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of the 


. Vol. III. No, 4, 

An Account of the 


based primarily on 


By lo-^^^'^^ 


.Tissiatant Superintendent, Indian J^useum. 


Calcutta : 


vSeptember, 1914. 

Miscellaneous Zoological Publications. 

Rs. As. 
Account of the Deep-sea liracliyura collected by 
the R.I.JI.S. "Investigator." By A. Alcock, 
M.H., C.SI.Z.S. .. .. .. ..60 

Account of the Deep-sea Madreporaria collected 
by the R.I.M.S. ' * Investigator. " By A. Alcock, 
M.B., C.M.Z.S. .. .. .. ..40 

Account pf the Triason (Hexactinellid) sponges 
collected by the R.I.M.S. " Investigator." By 
F. E. Schulze, Ph.D., M.D. .. .. .. 16 o 

Account of the Alcyonarians collected by the 
R.I.M.S. "Investigator." Part I. By J. Arthur 
Thomson, JI.A., and W. D. Henderson, M.A., 
B.Sc. .. .. .. .. .. 16 o 

Account of the Alcvonarians collected by the 
R.I.M.S. "Investigator." Part II. ByJ. Arthur 
Thom.sou, M.A., and J. J. Simpson, M.A., B.Sc. 20 o 
Aids to the identification of Rats connected with 

Plague in India. By W. C. Hossack, M.D. ..08 
Annotated List of the A.s'iatic Beetles in the Indian 
Museum. Part I. Family Carabidic, Subfamily 
Cicindehna?. By X. Annandale, D.Sc, and W. 
Horn . . . . . . . . ..10 

Biological collections of the R.I.M.S. "Investi- 
gator." List of Stations. 1884-191 3 
Catalogue of Indian Crustacea. Part I. — Introduc- 
tion and B'rachyura Priniigenia. By A. Alcock, 
M.B., LL.D., F.R.S. .. .. ■•70 

Catalogue of Indian Decapod Crustacea. Part I. — 
Brachyura. Fasciculus II. — Indian Freshwater 
Crabs— Potamonidac. Bv A. Alcock, C.I.E., 
M.B., LL.D., F.R.S. .. .. . . 14 o 

Catalogue of the Indian Decapod Crustacea. Part 
II. — Anomura. Fasciculus I. — Pagurides. By 
A. Alcock, M.B., LL.D., F.R.S., CLE. . . 14 o 

Catalogue of the Indian Decapod Crustacea. Part 
III.-^Macrura. Fasciculus I.— The Prawns of 
the Peneus Group. By A. Alcock, M.B., LL.D., 
F.R.S., CLE. .. .. .. ..70 

Catalogue of Indian Deep-sea Crustacea : Decapoda 
Macrura and Anomala in the Indian Museum. 
By A. Alcock, M.B., LL.D., C.M.Z.S... .. 10 o 

Catalogue of Indian Deep-sea Fishes in the Indian 

Museum. By A. Alcock, M.B., C.M.Z.S. . . 50 

Catalogue of Mammalia in the Indian Museum, 
Part I. By J. Anderson, M.D., LL.D., F.R.S. 
Part II. By W. L. Sclater, M.A., F.Z.S. .. 60 

Catalogue of Mantodea in the Indian Mu.seuni, 

Parts I and IL By J. Wood-Mason, F.Z.S., etc. 2 o 
Catalogue of Moths of India, Parts I to VII. By 

E. C Cotes and C Swinhoe, F.L.S., F.Z.S., etc. j J2 
Echinodcrma of the Indian JIuseum : Account of 
the Holothurioidea collected by the 
K.I.JI.S. " Investigator." By R. Koehler and 
C. Vaney . . . . . . ■ • ..160 

The above can be obtained from the Superintendent of the 
.V Sohu, 1 1 , Carlstrasse, Berlin. 

Rs. As. 

Kchinoderuia of the Indian Museum-: Littoral 
Holothurioidea collected by the R.I.M.S. 
"Investigator." By R. Koehler and. C Vaney 2 o 

Echiiujdernia of the Indian Musc\im : Deep-sea 
Ophiuroidea collected by the R.I.M.S. ' ' Investi- 
gator. " By R. Koehler .. .. . . 10 o 

Echiuoderma of the Indian Museimi : Shallow- 
water Ophiuroidea collected by the R.I.M.S. 
"Investigator." By R. Koehler .. ..40 

Echinoderma of the Indian Museum, Part \' : 
An account of the Deep-sea Asteroidea col- 
lected by the R.I.M.S. " Investigator." By R. 
Koehler . . . . . . . . ..120 

Echinoderma of the Indian Museum, Part VI : An 
account of the Shallow-water Asteroidea. By R. 
Koehler . . . . . . . . . . 20 o 

Echinoderma of the Indian Museum, Part VII: 

The Crinioids of the Indian Ocean. ByA. H.Clark 20 o 

Figures and Descriptions of nine Species of Squil- 
lida? from the Collection of the Indian Museum. 
By J. Wood-Mason, F.Z.S. , etc., edited by 
A. Alcock, M.B., C.M.Z.S. .. .. ..20 

Guide to tlie Zoological Collections exhibited in 
the Bird Gallery of the Indian Museum. By 
F. Finn, B.A., F.Z.S. .. .. . . o 12 

Guide to the Zoological Collections exhibited in 
the Fish Gallery of the Indian Museum. By 
A. Alcock, M.B., C.M.Z.S. .. .. ..08 

Guide to the Zoological Collections exhibited in the 
Invertebrate Gallerv of the Indian Museum. 
By A. Alcock, JI.B.,' C.M.Z.S. (Out of print.) 

Guide to the Zoological CoUection.s exhibited in the 
Reptile and Amphibia Gallery of the Indian Mu- 
seum. By A. Alcock, M.B., C.M.Z.S. (Out of 

Hand List of Mollusca in the Indian Museum 
Parts I and II, and Fasciculus E. By G. Kevill, 
C.JI.Z.S., etc. Index Parts I and II. By W. 
Theobold . . . . . . . . ..74 

Illustrated Catalogue of Asiatic Horns and 

Antlers in the Indian Museimi. By T. Bentham 2 o 

List of Batrachia ill the Indian Museum. By W. L. 

Sclater, M. A., F.Z.S. .. .. ..10 

List of Birds in the Indian Museum. Part I. — 
Corvidae, Paradiseidie, Ptilonorhynchidoe and 
Crateropodida;. By F. Finn, B. A., F.Z.S. .. 10 of Snakes in the Indian Museum. By W. L. 

Sclater, M. A., F.Z.S. .. .. ..10 

Monograph of the Asiatic Chiroptera and Catalogue 
of the Species of Bats in the Indian JIuseum. 
By G. E. Dobson, M.A., M.B., F.R.S. . . 30 

Mononraph of the Oriental Cicadidie, Parts I to 

VII. Ry W. L. Distant, F.E.S. .. .. ;i 14 


Calcutta, and from Messrs. Friedlander 

itthfi- Jnhlioitioiin eUiteil ami solil In/ the Siij>iiiiit»ii<lriil »/ thf Itulhin Miiseuiii (tilso 

;btiiinnble from Men.irs. Friedlander a Sohu) issued h,/ the Director 

of the Royal Indian Marine. 

Illustrations of the Zoologj- of the R.I.M.S. "Investigator," KS92. Fishes, Plates I to VII. Crustacea, Plates 
1 to V. if<94. Fishes, Plates VII to XIII. Crustacea, Plates VI to \III. Echinoderma, Plates I to III, 1895. 
Echinoderma, Plates IV and V. Fishes, Plates XIV to X\I. Crustacea, Plates IX to X\', 1896. Crustacea, 
Plates XVI to XXVII, 1897. Fishes, Plate XVII. Crustacea. Plates, XXVIII to XXXII. .MoUusca, Plates I to 
VI, 1898. Fishes, Plates XVIII to XXIV. Crustacea, Plates XXXIII to XXXV. ilollusca. Plates \II and Mil, 
1899. Fishes, Plates XXV and XXVI. Crustacea, Plates XXXVI to XL\", 19^'0- Pishes, Plates XXVi; to 
XXXV. Crustacea, Plates XLVI to XL^■III. Index, Part I, lyoi. Crustacea, Plates XLIX to LX. MoUusca, 
I'latts IX to XIII, 1902. Crustacea, Plates LVI to LXVII. Cru.stacea, Plates LXVIIl to LXXVI. Fishes, Plates 
XXXVl to XXXVIJI, 190^. Crustacea (Malacostraca), Plates LXXVII to LXXIX. Crustacea (Entomostraca), 
Plates I and II. JlolUisca, Plates XI\' to XVIII, 1907. Fishes Plates XXXIX to XLIIL Crustacea (Entomostra- 
ca), Plates 111 to \'. Mollusca, I'lates XIX and XX, 190S.— Re. 1 per plate. Mollusca, Plates XXI to XXIII, 1909.— 
As. 6 per plate. 


of the 


Vol. I, 1907. 

Part I —Contributions to the Fauna of the Arabian Sea. Records of Heniiptera and Hymenoptera from the Himalayas. 
Further notes on Indian Freshwater Entomostraca. The Fauna of Brackish Ponds at Port Canning, Lower 
Bengal, I— III. A Sporozoon from the Heart of a Cow. Miscellanea. 

Part II —Revision of the Oriental Stratiomyida;. Description of an Oligochsete Worm allied to Chistogaster. The 
Fauna of Brackish Ponds at Port Canning, Lower Bengal, IV. Further Note on a Polyzoon from the Himalayas. 
Reports on a coUection of Batrachia, Reptiles and Fish from Nepal and the Western Himalayas. The Fauna of 
Brackish Ponds at Port Canning, Lower Bengal, V. Notes on Oriental Diptera, I and II. Miscellanea. 

Part III —Report on the ilarine Polyzoa in the collection of the Indian Museum. The Fauna of Brackish Ponds at 
Port Canmng Lower Bengal, VI. A third note on E;arwigs (Dermaptera) in the Indian Museum, with the descrip- 
tion of a new species. Notes on Oriental Diptera, III. Description of a new snake from Nepal. Notes on a col- 
lection of marketable fish from Akyab, with a description of a new species of Lactarius. Description of two fresh- 
water Oligochaite Worms from the Punjab. Notes on Phosphorescence in Marine Animals. Notes on the rats of 
Dacca, Eastern Bengal. Notes on Freshwater Sponges, I — V. Miscellanea. 

Part IV —Nudiclava monocanthi, the type of a new genus of Hydroids parasitic on Fish. Preliminary descriptions 
of three new NycteribiidiE from India. Annotated Catalogue of Oriental Culicidae. Notes on Oriental Diptera. 
Notes on Freshwater Sponges, VI, VII. Description of a new Cyprinid Fish of the genus Danio from Upper 
Burma. Miscellanea. 

Vol. II, 1908-1909. 

Part I —The retirement of Lieut.-Col. Alcock, with a hst of his papers on Indian Zoology. The Fauna of Brackish 
Ponds at Port Canning, Lower Bengal, VII. Description of a New Dictyonine Sponge from the Indian Ocean. 
Notes on Freshwater Sponges, VIII. Remarkable cases of variation, I. Description of a new species of Lizard 
of the genus Salea, from Assam. The Fauna of Brackish Ponds at Port Canning, Lower Bengal, VIII. Des- 
cription of a new Cavernicolous Phasgonurid from Lower Siaiij. Descriptions of new species of Marine and 
Freshwater Shells in the collection of the Indian Museum. Notes on Oriental Syrphidae, I. Description of a new 
variety of Spongilla loricata. Notes on Oriental Diptera, V. Miscellanea. 

Part II — Gordiens du Musee Indien. The Fauna of Brackish Ponds at Port Canning, Lower Bengal, IX. Description 
of a new species of Danio from Lower Burma. Rliynchota Malayana, I. Cimex rotundatus, Signoret. Notes on 
Freshwater Sponges, IX. Fruit Bats of the genus Pteropus inhabiting the Andaman and Nicobar Archipelagos. 
A new species of Sun-Bird obtained near Darjiling. Three Indian Phylactolaemata. On two new species of Eagle- 
Rays (MyUobaUda:). Description of a new species of the genus Sesarma, Say., from the Andaman Islands. Des- 
criptions of new species of Land, Marine, and Freshwater Shells from the Andaman Islands. 

Part III —The Fauna of Brackish Ponds at Port Canning, Lower Bengal, X, XI. On some Oriental Solifugs with 
descriptions of new forms. The difference between the Takin (Biidorcas) from the Mishmi Hills and that from 
Tibet with notes on variation displayed by the former. On Caridina nilotica (Roux) and its varieties. Description 
of a new species of Charaxes from the Bhutan Frontier. First Report on the Collection of Culicidae and Corethridae 
in the Indian Museum, with descriptions of new genera and species. Miscellanea. 

Part IV Report on a collection of aquatic animals made in Tibet by Capt. F. H. Stewart in 1907, I. Notes on 

Aculeate Hymenoptera in the Indian Museum, I. Indian Psychodids. Description of a new species of mouse from 
the Madura District, Madras. Some Cleridas of the Indian iluseum. The Fauna of Brackish Ponds at Port Can- 
ning Lower Bengal, XII. Description of a new species of Saw-Fish captured off the Burma Coast. A new Sting 
Ray'of the genus Trygon from the Bay of Bengal. New Micro-lepidoptera from India and Burma. Notes on some 
ChrysomeUd Beetles in the collection of the Indian Museum. Six new CicindeUnse from the Oriental Region. Des- 
cription of a new slug from Tibet. 

Part F.— Revision of the Oriental Leptida;. Revised and annotated Catalogue of Oriental Bombylidas, with descriptions 
of new species. 

Vol. Ill, 1909. 

Part I. — The Races of Indian Rats. 

Part //.—Notes on Freshwater Sponges, X. Report on a coUection of aquatic animals made in Tibet by Capt F. H 
Stewart in 1907 II. Note on some amphibious Cockroaches. Description de quelques nouvelles Cecidomyies 
desTndes Desc'ript on of new land and marine shells from Ceylon and S. India Description of two new species 
oiCaranx from the Bay of Bengal. Remarks on some little known Indian Ophidia Remarks on some forms of 
Dipsadomorphus. A pelagic Sea-Anemone without tentacles. Rhynchota Malayana, II. 

Part ///—Notes on the Neuroptera in the coUection of the Indian Museum. New Indian Leptidae and BombyUdae, 
with a note on Comastcs, ol Sac, v. Heterostylum, Macq. Notes on the Trichoptera m the coUection of he Indian 
Museum Diagnoses of new species and varieties of Freshwater Crabs. 1-3. RfP°rt °n a smaU collection of . 
Lizards from Travancore. Descriptions of three new CicindeUna; from Borneo. The relation between fertility and 
n"rmahty°n Rats. Descriptions ^of a Barnacle of the genus Scalpellum from Malaysia^ Tndfa ^^e^hmTnar^'^te 
Polyctenids. Notes on Freshwater Sponges, XI. Descriptions of two new sheUs from S. India. Prehmmary note 
on a new genus of Phylactoltematous Polyzoa. MisceUanea. 

Part /F.-Description of a minute Hymenopterous insect from Calcutta. The Insect Fauna of Tirhut No '• descrip- 
tions of new species of Botia and N criaohilus . New Oriental Sepsinae. A new species of Fredencella from Indian 
lakes. oTalnSes of new species and varieties of freshwater crabs. No. 4- On some new or htUe-known Mygalo- 
morph spiders from the Oriental region and Australasia. 

Vol. IV, 1910. 

No. I. — Second report on the collection of Culicido; in the Indian Museum, with descriptions of new genera and species. 

Nos. II and III.— The Ihdian species of Papataci Fly (PhUbolomus). Taxonomic values in Culicidae. 

