring
about 0°45 mm. in length, whilst its maximum
thickness is about 0°15 mm. The nerve-ring sur-
rounds the oesophagus at about 0°09 mm. from
its anterior end. The excretory pore opens at
about 0°38 mm. from the head end in the male,

olmm.

Fie. 67.—Diaphanocephalus minutus
Head of female; dorsal view.

and 0°44 mm. in the female. The bursa of the b., wall of buccal capsule: d, duct of
male is completely campanulate and not easily dorsal oesophageal gland; p., papilla; s.,

spread out. There are three lobes, two lateral

wall of buccal capsule in optical section.

and anunpaired dorsal. Themain trunk of the dorsal ray (fig. 69) is extremely large,
measuring about 0:2 mm. in length, and up to 0:07 mm. in thickness just before it gives off

Oo-lmm.

Fic. 68.—Diaphanocephalus minutus.
female; lateral view.

Lettering as in fig. 67.

Head of

the large externo-dorsal rays. The lat-
ter are given off high up (about 0°16 mm.
from the tip), and at a wide angle.
Almost immediately behind this point the
main trunk gives off a pair of stout,
curved branches which together form a
large horseshoe; the tips almost reaching
the margin of the bursa. At its tip the
dorsal ray divides into two short curved
branches, the tips of which are again
bifurcated. The lateral rays originate
from a common root and reach almost to
the edge of the bursa. The ventral rays
are long and slender and closely applied
to each other throughout their length
They reach the margin of the bursa.
Just anterior to the origin of the bursa,
there is a pair of latero-ventral papillae,
which are stalked, and, though small,
quite prominent.

Thespicules are equalandslenderand
measure 0'255 to 0'275 mm. in length.

334 Memoirs ofthe Indian Museum. [Vor. VIT,

They are grooved on the ventral surface and slightly recurved at the tip. An access-
ory piece is absent. The anterior limit of the male genital tube is about 1°2 mm.
from the head.

The tail of the female is drawn out
to a rather acute point and measures 0°28
to 03 mm. in length. The vulva is
situated in the posterior half of the body,
dividing the latter in the proportion of
12°5: 5. It is fairly prominent. There
is a short vagina about o'14 mm. long,
running forwards, from which are given
off two well-formed ovejectors which run
anteriorly and posteriorly respectively.
The combined length of the ovejectors is
from 0'450 to 0'5 mm. The anterior

Om. uterus runs to within 08mm. of the base

Fie. 69.—Diaphanocephalus minutus. Dorsallobe of the oesophagus, whilst the posterior
of bursa; dorsal view. rs
runs to within about 0'4 mm. from the
tip of the tail. The anterior uterus is
commonly folded round the intestine, while the posterior has usually only a single
bend. The ovaries commence at the points where the respective uteri bend, and run
towards the middle of the body to cross each other just anteriorly to the vulva. The
eggs in the uterus are thin-shelled, and measure 0:068 mm. in length by 0'031 mm.
in breadth. Their contents are unsegmented.

e.d., externo-dorsal ray.

Diaphanocephalus, sp.

One female in poor condition and a fragment of another female were collected
from a cobra, Naja tripudians. The complete specimen is larger than the members
of the preceding species, measuring 15°5 mm. in length and 0°43 mm. in thickness.
The head, which is laterally compressed, measures 0°27 mm. in dorso-ventral diameter.
The buccal cavity is 0'Ig mm. deep. The structure of the head is similar to that
described for Diaphanocephalus minutus. The oesophagus measures 0°485 mm. in
length by 0'I6 mm. in thickness. The vulva is situated in the posterior portion of
the body, dividing it in the ratio of 6:5. Its lips are modified into a papilliform
outgrowth which measures 0'I4IX0'08 mm. The ova in the uterus are thin-shelled
and measure 0'065 X 0'045 mm. Their contents are unsegmented. The tail measures
0°45 mm. in length and is acutely pointed.

So far the only species described from the cobra is the preceding one, Diapha-
nocephalus minutus. The specimens under consideration differ from that species in
their size and in the form of the vulvar opening.

Diaphanocephalus, sp.
Large numbers of females of a species of Diaphanocephalus were collected on
nine different occasions from Russell’s viper in Calcutta. There were no males.

1922.] H. A. Bayuıs AND R. DAUBNEY: Parasitic Nematodes. 335

The females measure from 16 mm. to 18 mm. in length and 0:46 mm. to 0°51 mm.
in thickness. The head measures 0'225 mm. in dorso-ventral diameter, is laterally
compressed, and is followed by a slight constriction. There are the usual three
lateral pairs of parenchymatous bands, terminating anteriorly in three pairs of
papillae.

The buccal cavity is similar in shape to that of D. minutus and is 0°21 mm. in
depth. The oesophagus is en massue and measures 0°5 mm. in length and 0:22 mm. in
maximum thickness. The nerve-ring surrounds the oesophagus at 0°35 mm. from
the anterior end of the body. The excretory pore opens at 0'4 mm. from the head.

The vulva is situated in the posterior half of the body, dividing the latter in the
ratio of 10°5: 7. Its lips are modified into a papilliform process, the actual opening
being on the summit of the papilla. This “ papilla” is smaller than in the preceding
species, measuring 0°095 X0°085 mm. There is a short transverse vagina, at right
angles to which the well-developed anterior and posterior ovejectors are given
off. |

The eggs in the uterus are unsegmented and measure 0:080x0'048 mm. The
anus is situated at 0°72-0°75 mm. from the tip of the tail, which is drawn out to an
acute point.

Discovery of the males may prove that this species and the foregoing are identi-
cal. The differences observed in the single specimen from the cobra are too slight to
warrant definite separation.

Family ANCYLOSTOMIDAE, Looss, IQII.
Subfamily ANCYLOSTOMINAE, Looss, 1905, emend. Lane, 1917.
Genus Ancylostoma (Dubini, 1843) Creplin, 1845.

The collection contains all the species of this genus known to occur in India.
These are fairly well-known, and we propose only to give lists of the hosts from which
they were collected, with brief comments where necessary.

Ancylostoma duodenale (Dubini, 1843).

Hosts:
Tiger (Felis tigris).
Fishing cat (Felis viverrina).
Lane (1917 b) has given an account of the occurrence of this species in the tiger.
He observed that the specimens, though mature, were somewhat stunted in size,
which may be an indication that the worm, now occurring mainly as a human para-
site, finds the conditions of living in the tiger adverse to the attainment of its full
size. The same failure to reach the normal size is observable in the material in the
present collection. From the tiger there are several specimens of both sexes; from
the fishing cat only a single female. The fact that this species can live in wild
Felidae indicates that it may eventually be discovered in the domestic cat (cf. A.
caninum and A. ceylanicum).

336 Memoirs of the Indian Museum. [Vor. VII,

Ancylostoma caninum (Ercolani, 1859).

Hosts:

Wild dog (Cyon dukhunensis).

Indian wolf (Canis pallipes).

Indian jackal (Canis aureus).

Indian fox (Vulpes bengalensis).

Indian desert fox (Vulpes leucopus).

Sloth-bear (Melursus ursinus).

Tiger (Felis tigres).

Leopard (Felis pardus).

Fishing cat (Felis viverrina).

Domestic cat.

Ancylostoma ceylanicum, Looss, 19117.

Hosts :

Civet (probably Viverricula malaccensis).

Tiger (Felis tigris).

Lion (Felis leo).

Leopard (Felis pardus).

Fishing cat (Felis viverrina).

Leopard cat (Melis bengalensis).

Domestic cat.

Wild dog (Cyon dukhunensis).

Indian wolf (Canis pallipes).

Sloth-bear (Melursus ursinus).

Red cat-bear (Aelurus fulgens).

Ancylostoma malayanum, Alessandrini, 1905.
Hosts:
Malay bear (Ursus malayanus).
‘ Bear”’— probably the Sloth-bear (Melursus ursinus).

Genus Galoncus, Railliet, 1918.
Galoncus perniciosus (v. Linstow, 1885) Railliet, 1918.

The collection contains one female only of this species, collected from the intes-
tine of a leopard (Felis pardus) in the Zoological Gardens, Calcutta.

We have nothing to add to the description given by Railliet beyond remarking
that the head is evidently retractile, and in the extended state the buccal capsule
appears relatively much larger in proportion to the width of the body of the worm.
The specimen figured by Railliet (1918) is in the retracted state. Doubtless the
powerful muscles attached to the lateral walls of the buccal capsule, which were
noticed by Railliet and other authors, are connected with the retraction of the
head.

922.] H. A. BayLıs AND R. DAUBNEY: Parasitic Nematodes. 337

This genus appears to form a link between the subfamilies Ancylostominae and
Necatorinae.