No. H'. — Revision of the Oriental blood-sucking Sluscida; (Sloiwjxintg, PhiltBinatomyia, Aust., aud Prislirhyitcliomyia, 

(gen. nov.). 
No. I'. — A new arrangement of the Indian AnopheUnx. 
No. 17 — A revision of the species of Tahanus from the Oriental Region, including notes on species from surrounding 

-Vo. I'll. — New Oriental Nemocera. Miscellanea : — Synonymy in Corethrinae. Indian Phlebotomi. 
Nos. VIII and IX. — .-V levi.sion of the Oriental species of the genera of the family Tabanidae oilier than Tabanus. Con- 

tiibutions to the fauna of Yunnan, Part VII. 
.Vo. .Y— .\nnotated Catalogue of Oriental Culicidae — Supplement. 

Vol. V, 1910. 

Part I. — The Hydroids of the Indian JIuseum, I. Notes ou Freshwater Sponges, XII. Descriptions of new Shells in 
the collection of the ludiau Museum from Burma, Siam and the Bay of Bengal. Materials for a revision of the 
Phylactotematous Polyzoa of India. Studies on the aquatic Ohgochaeta of the Punjab. An undescribed Burmese 
Frog allied to Uana ttgrina. Miscellanea. 

Part II. — Description d'Ophiures nouvelles provenant des dernidres campagnes de " I 'Investigator " dans I'Ocean 
Indien. Description d HolotUurios nouvelles apparteuant au Musee Indien. The races of Indian rats, II. Descrip- 
tion of a new species of Scalpellum from the Andaman Sea. Descriptions of five new species of marine shells from 
the Bay of Bengal. Notes on tish from India and I'crsia, with descriptions of new species. 

Part III. — -A. new genus of Psychodid Diptera from the Himalayas and Travancore. The Indian barnacles of the sub- 
genus Smiiium, with remarks on the classification of the genus Scalpellum. On a subspecies of Scutigerella uuguicu- 
lata, Hansen, found in Calcutta. The flistribution of the Oriental ScolopendridEe. Notes on Decapoda in the Indian 
Museum, I. Description of a new species of Memachiliis from Northern India. Notes on the larva of Toxorhynchites 
immisericors, Wlk. Description of a South India frog aUied to Rana corrugata of Ceylon. Contributions to the 
fauna of Yunnan, Introduction aud Part I. Miscellanea. 

Part IV. — Notes and descriptions of Indian ilicrolepidoptera. On some aquatic ohgochsete worms commensal in 
5pongiUa carter!. On Uothrinneiirum iris, Beddard. Notes ou nudibranchs from the Indian Museum. On the 
classification of the Potamonida; (Telphusidae). Catalogue of the pheasants, pea fowl, jungle fowl and spur fowl 
in the Indian Museum. On certain species of Palaemon from South India. Alluaudclla himalayensis, a new species 
of degenerate (/) cockroach, with an account of the venation found in the genera Cardax and Alluaudclla. 
Rhvnchota Malayana, III. 

Vol. VI, 1911. 

Part I. — A Rhizocephalous Crustacean from fresh water and on some specimens of the order from Indian seas. Deca- 
poda in the Indian Museum, II. Coutiibutions to the tauua of Yunnan, Parts II lo V. I'edipalpi in the Indian 
Museum. I and II. Six new species of .-liells Iruiu Heayal and Madras. Miscellanea : — Fieas from India aud China. 
Flies found associated with cattle in the neighbourhood of Calcutta. Mosquito sucked by a midge. Large egg laid 
by a beetle. 

Part II. — Some sponges associated with gregarious molluscs of the family Verraetidae. Aquatic animals from Tibet, 
III. Cyprinidae from Tibet and the Chumbi Valley, with a description of a new species of Gymnocyprts. New 
species and varieties of Cru.stacea StomatopoJa in the Indian Museum. The developmeut of some Indian Ascala- 
phidae aud -Myrmeleouidae. Miscellanea: — The occurrence of ZJnc(>/o/'iHS ci^n, Risso, in the Himalayas. Note on 
.Aquatic Khynchota. 

Part III. — Nouvcaux Chironomides de I'lndian Museum de Calcutta. 

Part I V. — Indian Isopods. Systematic notes on the Ctenostoraatous Polj zoa of fresh water. Some aquatic Oligochaeta 
in the Indian Museum. Contributions to the fauna of Yunnan, Part VI. Asiatic species of Crustacea Anostraca 
in the Indian Museum. Freshwater sponges, XIII. Miscellanea : — Synonymy in Corethrinae. The distribution 
of the different forms of the genus Ibla. 

Part V. — Revision of the Oriental Tipulidae. Five new Indian species of Clavicoru Coleoptera. Les Chironomides 
(Tendipedidae) de I'Himalaya et d'Assam. The occurrence of .-ipus in Eastern Asia. 

Vol. VII, 1912. 

Part I. — Deep-sea Fish obtained by the R I. M.S. S. "Investigator" during 1910-11. Decapoda in the Indian Mussum, 

III. Fauna of Paresnath Hill. Observations on the sh dlow-waier Fauna of the Bay of Bengal made ou the 

" Goldi-n Crown," igoS-t*). Coccidae in the Indian Museum, I On the nervous system of Ampullana globosa. 

Description of a species of Phoridae that causes Myasis iu man. Miscellanea : — Mimicry of a Mutiliid by a spider. 

Capture of Limulus ou the surface. Development of the larva of Lingula. Distribution of s )!iie Indian aud 

Burmese Lizards. 
Part II. — The Mosquitoes of Calcutta. Freshwater sponges, XIV. I'edipalpi in the Indian Museum collection. III and 

IV New Preahwater Crab from S. India. Decapoda in the Indian Museum, IV. Fauna Symbiotica Indica, I 

and II. Invirteb.ate Fauna ol the Kumaon Lakes. Fauna bymbiotica Indica, III. The Indian Mud-Turtles. 

Anatomy of Atopo'i sangutnolenla. Cestodcs fiom Indian Fishes Miscellanea: — Entoprocta in Indian Water.s. 

Rearing of Leeches iu liara Banki District. Habits of Tiger- Beetles. Schnodactylus mons:rosus as bait for birds. 

Macrones menoda var. trachacanthus. Aquatic Tortoises of the Ganges and Brahmaputra. 
Part III. — Gordiens du Musuc Indien. A new Tortoise from South India. New species of Branchiodrilus aud other 

aquatic Ohgochaeta, with remarks ou cephalizalion iu the Naididae. Fauna Symbiotica Indica, IV. Freshwater 

Medusa from the Bombay Presidency. A new species of Indian Thrips. Aquatic Chelonia of the Mahanaddi. 

Recent Criuoids from the Indian Ocean. Fauna of Yunnan, VIII. Earthworms. Asiatic Naiades in the Indian 

Museum. Miscellanea : — Malaria .Mortality in the Fringe Area of Calcutta. Larval Habits of Toxorhynchites 

immisericors. East .-\sialic species of Apus. 
Part I V. — Notes on the surface living Copepoda of the Bay of Bengal. Freshwater sponges of the Malabar 2kme Notes 

on the Habits aud Distribution of Limnocnida indica. The Anatomy oi melo indicus. Two new species of Scolo- 

Part V. — Indian Psyllidae. Descriptions of .some new species of Freshwater Fish from North India. New Oriental 

Diptera, I. 

All Account of the 


P. 319 line 1, for " Taenioers" read " Taeniocerus" 
i ;i:^l line 21, for "Fwiiului," read " Ainitulue"' 


Assislanl Siipcnntendent, Indian Museum. 



Bv F. H, Gravely, M.Sc.^ Assistant Superliitendeiu, Iniiaii Miiseuiii. 
(Plates XI— XIII.) 



1. Introduction . . . . . . . . . . . . 177 

2. External anatomy with special reference to tasonomic values . . . . 179 

3. Classification of Indo-Australiau Passalidae . . . . . . . . 191 

4. Catalogue of specimens in tlie collection of the Indian Museum, with notes on those 

in certain other collections . . . . . . . . . . 204 

5. The variable species . . . . . . . . . . . . 262 

6. Geographical distribution and synonymy . . . . . . . . 265 

7. Appendix I. A revised classification of the Aceraiinae . . . . . . 316 

8. „ II. Keys for the determination of species . . . . . . 318 

9. ,, III. The genus r«y^;M»jMS and a remarkable new genus from New Guinea . 326 

10. ,, IV. Supplementary catalogue of specimens in the Indian Museum collection 330 

11. Summary and conclusions .. .. .. .. .. .. 336 

12. Bibliography . . . . . . . . . . . . . . 340 

13. Index . . . . . . . . . . . . . . 345 


The present Memoir is in e£fect Part II of the " Annotated List of the Asiatic 
Beetles in the collection of the Indian Museum." Its scope is, however, much larger 
than that of Part I ; on this account and for other reasons which need not be detailed, 
it has been decided to discontinue the publication of the " lyist " as a separate series. 

After I had commenced the catalogue of our Asiatic Passalidae it soon became 
evident, on account of the present confused state of knowledge of the family, that 
the value to specialists of the locality-records it was to contain would be greatly in- 
creased by the inclusion of full illustrated descriptions of all the species referred 
to; and that if these were included its value to collectors in the East wottld be still 
further increased by the inclusion of a key for the identification of all genera known 
from the Oriental Region. 

This led to an enquiry into the general principles on which the classification of 
the family has been based; and to the conclusion that, by a little modification of 
Kuwert's system, the Passalid fauna of the whole Indo-Australian area, together 
with that of China and Japan, could be shown to be much more homogeneous and dis- 

^7^ Mi-moirs of the Indian Mitsruni. [Vol. Ill, 

tinctive than that system at first sight suggests. I have accordingly gi\en a 
revised classification with keys to all the genera known from this area, except Tcti- 
ijuinins, Kuwert. This genus is as yet only known from one imperfectly described 
specimen from New Guinea which 1 have not seen,' and it does not appear to be closely 
related to any of the genera found in the Oriental Region/ If it should prove that 
I have attemi)ted more than I can successfully carry out in the limited state of 
my pe'rsonal knowledge of the family as a whole, I can only say that the attempt 
appeared to me to be worth making, and that the descriptions and figures of the 
species I have seen (which will lose none of their value thereby) will, I believe, 
prove sufficient to prevent iny work from adding in any way to the confusion which 
it is intended in some measure to clear up. References to genera or .species that 
are not known to occur in the Oriental Region (by which is meant here India, 
Formosa, the Philippines, Borneo, and intermediate localities) or in China or Japan 
are enclosed in square i)rackets, both in the keys and in the account of the 
zoogeography. of the Oriental genera of Passalidae. 

Although the Indian Museum collection of Asiatic Passalidae. as I found it, was 
in many ways a remarkably fine one, additional collections that have been sent to 
me for examination have enabled me to make this paper much fuller than would 
otherwise have been possible. How far this is so will be sufficiently evident from the 
notes included in the catalogue of our specimens. I am greatly indebted to 
Mr. T. 15ainbrigge Fletcher and Captain R. B. Seymour Se well, who have collected 
valuable series of vSouth Indian species for me; to Mr. S. W. Kemp, who, with the 
assistance of Captain the Hon. M. de Courcy and the 32nd Sikh Pioneers, made 
an equally important collection in the Abor country; and to Mr. E. E. Green, 
Mr. H. E. Andrewes, Mr. H. Stevens, the Sarawak Museum, the Colombo Museum, 
the Bombay Natural History Society and the Imperial Agricultural and Forest 
Research Institutes, for the loan of their collections. It must not be suppo.sed that 
these collections include only the species definitely attributed to them in this paper, 
in which, as it is primarily a part of the •' Annotated List of the Asiatic Beetles in 
tlie Collection of the Indian Museum", I have not thought it necessary to refer to 
other collections when there seemed to be no special reason for doing so.' 

' Also, I have since learnt, from two specimens in the Deutsches Entomologisches .Museum, 
recorded l>y Zang from New Guinea without any de-cription (19066, p. 23). These show that Kuwert's 
6gure of this species is less trustworthy than his description. In general apjjearance they closely 
resemble species of the genus Lcptaitlax, from which they differ chiefly in the absence of scars from 
the nientum and in the presence of si.K well- developed lamellae on each antenna. See also Appendix 
III, pp. 32'> 330 below. 

'■ The locality of Semicyclus redtenbacheri, SloWcvka. (1H73) is prol)al)ly not Ceylon hut Brazil (see 
Kuwert, 1S9K. pp. 203 and 279) ; while that of Paxilloidcs schmidlii and philippinensis, Kuwert (Deutsche 
Ent. Zeit=chr., iSgo) is probably also Brazil, not the Philippine Islands (sec Kuweit. 1891, p 1S2 and 
1898. p i8i). Consequently the genera Scmicydm and Puxilloides are not regarded here as belonging to 
the Oriental Region. Nor have I accepted the record of Muslochilus polilus from Madras (see Stoliczka. 
1873, p. 156). 

■ Since this paper was sent to press I have had access to the collections in l[ inihnru, Berlin, and 

igi4.| F. H. Gravkly : Aji Account o/ the Oriental Passciliddc. xyg 

The Indian Museum collection of Passalidae was first arranged by Dr. Stoliczka, 
who published an account of it in 1873. He himself presented a considerable 
number of the specimens which it contains. Many of these specimens belonged to 
undescribed species; and .Stoliczka's descriptions of them added considerably to the 
knowledge of the family. Had he lived to discuss Kaup's classification (published in 
1871), as he proposed to do in the monograph he was planning, there can be little 
doubt that it would have been greatly to our present advantage. Owing to his early 
death on his way back from Central Asia it happened that Kuwert was the first to 
attempt a revision of the family. Unfortunately Kuwert appears to have been 
anxious merely to define briefly and conveniently genera and species, and not to 
have cared greatly for the problems of phylogeny or distribution ; consequently he 
failed to show some of the close relationships that exist among Indo-Australian 
forms, giving an undue value to the presence or absence of asymmetry in the head, 
at the expense of other characters which seem to be really more important. 

The value of Kuwert's posthumously published work is further reduced by the 
fact that it had evidently not received its final revision at the time of his death ; 
so that the text is in parts very difficult to follow, and some of the figures appear 
to be unfinished, while others are either wrongly numbered or incorrect, unless his 
descriptions of the genera they represent are wrong. Nevertheless, his paper includes 
by far the most complete classification in existence. 


As yet no one appears to have made a study of the relative values, for diagnostic 
purposes, of the various parts of a Passalid beetle. Instead, it has been assumed 
that all specimens differing markedly from one another in size or in such conspicuous 
characters as the form of the various ridges of the head, necessarily belong to 
different species. Zang (igo^a, pp. 163-4) has, indeed, pointed out how misleading 
variations in the ridges of the head may be in specimens belonging to the genus Chilo- 
mazus ; but even he failed to notice that this had been a constant source of difficulty 
in other genera as well. As the views I have been led to adopt by my work on the 
collections described below have caused me to suggest considerable reductions in the 
number of recognized species, I propose to state them here in giving an account of 
the terms used to designate the different parts of the insects. 

the British Museum, and I have to thank Prof. Kraepeliii, Prof. Voti Brunn, Herr Gebien, Dr. Horn and 
Mr. Arrow for the help they gave tne when studying these collections. I have also, through the kindness 
of Mr. Rene Oberthiir, and the Directors of the Museums at Darmstadt, Dresden and Stuttgart, 
received for examination certain of the type specimens preserved in their collections. As a result 
of this I have been able to complete this paper by the inclusion of keys to all species known from 
the Oriental Region, and to Australian genera ; to examine a number of specimens namgd by 
Kuwert; to complete my account ol the widely-distributed Oriental genus Accraiits by notes on 
Zang's species, most of which I had not previously seen ; and to add to and improve the paper in 
several other respects. 

i8o Memoirs of the Indian Museum. [Vol. Ill, 

Size and Proportions. 