Subfamily NECATORINAE, Lane, 1917.
Genus Necator, Stiles, 1903.
Necator americanus (Stiles, 1902).

We have to record the occurrence of this species in a new host, viz. a young
African rhinoceros * (R. bicornis), which had lived in the Zoological Gardens, Calcutta
for a very short time. The animal was captured in the Tanganyika Territory
(formerly German East Africa) and was brought to India by Mr. E. W. Harper, to
whom we are indebted for this information.

Carefulcomparison of this material with specimens of Necator americanus from man
in the collection of the British Museum reveals no difference except that the female
specimens from the rhinoceros are slightly the longer. They measure from II to 13
mm. in length and 0°4 mm. in thickness, as against 10 to 12 mm. and o’4 mm. res-
pectively in the case of the specimens from man.

Tae subfamily Necatorinae was proposed by Lane (1917 a) to replace the older
subfamily Bunostominae, Looss, 1911. The difference between these two groupings is
that, according to Lane, the genus Uncinaria approaches more nearly to the Necator
and Bunostomum group than to the Ancylostominae, among which it was placed by
Looss. It is interesting to recall that of the subfamily Necatorinae, if Uncinaria be
left out of account, all the members except Necator occur in herbivorous animals
only,' and, in consequence, the occurrence of Necator in the rhinoceros is not so asto-
nishing as it might appear at first sight. All the species of Ancylostoma occur in
carnivores, and all except A ncylostoma duodenale and A. ceylanicum in carnivores only.
It seems probable, therefore, that the original hosts of the species now found in man

were carnivores. It is also almost certain that Necator americanus was introduced
into America with the African slaves, and if this is the case, man may have acquired
his earliest infestations with this parasite from some wild herbivore inhabiting Africa.

Family TRICHOSTRONGYLIDAE, Leiper, 1912.
Subfamily TRICHOSTRONGYLINAE, Leiper, 1908.
Genus Haemonchus, Cobb, 1808.
Haemonchus contortus (Rud., 1803).
This species occurred in the Markhor (Capra falconeri) in the Zoological Gardens,
Calcutta.
_ Haemonchus cervinus, sp. nov.
The collection includes several females of a species of Haemonchus from a spotted
deer (Cervus axis). The specimens are in poor condition. They measure from 13 to

1 Since the preparation of this paper, Necator has been recorded from the pig in Trinidad (Ackert and Payne, Amer.
Jl. of Hyg., 11, 1, Jan., 1922). The authors regard the form found in pigs as a new species, which they have named

N. suillus.

338 Memoirs of the Indian Museum. [Vor. VII,

15 mm. in length and up to 0-4 mm. in thickness. The body tapers uniformly ante:
riorly to a small head which measures 0'023 to 0°025 mm. in diameter. The mouth
contains the single small lancet characteristic of the genus. The cervical spines are
situated at about 0°37 mm. from the anterior end. The oesophagus is slender, and
measures about 1°2 mm. in length. It is encircled by the nerve-ring at about 0°25
mm. from the anterior end. The excretory pore is just behind the level of the nerve-
ring.

The vulva is slightly prominent, but possesses no overhanging anterior lip such
as is found in H. contortus. It is situated at about 11 mm. from the anterior end.
There is a short, transverse vagina, from which well-developed anterior and posterior
ovejectors are given off. The eggs in the uterus measure 0:08-0:09 mm. X 0'04-0'05
mm., and their contents are unsegmented. The tailis long and slender. The anus is
situated at about 0°37 mm. from the tip.

In addition to the females this batch of specimens includes one male, The bursa
of this specimen is incomplete, and the whole worm so badly damaged that we have
found it impracticable to give a description of the male.

Family METASTRONGYLIDAE, Leiper, 1908.
Subfamily RICTULARIINAE, Hall, 1913.
Genus Rictularia, Fröl., 1802.
Rictularia, sp.

A single female specimen, taken from the intestine of a palm-civet (Paradoxurus
hermaphroditus bondar,' though labelled “ Paradoxurus mger’’), caught in the
Museum compound, Calcutta.

Rictularia plagiostoma (Wedl) has been recorded as a parasite of a palm-civet
by Leiper.” The present specimen, however, does not agree with the careful descrip-
tion of R. plagiosioma given by Jägerskiöld (1909), and we refrain from attempting to
attach a specific name to it.

Fam. incert.
Genus Scolecophilus, nov.
Scolecophilus lumbricicola, sp. nov.
(Figs. 70, 71.)

Host: an earthworm (Perionyx m’intoshi, Beddard). Locality: Nepal Valley.

This nematode was found by Dr. J. Stephenson in the body-cavity of the host,
in the tenth and eleventh segments. He noted that in the former segment they
were surrounded by masses of what appeared to be the spermatozoa of the host.
The material is not in a perfect state of preservation, having, no doubt, been removed
from the host after death. We are unable, therefore, to give a very complete
account of the anatomy. The species, however, shows certain remarkable features
which make it worthy of a brief description.

1 See Robinson and Kloss, Rec. Ind. Mus., XIX, Part IV, 1920, p. 178.
2 Proc, Zool. Soc. London, 1911, p. 620.

1922.] H. A. BayrLıs AND R. DAUBNEY : Parasitic Nematodes. 339

The male measures 3'65-4°I5 mm. in length and 0°4-0°5 mm. in thickness; the
female 6°0-5°5 mm. and 0'5-0'6 mm. respectively. In general shape (fig. 70) the
worms are rather short and stout. The
male has its blunt tail strongly curled
ventrally, while the posterior end of the
female is straight and conical. The
anterior end tapers more gradually than
the posterior. The cuticle is thin and
smooth, except in the region of the lateral
fields. These are very conspicuous, being
very broad and of a granular appearance,
and the cuticle covering them, especially
near the anterior and posterior ends, is
thrown into strongly-marked transverse
furrows. The width of the lateral fields
is about oI mm. anteriorly, increasing
posteriorly to about 0°22 mm. Near the
tail they bend round towards the ventral
surface. The musculature is apparently
of the meromyarian type. The head is
somewhat abruptly truncate. The
mouth shows no recognizable lips or

Fic. 70.—Scolecophilus lumbricicola. Female ;
papillae. The oesophagus is slender, lateral view.

passing posteriorly into a relativelylarge, i., intestine ; oes.b., oesophageal bulb; ov., ovary ;
r., rudiment of second uterus; w., functional uterus ;
glandular bulb of almost oblong shape. „' vulva.

The entire oesophagus, including the

bulb, is 0°7-0°y6 mm. long inthe male, 0‘8-0°9 mm. in the female. The bulb measures
about 0°35 mm. in length and 0:17 mm. in diameter. It is connected with the
intestine by a narrow neck containing some kind of valvular apparatus. The
intestine is apparently modified into a “fat-body”’ somewhat resembling that of the
Mermithidae. An anal aperture appears to be absent, the intestine terminating
blindly behind. The nerve-ring is situated at about 0°13 mm. from the anterior end.
No excretory pore has been seen.

In the male, there are paired spicules and an accessory piece. The shape of the
spicules is highly characteristic, and is more readily conveyed by a figure (fig. 71)
than by description. Each spicule is broad at the base, and bent at right angles at
about its middle. In the males examined, the spicules were protruded as far as the
bend, and had their tips directed laterally, as shown in the figure. The tip of each
spicule is bifurcated, ending in two sharp points of slightly unequal length, separated
by a deep cleft. The spicules measure 0:18 mm. in length (following the bend).
The dorsal portion of the accessory piece is roughly triangular, broader behind than
in front, and appears to send down lateral processes at the sides of the spicules. No
caudal papillae have been made out.

340 Memoirs of the Indian Museum. (Vor. VII,

In the female, the vulva is situated at about o'r8 mm. from the anterior end.
Its position is usually marked by a sudden change in the diameter of the worm, the
portion anterior to it being much narrower than the rest of the body. The general
arrangement of the female organs is shown in fig. 70. There is no ovejector or
muscular vagina. Only one genital tube is complete and functional. The uterus
runs back almost straight for some three-quarters of the length of the body, and
shows little accumulations of imperfectly-formed ova here and there along its course.

Fig. 71.—Scolecophilus lumbricicola. Tail of male; ventral view.
a.p., accessory piece; s., left spicule.

The coils of the oviduct and ovary are confined to about the posterior third of the
body. The other uterus appears to be represented by a blind sac, lying alongside of
the anterior portion of the functional uterus, and serving as a reservoir in which the
fully-formed ova are stored before being laid. This sac may run back, in large
specimens, to a point about 2°5 mm. from the anterior end, and is generally crowded
with eggs. These are of oval shape, with a thin shell, measuring about 0°065 x 0°038
mm., and containing a crescentic embryo.