Arrow's statement {1907, p. 441) that " A rather striking feature in which the 
Passalidae differ from the Lucanidae, as from wood-feeding insects in general, is their 
constanc)' of size ' ' has proved not to be invariably true ; and the fact that it does 
appear to be true of most species renders exceptions to the rule the more remarkable. 
These exceptions are of special interest from many points of view and have been dis- 
cussed as fully as possible below (pp. 262-5). It will be sufficient to point out here, that 
since the very species which are most variable in size have also proved to be the most 
variable in structure, the most abundant, and the most widely distributed species in 
each of the several genera to which they belong, the fact of a great difference in size 
between two specimens cannot be held to indicate that slight differences in unimportant 
characters are likely to prove to have a definite taxonomic significance 

The actual length of a Passalid depends in some degree on the extent to which the 
head is drawn into the prothorax, and the prothorax over the mesothorax, but it 
appears to be the most convenient indication of size that can be given. All my 
measurements of total length have been taken from the front of the labrum (or, in the 
genus Ccrnaipes, from the tip of the big horn) to the tips of the elytra. As the pro- 
portion of length to breadth seems to vary greatly in almost all species, no breadth 
measurements are given. 


Oriental PassaHds may be either symmetrical or asymmetrical. The degree of 
asynmietry may vary slightly in different specimens of a single species ; it seems to 
be an indication of the degree of specialization attained by different species or genera 
rather than of the degree of their mutual affinity. 

The antennae are always ten-jointed. The basal joint is always long and thick. 
Joints 2-4 are thinner and only about as long as broad ; the fourth occasionally bears 
a rudimentar)' lamella. Joints 5-7 alwaj's bear at least the rudiments of lamellae, 
but these are often so constructed that they appear to form only a continuous thicken- 
ing of the shaft when the antenna is in the least degree curled (see text-figure i, A and 
B), as it almost invariably is at this point in dried specimens. The three terminal 
jomts always bear lamellae. The structure of the antennae of any one species is 
usually constant within very narrow limits. The only exceptions to this that I have 
met with are in the species Aceraius laevimargo the antennae of which are remark- 
ably variable in form, and in a specimen of Aceraius grundis subsp. Iiirsitlns in which 
the fifth joint of the antennae is devoid of the well-developed lamella that it normally 
bears in that species. The lengths of the different lamellae relative to one another are 
used in distiuguisting the species of Macrolinus one from another, and in separating 
the genus A'(//</)/o/»s from the genus Labicnus; and I have used the same character 
in defining the (ionutas group of genera. Beyond this I have rarely found either the 
relative or actual lengths of the lamellae of any practical use; for they do not seem 
to be iibsolnlciy constant and they are ditlicult to c-()nii)are in the case of insects which 


F. H. Gravely : An Account of the Oriental Passalidae 


Text-figure i. 

Fig. a. Lcptaulax dcntatus, from above. 
,, B. ,. from below. 

C. Left mandible of Accnims grandis, subsp. hirsutus, viewed obliquely from the inner side. 

D. Anterior margin of head of Leptaulax dentaius, from below. 
, 'E. ,, „ ,. Macrclimis nicobariciis, 

P. Front co.xae and p:othorax of Leptaulax deniatns, viewed obliquely from in front. 
G. Front coxae and prothorax of Comacupes cavicornis, var. laevicornis, viewed obliquely 
from in front. 

,(.((. anterior angle of head; (Iam. anterior intermediate area of metasteruum; a.lJ. anterior lower 
tooth: an.i. basal joint of antenna; a. prs. anterior plate of prosteruum; i.t'.'. base of elytron ; f. coxa; c.a. 
central area of metasteruum; c.a.j. anterior part of hind coxa; can. caathus ; c.p.;. posterior part of 
hind coxa; c.i. central tubercle; 'e. elytron; / femur; /;-.<;. frontal area; fr.r. frontal ridge; g./. fourth 
groove of elytron; i.t. inner tubercle; La. lateral area of metasteruum; ibr. labrum; /. />. fused lateral 
plates of low'er surface of prothorax; I.t I. lowest terminal tooth ; m. mentum ; m.Lt. middle lower tooth ; 
ms.em mesothoracic epimeron ; ms.cs. mesothoracic episternum; mss. mesosternum ; mt.ein. metatho- 
racic epimeron ; metathoracic episternun ; «{./.i. middle terminal tooth: o.f. outer tubercle; />.».(/.. 
posterior intermediate area of metasteruum ; p.l.t. posterior lower tooth;'i. posterior plate of prester- 
num ; p.r. parietal ridge; pni. pronotum ; s. scar; sc. scutellum; s.oc. supra- occipital ridge; s.or. supra- 
orbital ridge; snt. suture; /. tibia; Ir. trochanter; /./. terminal teeth ; u.t. upper tooth; ii.Lt. upper- 
most terminal tooth; v.t. ventral tubercle; L]'L abdominal sterna. 

i8_' Mniioirs of tin- hiilhiii Miiscidii. [\'()I.. III. 

differ from one another in size or in the extent to which their antennae happen to be 
unfurled. It is so difliciilt, moreover, to say exactly where the shaft of the antenna 
ends and the lamella begins, that it often depends entirely on the point of view of 
the observer whether a specimen appears to agree with a given description. 


The mandibles of a Passalid are triangular in section, and bear teeth on the 
upper and inner margins as well as at the extremity. There is a striking uniformity 
of plan in the mandibles of all species, and such deviations as occur appear as a rule to 
be constant for whole genera rather than for individual species. These deviations 
take the form of the suppression, bifurcation or duplication of particular teeth, 
suppression being as a rule most marked in the right mandible and duplication in the 
left, no matter which side of the head is most developed. These teeth (see text- 
figure C I) may be described as follows: — 

One upper tooth, situated on the upper margin of the mandible. 

Three termiuiil teeth. Of these the upper two are always compressed laterally; 
the lowest one is compressed either laterally or dorso-ventrally according to the sec- 
tion of the family to which the insect in (|uesti()n belongs.' It is set further l)ack in 
the latter case than in the former. 

Three lower teeth. Of these the anterior one varies considerably in form in 
different genera; the middle one is alwaj's a chisel-shaped lamella which is hinged 
on to the mandible except in cert^iin genera of Aulacocyclinae; and the posterior one 
is a strong hollowed cusp so completely hidden between the labrum and maxillae 
that it cannot be satisfactorily examined unless the mandible is extracted. 

Upper su. face of head. 

The ])rincii>al features of the upper surface of the head are shown in text-figure 
I A. This surface is usually marked, at least in part, with a number of hair-bearing 
punctures, which in certain species run together to form grooves. The nature of these 
markings seems to be of greater importance than their extent. 

The head is bounded laterally above by a pair of supra-orbital ruli^es, which 
extend from the anterior margin above the eyes directly backwards. These ridges 
are usually of almost uniform height throughout the greater part of their length and 
obliquely truncate in front. In species in which the ridge is a broad one, its crest 
commonly forks above the truncation, one branch extending along the outer and the 
other along the inner margin of the anterior face ; but in a few of these species the 
inner branch is found to be more or less ob.solete in certain specimens. The size of 

• ' The Oriental Passalidae have been found to belong to two distinct sections of the family, which 
differ markedly from one another in many ways. One of these sections consists entirely of members of 
the sub-fnniily Aulacocyclinae, and it is often convenient to refer to this as the first section of the 
family, as ojiposed to the remainint^ subfamilies which to.^ether constitute the second section. These 
subfamilies are considered in the present paper to be four in number, and I have termed them I'leur- 
arii'iae, Aceraiinae, Onaphalocneminae and Leptaulacinae. See below, p. 191 and onwards. 

igi4.] F. H. Oravely : An Account of the Oriental Passalidae. 183 

the angle at which the upper and anterior surfaces of the supra-orbital ridges meet is 
moderate^ constant for each species, and is therefore of some taxonomic importance, 
especially in the Aulacocyclinae. This angle may conveniently be referred to as the 
apical angle of the ridge. In some species it is peaked, and in others always rounded • 
but as the peak is very small and particularly liable to be rubbed away, this is not a 
character on which very much reliance can be placed. In the genera Cylindrocaitlns 
and Aurituhis this angle is developed into a conspicuous horn, which may be termed 
the supra-orbital tubercle. 

The anterior end of the supra-orbital ridge is produced forwards in certain 
species ; the process thus formed is as a rule relatively longer in small specimens 
than in larger ones of the same species (compare fig. 20 with fig. 20b, and fig. 28 with 
fig 28(0- The part of the anterior margin of the head from which this process is 
developed may be called the anterioy angle of each side of the head. The size of this 
angle is often of some taxonomic importance. From it a canthns extends outwards 
and backwards, usually about half-way across the eye. This canthus may be either 
rounded or truncate distally ; in the latter case the size of its external angle appears 
to be of some slight taxonomic importance at least in the Aulacocyclinae. 

In the Aulacocyclinae, the posterior end of each supra-orbital ridge seems usually 
to curve inwards and then forwards, towards the base of the central tubercle which 
is present in all genera except Cylindrocaulus and Auritulus. This part of their 
course is developed to a particularly striking extent in the genus Caidifer. Some- 
times, however, even in the Aulacocyclinae, the inward prolongations of the posterior 
ends of the supra-orbital ridges evihce a tendency to coalesce behind the central 
tubercle. In the second section of the family the posterior ends of these ridges are 
always united by what may be termed a supra-occipital ridge, except in the sub-family 
Macrolininae in which this supra-occipital ridge extends outwards beyond them on 
either side, curves forwards, and then disappears'. 

All the Passalids which come within the scope of the present paper, except 
the genera Cylindrocaulus and Atiritulus, have a more or less prominent median 
tubercle somewhere near the middle of the head. This is called the central tubercle. 
It is as a rule more strongly developed in the Aulacocyclinae than in the remaining 
sub-families, and assumes in the former a number of different shapes even the details 
of which seem to be, as a rule, extremely constant in individual species or their local 
races. In the second section of the family on the other hand, it is no more than a 
longitudinal ridge more or less distinct and pointed in front, and such .structural 
variations as it presents are usually small and not abso'utely constant, and are very 
frequently obscured by friction. 

In the second section of the family a parietal ridge extends outwards on either 
side of this central tuberdle. The angle between these ridges is as a rule moderately 
constant in any one species, but is less constant than has sometimes been assumed. 

In the second section too, there is a pair of frontal ridges, which are never 

' In Ophrygonius singapurae the supra-occipital ridge unites the supra-orbital ridges, and is also con- 
tinued outwards beyond them. 

1S4 Memoirs of the Indian Mtiscitm. [A'oi, III, 

found in the first section. These extend obliquely forwards and outwards from 
the front of the central tubercle. Close to their origin they are in some species 
more or less fused, to form a short median keel. The extent of this fusicm, as 
well as the precise course followed by the ridges, has been found to vary greatly in 
different specimens of a single species (see, for exam])les, fig. iq, and figs 52 and 
526, c and (/). In spite of the great differences in the general appearance of an 
insect produced by these variations, they have rarely been found to have any 
taxonomic value at all ; and in these rare instances they have always been found in 
association with more distinctive, though perhaps less striking, characteristics in other 
parts of the body. 

In some forms the frontal ridges are replaced, or defined on the anterior and 
inner side, by a fine groove which extends almost up to the tubercles in which the 
ridges normally end, a little behind which it either disappears or bends outwards and 
a little backwards (see, for examples, figs. 25-39 inclusive). Another groove (which, 
though easily seen in most specimens of Aulacocyclinae, is more obscure in the other 
subfamilies, , especially after the insect has become hard and black) extends backwards 
on each side of the head from the anterior margin close to the inner side of the supra- 
orbital ridge, and curves inwards as though to meet the former groove, when it too 
disappears. There is usually an abrupt bend in the course of each frontal ridge 
opposite the place where the lateral grooves disappear even in species in which no trace 
of a frontal groove is visible. Consequently there can, I think, be little doubt 
that the lateral grooves, and those which sometimes follow the course of the 
posterior parts of the frontal ridges, together represent the suture by which the frons 
is separated from the parts of the head that lie on either side of and behind it ; and 
that the posterior parts of the frontal ridges always follow the course of this suture 
even when the suture itself is no longer developed.' 

The course of the division between the frons and the clypeus, or j^late imme- 
diately in front of it morphologically, is less evident, and it will be convenient to 
describe the characters of taxonomic interest in the two ])Iates together before going 
on to enquire into this. 

The pair of tubercles in which the frontal ridges end may be called the inner 
Uihercles, as they are always found either on the folded anterior margin of the head 
between a second pair of tubercles, the outer tubercles, or else behind these away from 
the margin. That the inner tubercles are morphologically the same, no matter which 
of these positions they occupy, becomes evident as soon as the lower surface of the 
folded anterior margin of the head is examined.^ 

' In pupae there is often a deep groove running direct from the position of the lateral grooves to the 
angle between the central tubercle and parietal ridges of the imago; but this appears to me to be 
no more than a fold in the pupal skin caused by the broad depression beneath it. Even if this fold should 
prove to appeal, in the first instance, along the line of the sutures that bound the frons in the larva, it 
would not necessarily prove the above conclusion incorrect, as the plates which develop beneath it may 
well be of a diflcretit shape from those which they replace-. 

' Before this can be done the whole of th>- Inhnim must be removed. 

1914] F- H. Gravely : An Account of the Oriental Passalidac. 185 

It is then found that in addition to the inner and outer tubercles visible from 
above, there is a third pair which may be called the ventral tubercles ; that the two 
members of each of the three pairs are normally connected by a more or less distinct 
ridge; and that similar ridges tend to connect the different pairs together at either 
end (see text-fig. i, D and E). None of these tubercles seem to be developed in 
the Aulacocyclinae ; but below the fold which forms the anterior margin of the head 
there is a ridge which probably corresponds to the one between the ventral tubercles of 
the remaining subfamilies. From a taxonomic point of view the most important of 
the ridges found in the second section of the family is that between the inner 
tubercles. Its absence in the Pleurariinae is one of the characters by which this 
aberrant sub-family is distinguished from others; and the fact that it forms the 
anterior margin of the head as seen from above is the principal character by which the 
Leptaulacinae are distinguished. In most forms in which it is present other than 
the Leptaulacinae, it is straight or slightly curved inwards towards the central tuber- 
cle ; but when it follows any different course this is usually found to be extremely 
constant, and in the genus Macrolinus it seems to be a remarkably good character by 
which to distinguish one species from another. The area enclosed by this ridge and the 
two frontal ridges is termed the frontal area ; it is very variable in shape on account 
of the variability of the frontal ridges. The other ridges are not sufficiently well 
developed to be of use in the diagnosis either of groups or of species. 

The outer tubercles are usually of very great importance', particularly in the case 
of asymmetrical forms, though the characters they afford must always be used with 
considerable caution on account of the frequency with which they are modified by 
friction, sometimes in what appear in other respects to be remarkably fresh and 
perfect specimens. 

In many species the outer tubercles, when viewed from the outer side, are found to 
be horizontally grooved, and I am inclined to regard this groove as the dividing line 
between the clypeus and frons (see text-figs, i D and E, and figs. 36«, S?*^, 38«, 39«.. 4°^' 
and 41(7). This groove is always situated below all that can be seen of the outer 
tubercles from above, and in species in which these tubercles are forked or trunctate at 
the end it always crosses the apex of the lower angle — a fact which often enables one to 
distinguish fresh specimens of such species from worn ones of others in which these 
tubercles are normally conical. From this it follows that the whole of these tubercles 
as seen from above belongs in reahty to what is here considered to be the frons instead 
of to the clypeus. The course of the suture between the outer tubercles is very obscure. 
If it is visible at all (of which I am doubtful) it must follow the ridge direct from one 
of these tubercles to the other. But the pupae of Leptaulax dentatus which I have 
examined, suggest that the whole of the upper surface of the front of the head of that 
species is frons, and that everything below the folded margin is clypeus^i.t'. that 
the junction of the two is along the ridge joining the outer tubercles together by 
way of the inner tubercles, and not along the ridge joining them direct. 

' The genus Gnaphalocneniis affords the most important exception I know. 

iSG Memoirs of the Indian M iistiim. [V(ii. IIT, 

The labnim, or upper lip, is attached to the margin of the clj-peus, and projects 
beyond the fold which forms the anterior margin of the head to an extent which often 
varies considerably in different specimens of a single species. The shape of its 
anterior margin, too, is only moderately constant, though the minute tooth which it 
bears in the middle in the genus Gnaphulocncmis and its immediate allies appears to be 
a character of some importance. In asymmetrical forms the left anterior angle is more 
or less distinctly prolonged beyond the right even when the right side of the clypeus is 
more highly developed than the left. Its upper surface is always covered with punc- 
tures and long hair in specimens which are in really good condition ; and in the Aula- 
cocyclinae an indistinct ridge usually crosses it transversely not far from the anterior 
margin. In certain species of Aulacocyclinae e g. Ceracupes fronticomis and C. austeni) 
this ridge instead of being indistinct is very pronounced. 

Low'er Stir j ace of Head. 
The only plate on the lower surface of the head, that need be considered here 
is the mentum, a broadly U-shaped shield situated immediately behind the mouth- 
appendages. The structure of this plate is of great importance in the Aulacocyclinae 
in the definition of genera, and in some cases in the definition of species also. In the 
remaining sub-families it is convenient to refer to the transverse median part of the 
plate as the central part, and to the whole of the paired side pieces, right back to the 
posterior margin, as the lateral parts. The former area is usually smooth or more 
sparsely punctured than are the latter parts, from which it is often separated by a 
pair of depressions or primary scars. These scars are always more or less round when 
complete, but more often than not their antero-external margin is imperfectly developed 
or obsolete, when they appear crescentic, the concavitj' always facing outwards and 
forwards. Their presence or absence is a character of importance for the grouping of 
genera together ; but they do not seem to be of any use in separating species one from 
another. In certain genera the whole anterior margin of the central part of the 
mentum is apt to be depressed ; in others this depression is moved backwards and 
outwards on to the surface of the mentum, in some genera' so far as to replace the 
primary scars, which are obliterated. Thepair of depressions thus formed may be termed 
secondary scars. When so well developed as to obliterate the primary scars these 
secondary scars are of great importance in classification ; when the primary scars are 
present as well they may be useful but are less constant, are often found to be 
only of specific value, and sometimes to exhibit a considerable continuous range of 
variation within the limits of a single species. 

The whole upper surface of the prothorax is covered by the pronolum. In 
most species this bears at least a trace of a median groove, which in some species is 
very pronounced indeed. Differences in the distinctness of this groove are so readil}- 

' The scars on tht inentuni of these genera were unfortunately confused with primary scars in my 
I0I2 |)rc1iiniiiary ])a])er. 

I9I4-] F- H. Gravely : An Account of the Oncntul Passalidac. 187 

produced by friction however, that but Uttle reUance can be placed on them in the 
absence of other distinctive characters. In addition to the median groove there is 
always a pair of marginal grooves, whose structure is often of importance. These 
grooves are situated close to the lateral margin, and extend upwards along the anterior 
and posterior margins, beside which, in certain species, they meet in the middle line. 
Near the posterior angle on either side is a depression or pronotal scar, a structure 
which is rarely of use in taxonomic work. The general surface of the pronotum is 
either smooth or covered with punctures so small and shallow that they are quickly 
removed by friction and are of little or no use in taxonomy ; but the sides are often 
strongly punctured especially in the neighbourhood of the scars, and of the ante- 
rior angles. The extent of this puncturing is always subject to some variation, and 
in certain of the species which vary greatly in size (e.g. Episphenus indicus) it is much 
more intense in small than in large specimens; nevertheless it often affords a useful 
confirmatory character in the diagnosis of a species. 

The prosternum , or median ventral plate of the prothorax, seems to me to be of 
but little taxonomic importance. It consists of a median piece, which is slightly 
expanded, ni front of and behind the sockets {coxal cavities) in which the front legs are 
inserted, to form an anterior and posterior plate, from the former of which a pair of 
large plates {?episterna) spread outwards, each in the form of a triangle attached by 
its apex. The median piece is often keeled in front, especially in the Aulacocyclinae ; 
but in this subfamily the posterior part of the keel is usually hidden by the project- 
ing coxae, 01 basal joints of the legs. The posterior plate is pointed behind in the 
first section of the family and broadly truncate in the second; in some species it 
usually bears hair-carrying punctures, but the presence of these has never proved 
to be constant. 

The plates of which the lateral parts of the underside of the prothorax are com- 
posed are completely fused, and may be collectively referred to as the lateral plates. 
Differences in the sculpture and pubescence of these plates afford useful confirmatory 
characters, but they are never of any great taxonomic importance. 


The scutelltim, or median dorsal plate of the mesotliorax, is partly hidden under 
the bases of the elytra , and the triangular area exposed between them is all that need 
be considered here. The base of the triangle forms the anterior margin and is invari- 
ably finely striato-punctate and pilose. Whether this punctured pilose area is visible 
depends partly on the extent to which it is developed, and this differs in different 
individuals of a single species as much as in individuals belonging to the majority of 
different species. It also depends on the extent to which the pronotum is drawn 
back over the mesothorax, and the taxonomic value of this area appears to me to 
have been greatly overrated by Kuwert. In some species other parts of the scutellum 
are punctured as well; the position of such punctured areas appears to be of greater 
importance than their extent. 

Below the bases of the elytra, which do not appear to be of any taxonomic 

i88 Memoirs of the Indian Museum. |\()i.. Ill, 

importance, are the mcsothoracic episterna, which are also triangular. A band of 
strong punctures often extends obliquely across the polished surface of these plates, 
from about the middle of the upper margin to the middle of the anterior margin, 
which it follows down to the lower angle The upper angle in front of this band 
is smooth and polished, and the posterior angle behind it matt. This pattern 
seems to form the basis of the patterns of all species, such variations as occur 
being confined above to the extension of the punctured band forwards to cover the 
upper angles, and below to changes in its width and in the extent of the dulled area 
in the posterior angles. These variations show a considerable degree of constancy 
in individual species, provided that the pronotum can be pressed forwards sufficiently 
to ensure that the whole of the episternum is properly exposed ; but in rigid speci- 
mens they are apt to be misleading. 

The mcsothoracic epimera are small plates which do not api)ear to be of any 
taxonomic importance. 

The mesosterniim lies between the two episterna of the mesothorax. It is 
bounded behind by a transverse groove, which extends right across the ventral surface 
of the body a little in front of the place at which the middle pair of legs is inserted; 
it is roughly triangular, but the apex of the triangle is flattened, and forms the 
short anterior margin, which is finely striato-punctate and pilose to an extent fully as 
variable as that of the scutellum. The surface in the lateral angles is often matt; 
that of the middle part of the plate is usually polished. Either or both of these 
parts of the plate may be punctured, and such puncturing often affords a 'useful 
guide to the identification of a specimen. The central part often bears strongly 
marked grooves, keels or hollows, to which considerable importance has been attached 
by previous authors. I have never found these characters to be of any taxono- 
mic value, and in some species (e.g. Macrolinus andamanensis) the series before me 
proves conclusively that they have absolutely none. The mesosternal scars are a pair 
of depressions situated close to the lateral margins of the plate. These depressions 
are also more variable than has been supposed, but they sometimes exhibit 
unusually well-marked characters of undoubted value (e.g. in Leptaulax anipunctus). 
They are present in nearly all species of the second section of the family , but are not 
found in the first section 

The homologies of the T-shaped or I-shaped structure, situated behind the trans- 
verse groove which I have taken to mark the posterior margin of the mesosternum, are 
somewhat obscure. The antero-lateral parts seem *o be as fully divided from the 
median part as they are from the mesosternum and perhaps represent the trochan- 
tinc or subcoxa, though they are more firmly fused to the adjoining thoracic plates 
than to the base of the leg. They are of no taxonomic importance. The median part is 
probably a superficial portion of the mesosternellum or medifurca. In the Aulacocy- 
clinae it usually appears to be I-shaped, with a complete suture across the middle-line 
behind it and a more or less obscure transverse roughened patch across the narrow 
piece between the bases of the legs. In the genus Ceracupes, however, the transverse 
])iece behind this patch is usually found to be more oi less completely fused to the 

iqi_^.] F. H. GRAVtXY : All Account of the Oriental Passalidae. 189 

inetastenium. This is also the case in all genera of the second section of the family, 
in almost all members of which the indistinct transverse band between the coxae 
is replaced by a distinct suture. This plate is therefore of some interest in connec- 
tion with the classification of the fatnily; but I have not found it possible to utilize 

it for differentiating species. 


The metathoracic episterna and epimera do not appear to present any charac- 
ters of taxonomic importance ; but the meiastemum, which covers the whole of the 
ventral surface of this segment of the body, must be considered in some detail. Four 
different regions can easily be recognized in this plate. Firstly, there is a large 
flattened circular central area , whose boundaries are as a rule less clearly defined in the 
Aulacocyclinae than in the remaining sub-families. This area is almost always smooth ; 
but in a few species of Leptaulacinae it is rough and may bear strong punctures, 
very variable in number, whose presence is apparently a constant feature of all the 
species in which they are found except one, that one being the most variable species 
of Passalid known to me. The central area is also marked, not infrequently, with ill- 
defined but symmetrically arranged grooves and depressions, to which taxonomic 
importance has sometimes been attached. But I have found these much too variable 
to be of any use. There is, however, in many species of the genera Tiberioides, 
Epispheniis and Leptaulax, a very persistent and strongly marked roughened depression 
in the middle-line, just behind the anterior margin, to which special attention may be 
drawn. It s rarely entirely absent in species in which it is ever distinct, and never 
very distinct in species in which it is normally absent. 

The metasternum is usually bordered on either side by a pair of depressed lateral 
areas. These may either be of equal width throughout or broader behind than in front, 
and differences both in width and in shape are usually found to be extremely constant 
within the limits of each species. The surface of these areas is always roughened or 
punctured. Between the central and lateral areas are the intermediate areas, vihich 
are divided into an anterior and posterior part, more or less completely according to 
the size of the areas on either side of them. Differences in the extent and nature of 
the puncturing of the posterior, and to a less extent also of the anterior, interme- 
diate areas, afford useful confirmatory characters, but usually vary too much within 
the limits of a single species to be diagnostic by themselves. 

Abdominal sterna. 
On either side of each abdominal sternum a more or less distinctly triangular 
depression is often found. These depressions, which we may term scars, are less 
persistently found in the posterior sterna than in the anterior ones, and in the Aula- 
cocyhnae than in the other subfamihes. They are too variable in shape and distinct- 
ness to be of use in the differentiation of one species from another ; and I have been 
unable to find anything connected with the abdominal sterna, except puncturing, 
which has any taxonomic value— such other characters as have been made use of 
have proved either to be variable in themselves, or to be dependent on the extent 

igo MciHoirs of the Indian Mu^cnm. [\'ol.. Ill, 

to which the abdominal segments have telescoped and sunk in beneath the ends of 
the elytra. Even the puncturing is variable as a rule. It occurs with distinct- 
ness chiefly in the Leptaulacinae and in one or two genera of Aulacocyclinae. In the 
former subfamily two types of puncturing can be recognized. One, which occurs in 
a single species only (Leptaulacides planus t and is the principal character by which 
that species can be recognized, is a uniform, close, shallow and moJerately fine punc- 
turing which occurs over the whole lower surface of the abdomen. The other, 
which occurs to some extent in other subfamilies also, is a close, but finer and 
stronger punctur.'ng, centred in the scars, beyond which it extends to an extent 
which is apt to be extremely variable even within the limits of a single species; this 
at best affords evidence for the separation of species into somewhat vaguely character- 
ized varieties. The puncturing found on the abdominal sterna of certain Aulacocycli- 
nae (eg species of the genus Comampes) is of yet a third kind, being coarse 
and sparse, with a hair (of which all trace may have disappeared in a worn specimen) 
rising in fresh specimens out of each puncture. The presence or absence of this type 
of puncturing appears to be a useful confirmatory generic character, and its extent, 
when present, a confirmatory specific one. 


The coxae, or basal joints, of the first pair of legs present one of the chief charac- 
ters by which species belonging to the subfamily Aulacocyclinae can be distinguished 
from those belonging to the subfamihes of the second section of the family ; for 
although they are always transverse and embedded in the prothorax over the greater 
part of their length, they project downwards distally to a greater extent in the 
Aulacocychnae than in the other subfamilies (compare text-figures i F and i G). No 
other joint of the first leg has proved to have any taxonomic value in any of the 
species I have seen, except the tibiae which are occasionalh- important, at least in 
the genus Taeniocenis. 

The coxa of the second leg is very small and need not be considered here, but 
that of the third leg resembles that of the first in size. Its exposed face is divided 
longitudinally into a raised anterior and depressed posterior part ; and the presence 
or absence of punctures on the latter affords a useful confirmatory specific character. 

Neither the small trochanters nor the longer femora of the second and third legs 
are of any taxonomic importance; but in the Aulacocychnae there is often, about 
two-thirds of the way down the outer side of the tibia of each of these legs, a small 
spine, the presence or absence of which can frequently be utilized in checking a specific 
identification. Of these spines, those on the hind tibiae are the more useful, as the 
extent to which they are developed differs more in different species. 

Elytra . 

The elytra of all Oriental Passalids are marked each with ten grooves, with ribs 

between them. On the dorsal surface the ribs are always flat, and much broader than 

the grooves, but laterally the grooves may be as broad as, or broader than, the ribs. 

Each groove contains, as a rule, a row of more or less distinct punctures, which are 

iqi4.1 F. H. Gravely : An Account of the Oriental Passalidae. igi 

usually stronger at the sides than above ; and when the side-grooves are broad the 
punctures are drawn out laterally so that they come to form a series of short transverse 
secondary grooves within the primary grooves. These secondary grooves are not of 
absolutely constant form in all specimens of a single species; but they are sufficiently 
constant to be of great taxonomic value. I have rarely, however, found the slight 
variations that occur in the distinctness of the punctures of the dorsal grooves of any 
value whatever, though Kuwert seems to have placed great faith in them. Another 
character to which Kuwert attached greater importance than it deserved, was the 
puncturing of the sides of the elytra of different species of the genus Aceraiiis. This 
puncturing, unless it occurs on the eighth rib (counting the innermost rib as the first) , 
as well as on the seventh and ninth, is by no means constant in extent in each species ; 
and although it is often most useful (especially in worn specimens) as a confirmatory 
character, as a primary character it is apt to prove misleading. The presence of 
hair on the sides of the elytra seems to be a satisfactory character by which to 
distinguish two geoera, Acerai-us and Trichostigmus, from their respective allies. 


The preceding account of the external morphology of Oriental Passalidae has 
already indicated that all species which come within the scope of the present paper 
fall into one or other of two widely different sections of the family. The first of 
these sections includes the Aulacocyclinae of Kuwert, together with the genera 
Auritulus and Cylindrocaiihis,whos^ affinities with the Aulacocycline genus Ccracupes 
have been established by Arrow (1907, p. 446). Kuwert, who overlooked the 
only known species of the genus Auritulus, defined the Aulacocyclinae only accord- 
ing to the apparent structure of the pro sternum relative to the coxae of the 
first pair of legs, excluding the genus Cylindrocaulus which he placed next to a Mexi- 
can form. The part of the prosternum between the coxae of members of the second 
section of Oriental Passalidae does not, unless the coxae are dug out from their 
sockets, appear as a lamina. Consequently it may be concluded that the "strongly 
elevated lamina ' ' , that Arrow mentions as separating the front coxae of Cylindrocaulus 
bucerus from one another, is an exaggerated form of the keel which always extends 
along the middle-line of the central part of the prosternum of the Aulacocyclinae. 
Assuming this to be the case, the two sections into which the Oriental Passalidae are 
primarily divided may be distinguished by the following characteristics' : — 

' Although the present paper does not deal with internal structure, reference must be mide here to 
Sharp and Muir's works on "The Comparative Anatomy of the Male Genital Tube in Coleoptera " 
(Trans. Ent. vSoc. London, I'iT2, pp. 477-642, pi. xlii-lxxviii, Passalidae, pp 570-580, pi. xliv, 
figs. ii-ij(!). These investigators found that in representatives of the genus Aulxcocydm, the only 
genus of Aulacocyclinae they examined, -'the basal-piece and the lateral lobes form one piece, either 
by consolidation or the supression of the basal-piece," whereas in representatives of the genera 
Gnaphalocnemis ( = Eriocnemis), Labienns and Prolomococlus (Gnaphalociietninae), Z,i'/)/a/«/rt,r (r,eptaula- 
cinae), and of the American genera Proc«/»s and Helens --the tegmen consists of two distinct pieces, 
the basal piece and the lateral lobes." 