The absence of one branch of the uterus is not uncommon among nematodes, and is most frequently

seen in cases where the vulva has been displaced far from its “ primitive’? median position towards
either of the extremities of the worm. This modification is met with among certain free-living forms,
as well as in several families, or isolated genera and species, of nematodes parasitic in vertebrates. But
there is one group of forms (all of which are, at least during a certain phase of their existence, parasi-
tic in invertebrates), to which the present form is perhaps most closely related. This is the group
including Allantonema, Atractonema, Bradynema and Sphaerularia. These forms are generally placed
among the Mermithidae, though it seems doubtful whether such a classification would bear critical
examination. They do, at all events, share with Mermis and its close allies, with the present species,
and with one or two other isolated genera (Aprocta, Aproctonema), the peculiarity of having no posterior
opening to the intestine in the adult form. The alimentary canal is, moreover, in these forms, either

1922. | H. A. BAyLis AND R. DAUBNEY : Parasitic Nematodes. 341

entirely absent in the adult female, or reduced, as in the Mermithidae, to a more or less solid organ
funetioning as a reserve of food-material. Again, in all these forms the female genital apparatus is
reduced to a single uterus and ovary. The vulva, however, is not anteriorly but posteriorly situated.
The worms are either protandrous hermaphrodites, or the sexes are separate, only the females and
young larvae being parasitic. A position in some respects intermediate between the present form and
the group referred to might perhaps be assigned to the genus Aproctonema, Keilin, 1917. This is
represented by À. entomophagum, Keilin, of which the separate sexes are both parasitic in the larvae
of a Dipterous insect. The oesophagus of Aproctonema has a somewhat similar structure to that of
Scolecophilus. The intestine (which has an anterior diverticulum), is a solid ‘ fat-body ”, and has no
posterior opening. There are two opposed uteri in the female, and the vulva is situated somewhat
behind the middle of the body.

A sac-like vestige of the second uterus, somewhat like that described above for Scolecophilus,
has been observed in Sphaerularia, and also in some of the free-living “ monodelphic”’ forms. It
appears in some cases to act as a receptaculum seminis rather than a reservoir for developing eggs, but
possibly it serves both purposes.

FORMS OF FREE-LIVING (ANGUILLULID) TYPE.
Genus Cephalobus, Bastian, 1865.
Cephalobus seistanensis, sp. nov.

(hiss! 72, 73.)

A number of minute nematodes were collected by Dr. N. Annandale and Dr. S. W.
Kemp from the tissues of the water-snail, Gyraulus convexiusculus, in reed-beds in
the Hamun-i-Helmand, Seistan, E. Persia. These specimens
may have been in the pulmonary cavity. They had a reddish
colour when alive, like that of the blood of the mollusc.
They probably belong to the genus Cephalobus, of which one
member, C. bütschlii de Man, is known to be, at least at times,
a parasite of certain fresh-water snails (Succinea), though most
of the species are free-living or to some extent parasites of
plants.

The present species has a length of 0:95-1:08 mm. in the
male and 1°3-1'43 mm. in the female. The maximum
thickness of the male is about 0'025 mm., of the female 0'035
mm. The oesophagus is from 0:2 to 0'27 mm. long, and is
composed of a long anterior muscular portion, narrow ante-
riorly and posteriorly, but somewhat swollen for about the
middle third, and a rather elongated, fusiform, posterior
glandular portion, or bulb, of larger diameter. This bulb
is not distinctly marked off from the rest of the oesophagus.
The nerve-ring surrounds the muscular portion at the back
of its swollen middle region.

The caudal end of the male is strongly coiled ventrally.
The tail is 0'05 mm. in length and tapers sharply to a fine
point. There are two equal, broad spicules, 0:026 mm. long,

LEHREN

SET]

B
E

Gas

UUS0-0

STE

Fic. 72.—Cephalobus se-
istanensis. Anterior end.

342 Memoirs of the Indian Museum. [Vor. VII,

AUESO:O

Fıs. 73.—Cephalobus seistanensis.
of male; lateral view.

and an accessory piece 0'015 mm. long.
The only caudal papillae appear to be
two small ventral pairs near the extre-
mity of the tail.