Memoirs of the Indian Museum. 

[Vol. Ill, 

Section I . 

Section II. 


Plane of the lowest terminal tooth vertical, 
the tooth itself not situated well behind the 
two upper ones, directed forw.irds when 
the mandibles arc open. 

Parietal and frontul ridges, and the inner and 
outer tubercles with tho ridges associated 
with them, all absent. 

Plane of the lowest terminal tooth 
horizontal the tooth itself situa- 
ted well behind the two upper 
ones, directed inwards when the 
mandil)les are open. 

Parietal and frontal ridges, and the 
inner and outer tubercles with 
the ridges associated with them, 

Middle part of prosteruuni flat or 
with median groove or less dis- 
tinct keel, the whole surface 
between the coxae exposed, as 
the coxae do not project suffi- 
ciently from their cavities to meet 
above it. Posterior plate of pros- 
ternuni broadly truncate behind. 

Second and third | Tibiae each armed with a spine about two- Tibiae always unarmed except at 
pairs of legs. thirds of the way down the outer side, the tip. 

except in a few species in which the pos- 
terior tibiae are not so armed. 

Prothorax and its Middle part of prosteruuni with distinct median 
ap]>cndagcs. keel which is usually hidden behind by the 

coxae, as the distal ends of these project 
vertically from their cavities, their vertical 
inner faces touching one another in almost 
all species. Posterior plate of prosternuni 
more or less pointed behind. 

.Mention of other, but less distinctive, differences between the two sections of 
the family, which are referred to in the preceding part of this paper, need not be 
repeated here. 

Three genera of the first section [Ceracupes, Auritulus and Cylindrocauliis) differ 
markedly from all the rest; but although they are evidently related to one another 
more closely than to any other genus, they differ from one another so greatly that no 
single character not found in other genera of the section has yet been described as 
common to all three of them. I have therefore included these genera with all the rest 
in the s-ngle subfamily Aulacocychnae, although I think that a further examination 
of them may yet afford grounds for their separation. My principal reason for 
thinking this is that all known species of Ceracupes and Auritulus differ from all 
species I have examined of the genera Tristorthus, Taeniocerus, Comacnpes and 
Aulacocyclus , in having the middle lower tooth jointed on to the basal part of the 
mandible, as it appears to be in all genera of the second section of the family, 
instead of fused with it. Unfortunately I have not seen any specimens of the genera 
C'lii/tfer and Cylindrocaulus. 

The genera of the single subfamily Aulacocyclinae, of the first section of the 
family, may be distinguished from one another thus: — 

''Central tubercle not produced forwards to fuse with anterior margin 

of head ; upper tooth of mandibles not unusually long . . 2. 

Central tubercle very strongl>- developed, its apex fused with 
anterior margin of head ; upper tooth of both mandibles very long 
and slender; middle lower tooth moveable .. .. .. rcn/c;/ /'is, Kanp; pp. 2ij 

& -77 ■ 


F. H. Gravely : An Account of the Oriental PassaUdac. 193 

/^Central tubercle present; supra-orbital tubercles and anterior pro- 
1 longatioiis of supra-orbital ridges absent ; pronotum not drawn out 

into a bifid protuberance in front; middle lower tooth of mandible 
2. <(? always) immoveable •• •• •• •• ■■ • • 3- 

Central tubercle absent, head smooth and concave; supra-orbital 

tubercles and anterior prolongations of supra-orbital ridges present ; 
^pronotum drawn out into a bifid protuberance in front . . • ■ 7- 

'Supra-orbital ridges not united to form a median tooth behind 

central tubercle . ■ • . • • • • • • ■ • ■ 4- 

Posterior ends of supra-orbital ridges curved inwards to form 

together a median tooth situated immediately behind central tubercle ; 
^central tubercle elongated dorsally ; mesothorax smooth . . [Caultjcr, Kaup.] 

Meututn with strong median keel ; mesosteraum strongly punctured ; 

abdominal sterna with at least a few hair-bearing punctures . . Cninaaipcf;,^ Kaup ; pp. 

204 & 267. 
I Mentum not keeled; mesosternum at most feebly punctured; abdo- 
l minal sterna unpunctured . . . . • • • ■ • • • • 5- 

/ Central tubercle unidentate or tridentate above .. .. [Trisforl/i ns, ' Kawevt.] 

K Central tubercle bidentate above . . • • • • ■ • • • "• 

Central tubercle never pedunculate, often without any posterior face, 

anterior face never longer than dorsal, anterior end less highly elevated Taeniocenis, Kaup ; pp. 

208 & 270. 
Central tubercle usually pedunculate; otherwise always with dis- 
tinct posterior face, anterior face never shorter than dorsal, anterior 
end much more highly elevated than posterior . . . . Anliicocyclns, Kaup; pp. 

211 & 272. 
( Front coxae almost contiguous ; canthus extending about half way 
across the eye; supra-orbital tubercles flattened, expanded at the 
I apex, truncate .. .. •• •• •• .4»n7»/HS, Zang, p. 279. 

Front coxae widely separated ; canthus extending all the way across 

the eye : supra-orbital tubercles slender and pointed . . . . Cv/»f.!Voaj/«/;/.,s.Fairmaire; 

p. 279. 

The second section of the family appears to me to contain four Oriental sub- 
families (in addition to the Tarquiniinae, see below, pp. 326-330) which may be dis- 
tinguished thus:— 

/"Mentum without primary scars; secondary scars absent, or present 
only as depressions or small grooves close to anterior margin ^ ; outer 
j tubercles of head always simple . . . . • ■ • • ■ • . . 2. 

! Mentum with primary scars, except when these are replaced by 
more or less strongly developed secondary scars'; outer tubercles of 
l^head often complex . . . . • • • ■ • ■ ' ■ • ■ 3- 

I Since preparing this key I have examined the type of Comacupes minor. Heller, and specimens of 
C. /omcoft's, Kuw., from Borneo. These have proved to be, in several important characters, transi- 
tional between the genera Comacupes and Tristorthits (see below, p. 267). 

"^ I know of no really sharp distinction between these genera. The definition given here involves 
the transference of Taeniocenis deyrollei, and with it I presume T. masiersi (I have seen specimens of 
the latter determined by Zang, but not the original description), to the genus Aulacocyclus. These 
species appear to me to resemble A. rosenbergii more closely than they do any species of Taeniocenis. 

3 This distinction taken by itself is not altoj?:ether a satisfactory one, for in one race of Ophrygon- 


Memoirs of the Indian Museum. 

[Vol. Ill, 

/Only three antennal hmiellae recognizable when antenna is furled ; 
I ridge joining inner tubercles absent 

More than three antennal lamellae recognizable when antenna is 
furled ; ridge joining inner tubercles present 

PM?UR.\Rri>JAK; pp. 213 

Supra-orbital and supra-occipital ridges discontinuous 

.\cr:R.AiiN.\E ; pp. 215 & 

280.; pp. 240 


, Supra-orbital and supra-occipital ridges continuous ' . . . . . . 4. 

'Inner tubercles situated behind outer tubercles; ridge joining them 
not forming anterior margin of head; more than three antennal 
lamellae recognizable when antenna is furled . . . . OvTArnAi.ocNEMiNAE; pp. 

246 k 207. 
4,/ Inner tubercles situated between outer tubercles on anterior margin 
of head, the middle jiart of which is formed by the ridge joining 
them: only three antennal lamellae recognizable when antenna is 
furled .. .. -. ■- •• -• Lept.\ulacinae; pp. 251 

* 302. 

The first of these subfamilies, the Pleurariinae, contains only one Oriental 
genus, Pleurarins. This genus has been grouped bj' Kuwert with the genera Ninoides, 
Pertinacides and Epipcrtinax, all of which are confined to the New World. I have 
not seen specimens of any of these genera, so am not in a position to criticize his 

The next three subfamilies comprise between them the Macrolininae of 
Kuwert, together with all the groups which fall under the second number " O'' " of 
his table (1896, p. 219), which is trichotomous at this point. 

The definition of the Macrolininae given in the above key necessitates the 
removal, from the group to which Kuwert applied this name, of the genus Episphenus 
and one oriental^ species of the genus Tiberius, since these have no scars on the men- 

ius caniori the mentum bears ridges which somewhat resemble the margins of primary scars and are 
perhaps homologous with the margins of secondary scars. The form of these ridges is, howe\-er, quite 
unlike that of the secondary scars of any species of Gnaphalocneminae ; and the forni of the anterior 
margin of the head is in itself sufficient to show that the insect does not belong to any genus of that 
subfamily. On the other hand, the primary scars are sometimes so feebly impressed in the genera 
Cetejus and Aiialaches, that it is possible they may sometimes be absent, in which case it would be 
almost impossible to separate certain species from the genus Episphenus The fact is that these three 
genera are none of them, probably, very remote from the common ancestor of both subfamilies, 
although the countries inhabited by the last are so widely separated from those inhabited b)' the first 
two (see below pp. 313-5). The mentum of the genus Hyperplestheniis, Kuwert, which has only 
secondary scars and these not of very large size, closely resembles that of the genus Episphenus. The 
lateral and intermediate areas of the nietasternuni are, however, fusei as in all other genera of the 
Hyper plcsthcnus group, a fusion which is not known to occur in any genus of Aceraiinae. See also 
appendix III, p. 326, below. 

' See also p. 183, footnote. 

^ The so-called African species, Tiberius caffer, must also go. M. Oberthiir has shown me the 
type, which proves to belong to an Australian species, identified by comparison with the British 
Museum colkction as Pharochilus dilataUts, Dalm. 

igi4.] F- H. Gravely : An Account vf the Oriental Passalidae. 195 

turn and have the supra-orbital ridges continuous with the supra-occipital ridge. The 
Tiberius here referred to is T. ktiwerti, Arrow {=cancrus, auct.) and I have had to 
create a new genus Tiberioides for the reception of this species, of Chilomaziis borealis, 
Arrow, and of a new species {T. austeni) in the Indian Museum collection which com- 
bines some of the characters distinguishing the first two from one another. 

Kuwert's reason for including "Tiberius cancrus" and the genus Episphenus in 
the Macrolininae instead of among his "second ga ' ' groups, was that they were neither 
asymmetrical nor possessed of a mentum marked by grooves cutting off a small area 
("furchenbegrenztes Schild") from the rest. Either of these characters occurring 
without the other would have been sufficient to place them among the ' ' second ga ' ' 
groups, and they appear to me to be of much less importance than the two characters 
referred to above, in each of which they differ from the rest of the group in which 
Kuwert placed them. Moreover, the genus Episphenus closely resembles the genera 
Chilomazus {=Laches) and Basilianus in the texture of the upper surface of the head, 
and forms with them a series ranging from complete symmetry to a high degree of 
asymmetry, but otherwise remarkably alike. [See also below, p. 316]. 

Turning now to Kuwert's " second qa" groups, it is clear that of the grooves and 
depressions found in the mentum of different genera of his Ivachinae, only the lateral 
depressions, found nowhere except in the Far Eastern genera Mastochilus^ and 
Analaches,^ are primary scars. Consequently these two genera may be separated from 
the Oriental Chilomazus and partly Oriental Epilaches.^ On the same grounds the 
Oriental genus Heterochilus , which is without primary scars, can be separated from 
the remaining genera — none of thefn found in the Oriental Region — of the group in 
which Kuwert placed it, and placed near the Oriental genus Aceraius, a procedure 
which Zang has already advocated (1905a, p. 167) on different grounds. 

The new classification resulting from this rearrangement of genera , and necessi- 
tated by the definitions of the subfamilies Aceraiinae, Macrolininae and Gnaphalocne- 
minae given above, appears to me to be more natural than the old ; and all the 
changes advocated have the effect of bringing classification more nearly into line 
with distribution. 

The subfamily Aceraiinae is almost entirely confined to the Oriental Region, and 
its distribution, as will be seen later (pp. 311-313), follows such definite lines that the 
one striking case of discontinuous distribution which it appears to show — that of the 
genus Epilaches — wiU probably prove to be a mistake. E. filius, Kuwert, from the 

' Kuwert omits the scars in his figure of the type of meiitutn found in this genus. They are men- 
tioned, however, in Percheron's description of Passahis sexde.ntatus { = Mastochilus polyphyllus according 
to Kuwert) ; they are well developed in the specimens in the Hamburg Museum determined by Zang 
M. polyphyllus ; and Kuwert himself mentions in his descriptions of both species of the genus the presence 
of scars, which he describes in terms that seem unlikely to refer to the small grooves he figures close to 
the middle of the anterior margin. Concerning the correct form of the name of the genus (whether 
Mastochilus or MastacJiilus, see Zang, 19036, p. 418. 

* Concerning Kuwert's confusion of the names Analaches and Epilaches, see below p. 283. 


Memoirs of the Indian Museum. 

[Vol. Ill, 

Nilgiris, is almost certainly a synonym of Basilianus neelgherriensis ' ; and, as the genus 
Basilianus is most unlikely to occur in the St. Cruz islands, the probability is either 
that there has been some mistake about the locality from which the only other 
species — E. infantilis — comes', or else that this species has true scars on the mentum 
which have not 3^et been described. Consequently the genus Epilachcs is not included 
in the following key. The genus Heterochilus, for which Zang (19056, p. 154) has 
suggested the new name Rhipsaipis , does not appear to me to be sufficiently distinct 
for recognition, especially in view of Zang's notes upon it (1905^, p. 167). The genus 
Ophrvgonius, Zang, is very close to Basilianus, Kaup; but as the characters by which 
the two are separated do undoubtedly distinguish all the Indian forms on the one 
hand from the only Malaysian form with which they could be confounded on the other, 
I have found it convenient to retain it." 

The genera which I recognize in the subfamily Aceraiinae may be distinguished 
from one another, thus^: — 

; i-A as long as that separating 


times as long as that 

( Inner tubercles separated by a sf: 
outer tubercles 

Inner tubefcles separated by a 

' separating outer tubercles 
Anterior margin of head symmetrical or but slightly asymmetrical, 
both right and left outer tubercles simply pointed 

^ Anterior margin of head markedly asymmetrical, left external 

i tubercle longer than right, rounded or more or less obliquely truncate 

V^at apex 
Central area of mentum smooth, neither grooved nor depressed near 
middle of anterior margin ; anterior margin of head quite symme- 

Central area of mentum grooved or depressed round a median (rarely 
paired) process close to anterior margin; anterior margin of head 
more or less asymmetrical 

Frontal ridges represented by line grooves (which in certain speci- 
mens are somewhat obscured by their posterior margins being deve- 
loped into ridges} ; antennae with five or six pubescent lamellae ; 
lateral grooves of elytra narrow, their punctures never linear 
Frontal ridges neither represented nor accompanied by grooves ; 
antennae with only four pubescent lamellae ; lateral grooves of elytra 
broad, their punctures transversely linear .. 

Tibcrioidcs, Gravely 
2X5 & 280. 

217 & 


Chiloiiuiziis, Zang ;.pp 

& 28 1. 

227 & 285 


Zang ; pp 

' I have since been able to con6rm these two suggestions by reference to the type of /:. iiifanlilis 
and cotypes of /; fdiiis; see below, p. 283. 

'^ For a modification of the views e.'<pressed here see Appendix I, pp. 316-318. The page- references 
given in this key refer to the pages of the present paper on which are described the first species of each 
of the genera as here defined. The shifting of the points of division between the genera has not 
necessitated any change in the oider in which the species would have been described. 

1914] ^- ^- Gravely : .4/; Account of the Oviental Passalidae. 197 

Sides of elytra hairless, unpuactured except in the grooves . . Basilianiis, Kaup ; pp. 

220 & 282. 
Sides of elytra clothed with hair, lateral ridges more or less exten- 
sively punctured especially near the shoulder .. ^ .. Aceraiiis, Kaup ; pp. 228 

& 286. 

All species of the subfamily Macrolininae as defined here were placed by Kuwert 
in one or other of the genera Macrolinus and Tiberius. Zang (1905a, p. 163) has 
pointed out that Kuwert' s definitions of these genera are based on a distinction 
which does not really exist. Zang's definition, however, is based on a character 
which appears to me to be of specific rather than of generic importance. If the 
subfamily is to be divided into genera at all it must be along the lines indicated by 
Kuwert, but the definitions will have to be differently expressed, and Kuwert' s genus 
Macrolinus will have to be further subdivided into three sections, to each of which 
generic value must be given. It seems to me preferable, therefore, to reunite the 
genera Macrolinus and Tiberius. The single genus thus formed may then be divided 
into groups of species as follows : — (i) Macrolinus urus and diuvenbodei from Celebes ; 
(2) M. sikkimensis, nicobaricus, and andamancnsis \ (3) the Ceylon forms: and 
(4) the Malaysian and Philippine forms with M. sulciperfcctus from Celebes. The so- 
called African species, Tiberius caffer, of which I have examined the type, has proved 
to belong to the genus Pharochilus ; it is probably P. dilatatus from Australia. 

The subfamily Gnaphalocneminae contains the remaining genera belonging 
to the "second 9a" groups of Kuwert's classification, i.e. all except the genera 
Chilomazus { = Laches), Epilaches, Heterochilus , Basilianus, and Aceraius. It is most 
abundantly represented in the Far Eastern part of the Indo-AustraHan area, and the 
following revision of its classification is based on the collections in the Berlin and 
Hamburg Museums, in which Pelopides and Tatius are the only genera that are not 

In the Gnaphalocneminae, as in the Aceraiinae, some species are symmetrical 
and others highly asymmetrical; it is probable that the two subfamilies are 
very closely alUed, and that the former occupies the same position in the fauna of the 
Austrahan Region as the latter does in that of the Oriental Region. 

The classification of the Gnaphalocneminae has hitherto been based principally 
on the sculpturing of the mentum and of the anterior margin of the head. This 
sculpturing appears to be of primary importance, but it is very difficult to define 
some genera precisely with its help alone, and I have found it necessary to use also the 
structure of the mandibles. In the less highly specialized genera of both Gnaphalo- 
cneminae and Aceraiinae the dentition of both mandibles is complete and normal, 
all the terminal and lower teeth being present and approximately equidistant one 
from another; whereas in the more highly specialized genera the lowest terminal 
tooth and anterior lower tooth tend to fuse or to disappear, on one or both of the 

1 I have also, through the kindness of Prof. I^ampert, been enabled to examine the type specimen 
of the genus Hyperplesthcnus, Kuwert, which is preserved in the Stuttgart Museum. Without reference 
to this specimen I could not have determined the characteristics of the genus. 


Mt'inoii's 0/ tlic I ml in It Miiatiiiii. 

[Vol. Ill, 

'iiandibles. In the case of the Aceraiinae it was useless to introduce any reference to 
this character into the key to the genera, as these were found to be well defined and 
to form a linear series of increasing specialization.' But in the Gnaphalocneminae it 
is more useful; for the genera, besides being less sharply defined by other charac- 
ters, form a branching series with one or more genera in which the dentition is 
reduced at the end of each branch; and the nature of the reduction , which is the 


\°o \ \ 

o \ \ / 

\%' \ ^. i^^/teyt/e-ifftcnai^ 


\ /* / // 

\ I / / ^^ 


I ^ 

I ? 

\ I "a. 

same in all genera in which it occurs in any one branch, is difierent in different 

Zang has attached great importance to the fusion of the (anterior) intermediate 
and lateral areas of the metasternum in certain genera of Gnaphalocneminae, and here 
we have another character, usually very sharply defined, which is most useful for the 
determination of these genera. But it does not appear to me to be of such primary 
importance as the sculpturing of the mentum and of the anterior margin of the head ; 
and Zang's " Kmipiolus Group," which includes all genera so characterised, does not 

' I have had to modify my views ou these points, see Appendix I, p. 316. 

igi4.] F. H. Gravely : An Account of the Oriental Passalidae. igg 

appear to me to be much more natural than a group containing all the genera 
with reduced dentition would be. In the accompanying diagrammatic representa- 
tion of the grouping of the genera of Gnaphalocneminae that appears to me 
to be most natural (see previous page), the names of the genera so characterised 
are marked with an asterisk ; and it will be seen that they appear in two different 

When thus arranged the genera fall into six groups (each enclosed, in the diagram, 
by a dotted hue), and it will be convenient to refer to these groups respectively as 
the Kaupioloides, Protomocoelus , Hyperplesthenus , Gnaphalocnemis , Plesthenus and 
Gonatas groups. The diagram indicates only the general course of the evolution of 
asymmetrical forms with reduced dentition, from symmetrical ones with complete 
dentition, as this is illustrated by the genera now in existence. Many of the genera 
in the lower parts of the diagram are distinguished from those above, as well as from 
those below them, by characteristics pecuHar to themselves ; and the diagram must 
not be taken to imply the actual descent of one genus from another now existing, but 
only from one the general characteristics of whose head and mandibles were very 

The Kaupioloides group appears to be the most primitive, and as its members 
closely resemble those of the subfamily Macrolininae, it may be supposed that the 
two originated from a common stock, and have not departed very widely therefrom 
in general appearance. The Protomocoelus group, in spite of its possession of 
primary scars, appears to be related more closely than any other to the sub-family 
Aceraiinae. The affinities of the tWo remaining subfamilies of the second section 
of Oriental Passalidae, the Pleurariinae and Leptaulacinae, are more obscure ; but 
it seems likely that the former is related to the Aceraiinae. 
The genera of Gnaphalocneminae may be defined thus : — 

Primary scars almost always present^ ; when absent, area between 

anterior part of left supra orbital ridge, and ridge joining left inner 

and outer tubercles, deeply excavate, the excavation bounded 

behind by a sharply defined ridge ; dentition of right mandible never 

more reduced than that of left . . . . . . • • • • . . 2. 

Primary scars never present ; surface of head wit'iout deep or 

sharply defined anterior depressions, or dentition of right mandible 

, more reduced than that of left . . . . . . • • ■ • . . 9. 

' I follow Zang (1905&, p. 227) in supposing the lateral and intermediate areas of the metasternum 
to be fused in the genus Pt'/o/)f£/.'s ; but Zang himself had not seen a specimen (igoSir, p. 316). See 
also following footnote. 

* It is, for instance, very possible that Protomocoelus and Pdopitks have sprung more directly thai: 
is here indicated from KaiipioUdes, which they resemble in the structure of the metasternutu more th.i ' 
they do AnalacJies-Cetejus. 

3 Perhaps always; but in the most highly specialized forms they are often very small and obscare. 
In all the more highly specialized forms, and also in some les5 highly specialized, the excavation behind 
the anterior margin of the head is present on the left and often on both sides; and in these forms it 
supplies an excellent mark of recognition; it faces more upwards and less forwards, and when well 
developed is much deeper than the somewhat similar hollow often found in the genus Gnapltalocnemis, 

Memoirs of the Indian Museum. 

(Outer tubercles at least as far from one another as from anterior 
angles of head ; never asymmetrical, except when lateral and inter- 
mediate areas of metasternum are fusel ; surface of head usually 
rough ; dentition always complete 
^' ' Outer tubercles at most as far from one another as from anterior 
angles of heal ; often asymmetrical, always so when lateral and inter- 
mediate areas of metasternum are fused ; surface of head smoother, 
polished ; dentition complete or reduced 
i Anterior intermediate and lateral areas of metasternum distinct 
3. from one another 
\ Anterior intermediate and lateral areas of metasternum fused 
r Secondary scars feebly developed (sometimes absent), transverse, 
j never extending back as much as half way to posterior margin of 

Imentum . . 
Secondary scars distinct, variable 

Secondary scars represented by a pair of very small grooves, meet- 
ing in an acute angle . . . . . . . . 

5.^ Secondary scars represented by a single long transverse crescentic 
or semicircular groove, crossing middle-line about halfway between 
anterior and posterior margins of mentuni . . 

I Anterior intermediate and lateral areas of metasternum distinct 
from one another* ; dentition complete, normal 
Anterior intermediate and lateral areas of metasternum fu-ed ; 
dentition of mandibles reduced on both sides, but especially on 
the left, anterior lower tooth (when present) widely separated 
from middle lower tooth, and partially fused with lowest terminal 

[Vol. Ill, 

{Kciiif^ioloidcs Group) 3. 

(Protomococlus Group) 

[ Kiuipioloides, Gravely ']. 



[Maslochilus, Kaup]. 

[Phiirnchilus, Kuwert ']. 

in which genus moreover, the dentition of the right mandible is always much more reduced than that 
of the left. 

' These two groups are not very sharply separated. The asymmetrical species of the J'ro/niiiocochis 
group can easily be recognized by the form of the anterior margin of the head (see Kuwert, 1S96, pi. vii, 
figs. 70, 71, 77 and 82). Some of the symmetrical forms, however, come very near certain members of 
the Kaupioloides group, but are smaller, with the upper surface of the head smoother and the middle 
part of its anterior margin usually somewhat more prominent and never very broad. The genus 
Kaupioloidcs. seems to be almost exactly intermediate between the two groups in the=e characters, but, 
being itself very slightly asymmetrical, it is distinguished from all their symmetrical forms and many of 
their asymmetrical ones by the fusion of the lateral and intermediate areas of its metasternum and from 
the rest of their asymmetrical ones by the complete dentition of its mandibles. The lateral and inter- 
mediate areas of the metasternum are also fused in all genera of the Ilypcrplcsthcuus group; in this 
respect the genus Kaupioloidcs connects the archaic group in which I have placed it with this group, 
which must in any case be supposed to have originated from some such archaic forms ; and in the 
structure of the head it connects it with the members of the Protomococlus group. 

^ Mitt. Naturhist. Mus. Hamburg XXX, 1913, p. 103. 

■" Concerning the genera Episphenoidcs and Pharochiliis see also Heller, 1910, p. 17. The general 
sliape of the scars of the ftjrmer is correctly shown in Kuwert's figure, but they appear somewhat 
too sharply defined and at too great a distance from the anterior margin of the mentum ; so that the 
effect is that of the form of mentum found in certain species of the genus Maslochiliis, rather than 
of that characteristic of the genus Episphenoidcs. 

* The dividing line is less pronounced than usual in Auulnchcs schciikliiigi, Heller, but it is there; 


F. H. Gravely : An Account of the Oriental Passalidae. 


Upper margin of lett mandible, between upper tooth and base, 
straight or lightly curved so as to contain not less than 120° ; ridge 
joining inner tubercles usually straight and not strongly crested ; 
prouotum usually very strongly punctured in anterior angles ; 
posterior angles of lower side of prothorax usually not very hairy.. [Cetejus, Kaup]. 
Upper margin of left mandible, between upper tooth and base, 
abruptly bent making a distinct angle of at most 120° ; ridge join- 
ing inner tubercles usually concave and sharply crested, the crest 
projecting forwards ; anterior angles of pronotum almost or quite 
unpunctured; posterior angles of lower side of prothorax usually 
very hairy .. .. .. ., .. [Analachcs. Kuwert]. 

Left outer tubercle broader, but little longer than right; when a 
ridge is present between left inner tubercle and supra-orbital ridge, 
it is directed towards anterior end of latter . . . . [Pclopides, Kuwert]. 

Left outer tubercle much longer than right ; a ridge extending be- 
hind deep excavation from left inner tubercle towards middle of 
supra-orbital ridge . 

[Protomocoehis,Zang (■■ 
Pelops, Kaup)]. 

Lamellae of antennae usually all rather short and stout, never more 
than four very long and slender; mandibles normal or with denti- 
tion reduced on right side ; secondary scars almost always longitu- 
dinally linear and widely separated 

All sLx lamellae of antennae long and rather slender ; mandibles 
sometimes normal, more often with anterior lower tooth absent or 
fused at base with lowest terminal tooth, especially on left side ; 
secondary scars semi-circular or V-sUaped, more or less transverse, 
often more or less distinctly continuous with one another across 
middle line 

Lateral and intermediate areas of metasternum fused ; mandibles 
normal on both sides . . 

Lateral and intermediate areas of metasternum distinct ; anterior 
lower tooth of right mandible smaller than that of left, usually 

(Gonatas Group) 17. 

(Hyperplestlicn f/s Group) 

and the dentition is clearly complete and normal, though the anterior lower and lowest terminal teeth 
are broken on the right side in the type. 

' See Heller, 1910, p. 14. These two genera cannot be sharply separated from one another, and 
should perhaps be regarded as sub-genera only. 

* Kuwert separates these two genera thus, but his figure scarcely bears out his statement that the 
left outer tubercle is much longer than the right in Protoinocoelus ; and the material before me seems to 
indicate that his figure is better than his description. I am inclined to think that Pelopides will ulti- 
mately have to fall as a synonym either of Analacli's or of Pyotomococliis ; but which cannot be deter- 
mined before the mandibles and metasternum are described after examination and not, as in the above 
key, by inference. vSo I prefer to retain the name for the present. Zang (19056, p. 227) who had also 
not seen a specimen (1905a, p. 316) regards P. schraderi, Kuwert, from New Guinea, as the type of the 
genus, which he considers distinct ; and suggests that the remaining species, P. gravidas, Kuwert, from 
Mindanao, ought probably to be transferred to some genus of the Gnaphalocnemis group on account of 
its toothed labrum. 

Memoirs of the Indian Museum. 

[Vol. Ill, 


/ Central tubercle of head free, directed forwards 

I Central tubercle of head normal 

( Secondary scars widely separated 

I Secondary scars not widely separated, directed inwards so that 

I their posterior ends nearly meet in luiddle-line without reaching 

. posterior margin of mentum ; antennae normal ; left outer tubercle 

truncate or complex, right similar or simple and bluntly pointed ; 

pronotal scars with or without hair 

Antennae normal ; outer tubercles variable, simple or complex, not 
markedly asymmetrical ^ ; pronotal scars hairless 

i Antennae with lamellae of last four joints abnormally long and 
slender, left outer tub;rcle broadly truncate, right more or 
pointed but no longer than left ; pronotal scars hairy 
TOuter tubercles simple, very variable, acute or truncate, highly 
I asymmetrical (that of the right side being always the larger) or 
I almost ( ? sometimes quite) symmetrical 

At least one of the outer tubercles comple.K, or truncate with an ad- 
ditional tubercle between itself and anterior angle of same side of 

'Secondary scars united in middle-line bahind, together forming an 
ou-shaped depression much as in Omegarius ; anterior lower tooth 
of right mandible small but distinct 

[Aiireliiis, Kuwcrt 

Secondary scars completely separated as a rule, anterior lower tooth 
V^of right mandible absent 
Outer tubercles equal ' 

i6./ Outer tubercles unequal, that of right side always larger than that 
of left 

[Hypt-rplcsl/toi IIS, Ku- 

[Kaiipiolus, Zang* 

(= I'ellejiis, Kaup)j. 

[Ldhicniis, Kaup]. 

[Plestheniis, Kaup ( = £")i 
hryiilciis, Zang ■)]. 

{Giiiiphalocncmis Grou])) 

Parapelopidcs, Zang ; 
pp. 24() & 297. 


Tnipczuchiliis, Zang; 

pp. 247 & 297. 

Gnaphalooiemis,' Heller ; 
pp. 248 & 298. 

' The genus Aurelins, specimens of which I have myself examined, possesses all the characteristics 
of the Hy pcrplesthenus group. 