The tail of the female is tapering,
0:08 mm. long, and curves towards the
dorsal side. The vulva opens at about
0:55 mm. from the posterior end, %.e.
somewhat behind the middle of the
body. The single genital tube runs for-
ward for about 0°3 mm., and then doubles
upon itself to run straight backward, the
blind end of the ovary being situated at

Posterior end „pout 0'I-O'I5 mm. behind the vulva.

a.p., accessory piece; s.. right spicule. The uterus, in mature females, never

seems to contain more than one fully-

formed ovum at a time. Such ova have an unsegmented content and measure 0'045

x 0'025 mm.

The relative dimensions of this species, expressed according

to the formula of de Man, would be somewhat as follows: a = 38-

40:85 ; B=4 75-53 ; y=19 in male,
Genus Monhysterides, nov.

Monhysterides piscicola, sp. nov.

(Figs. 74, 75.)

Host: Mahseer (Barbus

River, Falakata, Eastern Bengal.
This is a very small form, the male measuring

17°8 in female.

tor). Locality: Torsa

3'5-4°0 mm. in length, the female 37-44 mm. The
maximum thickness is 0'I5-0'2 mm. The body is
slender and tapering towards each extremity, the mid-
dle region being relatively stout. The diameter of
the head is 0':03-0'04 mm. The cuticular striation is
exceedingly fine. Owing to the very small size of the
head the characters of the mouth are difficult to
determine. It appears to be surrounded by six small
nodules, two of which are probably lateral, two sub-
dorsal and two subventral. There are indications of
minute papillae near their bases. The oesophagus
consists of a short anterior portion which is transparent
and purely muscular, about 0:2 mm. long, and a longer
posterior portion which is partly glandular and partly
muscular, and is swollen behind. This portion may

Fic. 74.— Monhysterides pisci-
cola. Anterior end of female; lat-
eral view.

e., excretory pore; oes.b. oesoph-
ageal bulb.

1922.] ' H. A. Bayuıs AND R. DAUBNEY : Parasitic Nematodes. 343

probably be regarded as a pear-shaped bulb of an unusually elongate shape. It
measures about 0°32 mm. in length and 0'065-0'09 mm. in thickness posteriorly.
The anterior, musclar portion of the oesophagus appears to be divided transversely near
its posterior end, by a kind of diaphragm, into two parts of slightly different histological
appearance. The nerve-ring surrounds the neck of the “bulb” (not the muscular
part of the oesophagus, as is usually the case), near its origin, i.e.at about 0°22 mm.
from the head-end of the worm. The excretory pore is situated at about 0'4 mm.
from the anterior end.

In the male, the tail measures 0°34 mm. in length, and tapers to a very fine
point. There are two very unequal tubular spicules, of which the left measures 0:21
mm. in length, the right 0:08 mm. The
tip of the long spicule is bluntly rounded,
that of the short spicule more pointed.
There is no accessory piece. There are
nine pairs of caudal papillae, of which
four are preanal and five postanal. The
preanal papillae are all very close to-
gether, near the cloaca, and these and the
most anterior pair of postanal papillae are
very large and prominent, projecting
ventrally. Of the remaining postanal
pairs, which are smaller, one is laterally
situated, not far from the cloaca, and
the rest form a triangle, two being ventral

and one lateral, at about the middle of

the tail. Fie. 75.—Monhysterides piscicola. Posterior end
of male; lateral view.

The female has a finely tapering tail,
0°55-0°65 mm. long. The vulva is situat-
ed at about 0:45 mm. in front of the anus, and the uterus andovaryare single. The
latter is situated anteriorly, and is reflexed at about 1:2 mm. from the anterior end.
The worm is viviparous, the embryos being at first enclosed in large, oval, membranous
shells, measuring 0°275X0125 mm., but subsequently hatching in utero. The
voluminous uterus may contain at one time some 15 to 20 eggs containing embryos
in various stages of development, and about four or five free embryos. The latter
are about I mm. long, or roughly a quarter of the length of the parent.

1., left spicule; r., right spicule.

The systematic position of this species is somewhat doubtful, but we are inclined to regard it as a
member of the usually free-living family Anguiilulidae that has recently taken te a parasitic mode of
life. It does not show the specialization of the female genital apparatus which usually takes place in
true internal parasites, One of us (Baylis (1915)) has described two semi-parasitic species apparently
belonging to the genus Monhystera, and it is to this genus that the present form seems to approach
most closely in its general anatomy.

344 Memoirs of the Indian Museum. [Vor. VII,

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