^ The genus Hyper plestheniis does not appear in Kuwert's table for the determination of genera 
(1896), being described, apparently for the first time, in 1898 (p. 318) in the description of H. impar, 
Kuwcrt, without any separate generic definition. An examination of the type specimen has enabled 
me to define the genus more precisely, and to find that Lrt/j!'c»;/s gracilis, Heller (Abh. Mas. Dresden 
xiii, 3, p. 16), of which I have also seen the type, is really a Hypcrplesthenus. See Mitt. Naturhist. 
Mus. Hamburg XXX, 1913, p. 105. 

« Kuwert says always asymmetrical, but they are symmetrical in his figure and in the majority of 
specimens before me. 

* Kaupiohis irigonophoriis, Zang, as Zang himself suspected, is not a true Kjiipioliis, and I have 
made it the type of a new genus Kaiipioloidcs (see above, p. 200). 

' Zang considers the name Plestheniis to be preoccupied by Pleslhenes, Stal. 

' Not alway.^^ quite symmetrical in structure however : so it is possible that the genus Trapezochilits 
will eventually have to be merged in Gnaphalocnemis. 

"^ Kuwert I189G, p. 229) states that there is always a depression between the left outer tubercle 

IQI4-] F- H. Gravely : .In Account of the Oriental Passalidae. 203 

! Secondary scars curved 
Secondary scars V"Shaped, forming together a VV'Shaped depres- 

Guiuitas, Kaup, pp. 250 
& 299. 

/'Secondary scars appro>:iniate, forming together an <jj shaped de- 
i8.<^ pression .. .. .. .. .. .. [Oniegarius, KnwertK] 

(^Secondary scars widely separated .. .. .. [Tatitis, Kuwext'.] 

The last Oriental subfamily, Leptaulacinae, contains only two of Kuwert's 
Oriental genera, these being Leptaulax and Trichostigmus. Of these the former is by 
far the larger, and Zang (1905a, p. 106) has split it up into two. The species belonging 
to it seem capable of arrangement in quite a definite linear series. At one end of this 
series we have forms in which the intermediate areas of the metasternum, and as a rule 
the sides of the pronotum, are thickly and extensively punctured, the lateral grooves of 
the elytra being very broad with transversely linear punctures ; while at the other end 
come forms in which the lateral grooves of the elytra are narrow with small round punc- 
tures, the sides of the pronotum bearing a few punctures only, while the punctures on 
the intermediate areas of the metasternum are absent, or obsolete and confined to the 
inner posterior angles. All species belonging to the first-mentioned end of this series 
apparently belong to the genus Leptaulax as restricted by Zang, and those belonging 
to the last-mentioned apparently belong to his new genus Leptaidacides ; but as the 
character on which these genera are separated — the form of the parietal ridges of 
the head— is only once referred to by Kuwert, it is impossible to be quite certain 
whereabouts in the series the restricted genus Leptaulax ends and the new genus 
Leptaulacides begins. I have, therefore, recagiiized the genera as distinct, only when 
describing tlie species before me,^ and have dealt with both under the old name Leptaulax 
(as defined by Kaup and Kuwert) in the subsequent synonymmic revision of the 
Oriental Passalidae as a whole. The threa accepted genera of lycptaulacinae may be 
distinguished as follows : — 

r Sides of elytra hairless . . . . . . . . . . 2. 

I. Sides of elytra hairy .. .. Trichostigmus, Kaup, pp. 

I, 261 & 3T0. 

and supra-orbital ridge in this genus. This character is not found in any other accepted genus of the 
group but it is shared by "Pelopides" gravidas, Kuwert, which must be transferred, in all prob- 
ability, to this group (see Zang, 19053, p. 316 and 19056, p. 227). "Pelopides" gravidas differs, 
however, from all known species of Gnaphalocnemis, in that the left outer tubercle is larger and not smaller 
than the right. One species of Gnaphalocnemis is known to me in which there is no depression between 
the left outer tubercle and supra-orbital ridge. This is described below under the name G. simplex. 
It further differs from other species of Gnaphalocnemis in the form of the outer tubercles, and in 
having the anterior intermediate and lateral areas of the metasternum very imperfectly separated. A 
new genus may ultimately be required for its reception . 

' I very much doubt the distinctness of these two genera one from another. 

^ An examination of further material has shown that this distinction does not always hold good, 
for L. himalayae and certain species immediately succeeding it in my list may have parietal ridges of 
the kind found in either genus, the two kinds being souaetimes found on opposite side of a single speci- 
men. The two genera have therfore been united in the following pages. 

204 Memoirs of the Indian Museum. [\'oi.. Ill, 

/ Parietal ridges of head short and laterally truQcate, being separated 
1 from supra orbital ridge by a deep hollow ; insects strongly and 
2 I J more extensively punctured .. .. .. .. L('/)/:(/(/((.v, Kaup, pj). 251 

& 302. 
Parietal ridges long, laterally continuous with supra-orbital ridges; 
insects sparsely and less extensively punctured .. .. l.cptaitLuidcs, Zang. 


Localities are recorded as briefly as possible in this list. For further geograjjhical 
iuformatioa regarding them, and also for all information regarding the synonymy 
adopted here, the sixth part of this paper should be referred to. I have thought it best 
to retain, for the present, all names already in use which can be applied to forms satis- 
factorily distinguishable from their allies in the collection before me, even when (as 
in the case of Ephisphcnus comptoni and its varieties) there is evidence to show that 
a larger series of specimens is almost certain to prove that the characters, which now 
seem to be distinctive, are of no real taxonomic importance*. But as full species I 
have only recognized forms differing in at least one constant and definite character, 
or combination of characters, from every other form known to me. 

The descriptions refer only to the specimens before me, and the measurements 
given for each species do not always show the total range in size indicated in the 
descriptions of previous authors. 


Genus COMACUPES, Kaup. 

Comacupes cavicornis, Kaup. 

1. C. CAVICORNIS, Kaup,s. str. 
Regd. No. '"^^' '■■■'■^-'"' Sinkep Island Moti Ram. 

Description. — Length 23-25 mm. Labnim about twice as wide as long, anterior 
margin slightly concave, angles rounded, sides slightly convergent behind Upper 
toot\\ oi mandibles weak, obtuse; anterior ower tooth conical, usually truncate or 
bifid at apex, that of left side partially fused to lower margin of lamelliform middle 
lower tooth. Mentum hairy and coarsely punctured, with a strong median keel, 
which is obliquely truncate behind and extends into a slight prominence in middle of 
anterior margin. Head hairy and strongly punctured, except on supra-orbital ridges 
and anterior margin ; central tubercle narrow (usually more so in front than behind), 
pointed before and behind, hollowed out above, the upper margins of its raised sides 
straight and usually somewhat elevated behind and dipping at an angle of about 120° 
above the somewhat elongated anterior overhanging portion; anterior margin of head 
thickened, approximately straight except close to lateral sutures of frons on inner side 

' See footnote 2 on i)receding page. 

■' A list of these collections will be found above p. 178. 

■' This has since been confirmed; see p. 2S2. 

1914-] !'• H. Gravely : An Account of the Oriental Passalidae. 205 

of which it is a Httle convex; lateral sutures of frons meeting anterior margin of 
head in an angle of about 60°, and crest of supra-orbital ridge in an angle of about 
30°; crest of supra-orbital ridge meeting canthus in an angle of about 30°; external 
angle of canthus acute, usually little more than 60°; supra-orbital ridge unpunctured, 
apical angle about 120°, crest defined by a groove on inner side ; a more or less distinct 
transverse groove present between supra-orbital ridge and central tubercle. Pronotiim 
without strong punctures except in the marginal groove, which is not interrupted 
before or behind in the middle-line, and is broader on the inner side of each rounded 
and slightly prominent anterior angle than elsewhere ; sides parallel ; posterior margin 
rounded; scars curved (the concave face anterior), slightly oblique, with a single 
smaller round depression immediately in front of them. Lateral plates of lower side of 
prothorax with hair-bearing punctures behind, and a few along outer margin in front. 
Mcsothoracic plates punctured throughout with the exception of the posterior angles of 
the episterna; sternum covered with long hair; the rest of the mesothorax hairless. 
Metasternum with the central area moderately distinct, this and the posterior inter- 
mediate areas smooth and hairless; lateral and anterior intermediate areas punctured 
and more or less hairy; lateral areas much broader behind than in front. Second 
and third abdominal sterna covered with hair-bearing punctures all over, fourth to 
sixth at the sides. Anterior margin of front coxae, and posterior part of hind 
coxae, punctured. Tibiae of middle and hind legs each armed about two-thirds of 
the way down outer side with one spine. Elytra somewhat punctured and very 
hairy (in perfect specimens) at the shoulders; all the grooves strongly punctured; 
the rest smooth. 

2. C. CAvicoRNis, var. borneensis, n. var. 

' PI. xi, figs, i-ifl. 

Regd. No. ■'Jl'" Kuching, Sarawak Sarawak Museum. 

I have examined three specimens of this form. Of these one has been kept 
in the Indian Museum, one has been returned to Sarawak, and one (the type) has 
gone to the British Museum. 

Description. — Length 26'5-28'5 mm. Differs from C. cavicornis, s. str ,, only in 
that the head is smooth as in the following variety and has all its angles more or less 
produced, while the puncturing of the metasternum is intermediate in extent between 
that found in C. cavicornis, s. str.. and that found in the following variety. 


PI. xi, figs. 2-2a. 
Regd. No. '■^" Malacca ? 

1 1 1 

9-t23-S 9430 9432 8 9437^ | 

1 1 1 1 I 

94.43-5 9480 6381-2 ' 

I Stoliczka bequest, 
( J. Wood-Mason, etc. 

J. Wood-Mason. 

2o6 Memoirs of the Indian Museum. [A'oi,. Ill, 

Dcscripiion. — Length 2^-27 mm. Differs from C. cavicornis, s. str., only in having 
the upper surface of the head entirely smooth, hairless, and unpunctured except right 
at the back behind the supra-orbital ridges ; in having the central tubercle of the 
head shorter and more abruptly truncate in front; in having somewhat wider enlarge- 
ments of the anterior part of the marginal furrow of the pronotum ; and in having 
the lateral and anterior intermediate areas of the metasternum more sparsely and less 
extensively punctured behind. 

Comacupes stoliczkae, n. sp. 

PI. xi, figs. ^-^a. 
Regil. Nu. "I^ '•'*'■' (/y/)c) Johore, Malay Peninsula J. Wood-Ma.son, etc. 

-^ Sinkep Island Moti Ram. 

This species is intermediate between C. cavicornis and C. niasoni \ but it closely 
resembles C. cylindraceiis also, agreeing with it in every detail referred to by Kaup 
in his monograph. Kaup notes, however, that he includes under the one name cylind- 
raceus two forms differing considerably in the shape of the central tubercle, both of 
which forms he figures; and I have little doubt that his " 2nd form" was in reality 
the present species. Perty's original description and figure of C. cyhndraceus would 
perhaps do almost equally well for either form ; but I think that these, and also those 
subsequently given by Percheron, apply best to the form referred to by Kaup as the 
typical one. As the present form differs considerably from this "typical" C. 
cylindraceus in the shape of the central tubercle, I have no hesitation in regarding 
it as a distinct species ; and I name it after Dr. Stoliczka in recognition of his valu- 
able work on the Oriental species of Passalidae. 

Description. — Length 25-27 mm. Labrum often nearly as long as it is wide, 
punctured, hairy, anterior margin approximately straight, angles strongly rounded, 
sides straight and parallel. Mandibles as in C. cavicornis, the upper tooth even 
more reduced. Head above and below also as in C. cavicornis in general shape, but 
with the central tubercle more elevated (especially in front), much narrower, thickened 
behind the apex, and simply rounded, not excavate, above; there is, moreover, 
a pair of transverse ridges at the base of the central tubercle much as in Taeniocerus 
bicanthatus, but more pronounced and covered like the rest of the head with a 
coarse punctuation slightly finer and closer than that of Comacupes cavicornis. 
Prothorax as in C. cavicornis, but with somewhat smaller scars, and still more hairy 
sternum. Mesothorax also as in C. cavicornis. Metasternum differing from that of 
C. cavicornis in that the punctuation is somewhat finer and closer, and extends 
over the whole surface outside the central area. Abdominal sterna (? always) less 
hairy than in that species. Posterior part of hind coxae punctured ; middle tibiae 
armed with one spine about two-thirds of the way down ; hind tibiae similarly 
armed, but with the spine usually very slightly smaller. Elytra sometimes with 
some small punctures along the lower side of the anterior half of the tenth rib ; 
otherwise as in C. cavicornis. 

I9I4-] F. H. Gravely ; An Accoimt of the Oriental Passalidae. 207 

Comacupes masoni, Stoliczka. 
PL xi, figs 4-4^. 

Regd. No. '^' {type') Johore, Malay Peninsula J. Wood-Mason. 

Description. — Length 30 mm. Labrum as in C. stoliczkae, but with anterior 
margin slightly concave. Mandibles also as in C. stoliczkae. Mentum perhaps differ- 
ing from that of that species in that the median keel is scarcely as high. Head 
somewhat more finely punctured, otherwise similar; central tubercle quite as 
prominent, not depressed at tip as in both C. stoliczkae and C. cavicornis, scarcely 
excavate along middle-line, outline in side view straight (perhaps very faintly 
convex) and horizontal above, oblique beneath. Pronotum with anterior angles a 
little more prominent, sides slightly divergent behind ; enlarged areas of anterior 
marginal groove rather abruptly terminated at their outer ends. Prothrorax ventrally 
as in C. stoliczkae. Mesothorax, metastenium , elytra and legs also as in C. stoliczkae. 
Second abdominal sternum strongly punctured throughout ; third with a few punc- 
tures at each end; rest of lower surface of abdomen smooth and highly polished. 

Comacupes cylindraceus (Perty). 
Pi. xi, figs. 5-6(? ; text- fig. 2. 

Regd. No. '^ '^ Sinkep Island Moti Ram. 

-^ Klingkang, Borneo Sarawak Museum. 

Also specimens from the Larut Hills in the British Museum, from Deli (Sumatra) 
in the Hamburg Museum, and from Padang and St. Rambe (Sumatra) in the 
Deutsches Entomologishes Museum. 

Under this name, as explained above (p. 206), I include, of the two forms 
included under it by Kaup, only the " typical " one. But I 
also include everything referred by Zang to C. masoni and 
C. cormocerits ; and the series of specimens before me shows 
the species to be so variable that I am inclined to think it 

possible that C. masoni, Stoliczka, may ultimately have to 
Text figuer 2. , ^ 1 .^ -.l 1 

„ J , , .be referred to it also. 

Head of a form ot ^.„ . ^, ,. / 

C. cylindraceus, from Description. — Length 24-26 mm. Diflers from C . stoliczkae 

the left side, x 4. ^^^ (- j^^soni only in the form of the central tubercle of the 

head ; this is somewhat less prominent in front as seen from 
the side, and more or less vertically truncate; its anterior margin may be straight 
or markedly concave, and either the upper or the lower angle may project some- 
what — but never very much — beyond the other (see pi. xi, figs. 5^7 and 6a, and text- 

' This specimen appears to have been entered in our register originally under the No. " ,-' , to 
have lost the label bearing this number, and then to have been entered again under another number. 

2o8 Memoirs of the Indian Museum. [\'oi.. Ill, 

fig. 2'. The form of the anterior parts of the margmal groove of the pronofum 
and the puncturing of the ahdomiiud slriiui are both extremely variable. 

Taeniocerus bicanthatus (Percheron). 
PI. xi, figs. 7-7/;. 
Regd. No. -*-' ? J- Wood- Mason. 

9177 MBi-S «38S 

Johore, Malay Peninsula 

No. ""' is markedly smaller than any of the others, but in structure it agrees with 
them perfectly. In all the specimens I have seen the external angle of the canthus 
is more or less obtuse, scarcely rectangular as it is shown in Percheron's figure, and 
certainly not acute as in Kaup's figure. 

Description. — Length 24-29 mm. Labrum about twice as broad as long ; anterior 
angles rounded, laterally prominent; anterior margin concave. Upper tooth of 
mandibles somewhat obtuse and not very prominent ; uppermost terminal tooth 
obsolete; anterior lower tooth of left mandible wider than that of right ; the former, 
and often the latter also, more or less bifid, the upper denticle being longer than the 
lower. Mentum with a considerable angular prominence in middle of anterior 
margin; this prominence continued backwards to posterior margin as a hairless, and 
usually smooth and more or less hollowed, triangular area ; lateral portions of 
mentum flattened, hairy and coarsely punctured. Head smooth and polished as a 
whole, but punctured behind supra-orbital ridge, beside central tubercle, and 
in area enclosed by the U-shaped ridge with which this tubercle is crowned ; arms 
of this ridge parallel or slightly divergent,' their extremities somewhat promi- 
nent. Anterior margin of head slightly concave, with a slightly convex trans- 
verse groove immediately behind it, meeting crest of supra-orbital ridge at 
an angle of not more than 90°, and margin of canthus at an angle of about 120° ; 
external angle of canthus about 120°; apical angle of supra-orbital ridge somewhat 
more obtuse than this ; crest of supra-orbital ridge defined on inner side by distinct 
groove ; a pair of more or less distinct ridges extending outwards and very slightly 
forwards from base of anterior angles of central tubercle. Pronotiim smooth, 
polished ; anterior margin almost straight, anterior angles rectangular, pointed but 
not prominent ; sides parallel, posterior angles much rounded, posterior margin some- 
what convex; margmal groove punctured, incomplete in the middle-line in front, 

' The type specimen of C. cormocerus, Zang, has the central tubercle of the form shown in the 
last-mentioned figure, which is from a specimen from the Larut Hills, but the upper angle does not project 
upwards, and occupies a much smaller part of the front of the tubercle than does the lower. 

* The width of this horn is very variable in proportion to its length even in our few specimens, all 
of them ])robably from Johore. So Zang's suggestion (igSfl, p. 105) that Bornean specimens are 
distinguished ))y a broad horn such as is figured by Kaup, and Malayan ones by a longer horn, can no 
longer be maintained. 

1014.1 F- H. Gravely : An Account of the Oriental Passalidae. 209 

widest a little more than half way from this point to the angles; median groove 
strongly developed; scars obUquely S-shaped, punctured in lower two-thirds 
of their length. Lateral ventral plates of prothorax sparsely and obscurely punc- 
tured in front, closely and distinctly punctured and hairy behind. Scutellum more or 
less sparsely punctured in antero-lateral angles ; oblique punctured band of mesotho- 
racic episterna very broad, covering almost the whole of the plate, posterior angles 
polished ; anterior parts of lateral angles of mesosternum punctured, the rest smooth, 
middle-line marked by a broad and somewhat Y-shaped groove whose fork is 
directed forwards. Metasternum unpunctured except in anterior angles, lateral 
areas not very broad behind. Abdominal sterna smooth and polished, unpunctured. 
Posterior coxae matt behind, not distinctly punctured. Tibiae of middle legs with one 
spur situated externally slightly beyond the middle; those of hind legs without these 
spurs. Elytra unpunctured except in the grooves. 

Taeniocerus pygma2;us, Kaup. 
PI. xi, figs. 8-8(?. 
Regd. No. "^ "Malacca" ? 

I have also examined two specimens from Kuching in the Sarawak Museum 

Description.— Length 15-16 mm. Labritm twice as broad as long behind, but 
three times as broad as long in front, the rounded anterior angles being much more 
prominent than in the preceding 'species. Upper tooth and uppermost terminal 
tooth of mandibles obsolete; anterior lower tooth conical, that of left side united by 
a ridge to middle lower tooth. Smooth central part of mentnm broader in front than 
in preceding species, being distinctly pentagonal; mentum otherwise as in that 
species. Head more or less finely roughened, except around central tubercle where 
it is smooth, and further back where it is punctured; central tubercle almost 
completely divided by a longitudinal groove into a pair of conical crests apically 
rectangular in profile; these crests obscurely joined together behind into a single 
horse-shoe shaped structure. Anterior margin of head slightly concave, sometimes 
bordered by an obscure marginal furrow, meeting supra-orbital ridge in a much 
rounded angle of a-bout 60° and side margin of the head at about 90° ; side margin 
of head making a somewhat rounded re-entrant angle of about 120° with canthus; 
external angle of canthus and apical angle of supra-orbital ridge also about 
120°, the latter situated at about the middle of the ridge. A slight hollow present 
on each side of central tubercle. Pronotnm polished, anterior niargin wavy or almost 
straight, anterior angles acute and often very slightly prominent ; sides convex, diver- 
gent; posterior angles rounded, posterior margin convex, marginal groove well deve- 
loped across middle-hne in front, otherwise as in preceding species ; median groove 
strongly developed; scars /- or y-shaped, punctured. Prothorax ventrally as in the 
preceding species. Scutellum entirely smooth except for usual finely punctured 
patch in middle of anterior margin, or faintly grooved in middle-line immediately 

2IO Memoir<i 0/ the Indian ^fllseum. [Vol. Ill, 

behind this patch ; mcsothoracic episterna crossed by oblique band of punctures or 
with their upper angles punctured throughout; mesosternum finely striatopunctate at 
sides in extreme front, otherwise smooth and polished, sometimes more or less clearly 
grooved or keeled in the middle behind; metasternum as in the preceding species 
except that an obscure median longitudinal keel is present and the lateral areas are 
even smoother. Abdominal sterna and hind coxae polished, unpunctured. Tibiae of 
middle legs with one blunt tooth slightly below the middle; those of hind legs 
without an)- such tooth. Lateral grooves of elytra distinctly punctured. 

Taeniocerus bicuspis, Kau]). 
PI. xi, figs. 9-9(/. 

Regd. No. 

Stoliczka bequest, Asiatic 
Society, etc. 

Gopaldhara, Rungboiig Valley, Dar- H. Stevens (coll. \V. K. 
jeeling District Webb). 

Choiigtang, Darjeeling District J. Wood-Mason. 

Upper Tista, 4000 ft. ? 

vSurei!, Darjeeling District A. Alcock. 

N. E. Frontier ? 

Dafla Expedition 1 

Dafla, Camp q xx t^ ^< , • « ^ 

' ^ ^ ^ H. H. Godwin- Austen. 

Dikrang Valley I 

Harmutti, base of Dafla Hills j 

Kobo, Abor Country, 400 ft. 1 

Rotung, Abor Country, 1400 ft. | Nov. 1911— Feb. 1912. 

Upper Rotung, Abor Country, 2000 ft. ? S. W. Kemp. 

Upper Renging, Abor Country I 

Cachar j. Wood- Mason. 

Mr. Stevens' collection includes a specimen from .Silonibari at the l)ase of the 
North Lakhimpur Hills in Assam. 

Description. — Length 20-22 mm. Labriim as in T. bicanthatiis. Mandibles as in T. 
Pygmaeus but with anterior lower tooth slightly more compressed. Mentum not unlike 
that of T. pygmaeus, but with median prominence of anterior margin almost absent 
or entirely so, the central area being conseqtiently quadrangular; distinctness of central 
area from adjoining parts somewhat variable as (in specimens from the Darjeeling 
District especially) it is apt to be punctured, and in extreme cases (e.g. "","' and '""') 
the whole surface of the mentum appears uniformly punctured and hairy. Head 
smooth, with a transverse patch of strong punctures at the back on each side behind 
supra-orV>ital ridges. Anterior margin bordered by a strong groove, slightly convex in 
the middle and slightly concave on each side ; meeting crest of supra-orbital ridge 








:U 29-30 



841 2. -. 

igi^.] F. H. Gravely : An Account of the Oriental Passalidae. 211 

in an angle of 90° and canthus in an angle of about 120°; external angle of canthus 
about 90° (sometimes, especially in Cachar specimens, this angle is reduced by 
an intensification of the concavity in the anterior margin of the canthus) ; apical angle 
of supra-orbital ridge about 100°, situated above and a little in front of anterior 
margin of eye ; crest of supra-orbital ridge defined on inner side by a strong groove ; 
central tubercle distinct, divided by a longitudinal concavity above into a pair of 
parallel or slightly divergent crests whose front margins are vertical and straight or 
slightly convex, their upper ones meeting these at an angle of about 90° (usually 
somewhat less) and being straight or more frequently slightly concave ; hollows at 
sides of the central tubercle somewhat as in T. bicanthatus but more extended. 
Prothorax as in T. pygmaeus, but with anterior angles of pronotum a little more 
prominent and sides approximately parallel. Sciitellum without (cr with very few) 
punctures other than those usually found on anterior margin ; mesothoracic episternum 
with broad band of punctures extending obliquely across it; mesosternum more or 
less smooth, punctured close to lateral margin near the front, usually more or less 
obscurely grooved or keeled in the middle-line behind, the extent of all these mark- 
ings being apparently variable and in no way characteristic of specimens from parti- 
cular localities. Metasternum as in T. bicanthatus. Second and third abdominal 
sterna finely roughened throughout. Posterior coxae finely roughened behiad ; middle 
tibiae armed with one more or less strongly developed tooth or spine; hind tibiae with 
a very rudimentary tooth or unarmed. Elytra with all grooves punctured, the ribs 

Habits. — Mr. Kemp informs me that this species lives under the bark of moder- 
ately hard logs in very wet jungle. 


Aulacocyclus andrewesi, n. sp. 

PI. xi, figs lo-ioa. 

Described from a single specimen, collected by Mr. H. L. Andrewes in June from 
rotten timber in the Anamalai Hills at an altitude of 3500-4000 feet ; now in the 
collection of Mr. H. E. Andrewes. 

Description.— Length 23 mm. Labrum about twice as broad as long, anterior 
margin strongly concave, sides convergent behind, angles rounded and prominent. 
Right mandible missing, the left with upper tooth small but distinct, uppermost of 
three terminal teeth still smaller, remaining two terminal teeth larger, anterior lower 
tooth somewhat L-shaped in section, but fused with middle lower tooth to form with 

it a L l-shaped structure with a minute denticle on the connecting ridge. Mentiim 

punctured and hairy, with a smooth anterior prominence in middle which is medially 
grooved in front. Antennae with first two lamelliform lobes about four times as long 
as broad. Head above smooth and polished; anterior margin smooth above and 
hairy beneath, strongly concave in middle, slightly convex and very finely granular 
laterally, bordered by a strongly marked groove, meeting crest of supra- orbital ridge 

2J-2 Memoirs of the Indian Museum. [Vol. Ill, 

at an angle of about 120°, and canthus in a straight line ; external angle of canthus 
about 120"; apical angle of supra-orbital ridge also about 120°, but somewhat 
rounded; central tubercle with proximal vertical part about twice as thick as distal 
horizontal part when seen from the side, but only about two-thirds as thick when 
seen from above, extremity concave. Pionotnm polished with small deep circular 
scars; anterior margin slightly undulating, distinctly concave as a whole; sides 
parallel ; marginal groove complete, rather indistinctly punctured in front ; median 
groove complete and deep. Prothorax beneath much as in the following species. 
"scutellum smooth; mesothoracic episterna punctured above (rather sparsely except 
close to upper margin), smooth below and behind; mesosternum smooth and polished 
throughout. Metasternum faintly ridged in middle line, lateral areas somewhat 
broader behind than in front; lateral and anterior intermediate areas finely (but 
not very closely) punctured and hairy, the rest smooth and polished. Second 
abdominal sternum somewhat rugose between posterior coxae and along middle of 
extreme posterior margin ; succeeding abdominal sterna polished throughout. Elytra 
unpunctured except in the grooves. Tibiae of second legs armed with a small spine 
on the outer side about two-thirds down, those of the third pair with this spine 

Genus CERACUPES, Kaup. 

Ceracupes fronticornis (Westwood) . 

PI. xi, fig 12. 

Regd. No. '",^" Gopaldhara, Rungbong Valley. Uaijediug District. H. Stevens (coll. \V. K. Webb). 

Description.— Length 225 mm. Differs from C". aiisteni in the following respects 

only. Anterior margin of mentum not prominent in middle line, without marginal 

groove. Central tubercle of head broader above, broadly bifid at apex ; external 

angle of canthus less acute, containing slightly more than 90°. 

Ceracupes austeni, Stoliczka. • 
PI. xi, figs ii-ii((. 

Regd. x\o. f^* ''^} I^'ifla, Camp 9 x 

"••>!m-7 1"" Dikrang Valley 

i» "< ) H. H. Godwin-Austen. 

!i|L(/y/,e)i Naga Hills 

ffiiM^ti" Mauipur / 

Description.— Length 20-25 mm. Lubriim approximately rectangular, transversely 
ridged behind a concave anterior portion which faces forwards and upwards and is 
fringed, but not itself covered, with long brown hair. Upper tooth of mandibles very 
long, straight, slender, bluntly pointed distally, flattened and rugosely striate above 
and on inner side, at least twice as long as terminal part of mandible, and often 

' This specimen seems to have been entered in our register originally under the No. '"^^ and subse- 
quently to have been entered again under a fresh number. 

1914-] F. H. Gravely : An Account of the Oriental Passalidae. 213 

three or four times as long; terminal portion of mandible curved inwards; middle 
tooth somewhat the largest, and lowest somewhat the smallest, of the three terminal 
teeth; anterior lower tooth of right mandible slenderer than that of left, both L-shaped 
in section with the short limb in front and the long one below. Mentum thick in 
front, with vertical or bilaterally excavate anterior face slightly prominent in middle 
line ; anterior margin as seen from below bordered by a distinct groove. Head above 
smooth and pohshed; central tubercle very large, directed forwards, confluent with 
anterior margin, about equal to upper teeth of mandibles in length, flattened on either 
side and above, upper margin S-shaped in profile, being convex (and usually very obtusely 
angular) behind and concave in front, long narrow postero-dorsal surface granular, 
antero-ventral surface fiat and very strongly granular except immediately below the tip 
which is smooth. Supra-orbital ridges tooth-like, apical angle 6o°-9o°, ridges not 
extending forwards as far as anterior angles of head which contain about 120° each. 
External angle of canthus prominent, rounded, somewhat less than 90°. Pronotum 
polished; anterior margin slightly sinuate in the middle; lateral angles prominent; 
sides straight, slightly divergent; posterior angles strongly rounded; posterior margin 
somewhat convex ; marginal groove punctured throughout, more or less incomplete 
in middle-line, anterior transverse portion broader than the rest ; median groove 
present, very pronounced; lateral scars usually '-shaped, but somewhat variable, 
usually punctured, a few additional punctures present a little above and in front of 
them. Lateral plates of lower surface of prothorax punctured and more or less hairy 
in front of the coxal cavities, smooth and hairless behind. Scutellum more or less 
finely and sparsely punctured in front, otherwise smooth and polished. Mesothoracic 
episterna punctured throughout. Mesosternum with strong median concavity behind, 
whose sides may be raised into ridges further forwards ; punctured in front on either 
side of middle-line and beside lateral sutures, the rest of the surface being polished 
and smooth, or slightly rough in parts. Metasternum with central area not defined 
except near posterior margin, and there but vaguely; anterior intermediate areas 
closely punctured; lateral areas rough, narrow throughout; central area and 
posterior intermediate areas smooth and polished. Posterior parts of hind coxae and 
scars of abdominal sterna (including almost the whole of the second sternum) roughened 
or indistinctly punctured. Tibiae of middle legs with 1-3 spines on the outer side 
before the apex, those of the hind legs with at most one .such spine. All furrows 
of elytra distinctly punctured, ribs unpunctured. 


Genus PLEURARIUS, Kaup. 

Pleurarius brachyphyllus, Stoliczka. 

PI. xi, figs. I3-I3«- 

fii3i7 ftis.i "i C Beddome. 

\^^'* (panilypes) ' ^ S. India Stoliczka. 

~^ ) V ? Madias Museum. 

' Here again it is evident that the original numbers have been lost at some time and replaced b / 


I'uddutottam, Ana- A 


malais, 4200 ft. | 



Ananialais, a. 5500 ft. j 



Kalyaua, Pandal, Aiiamalais, 3000 ft. 

"^' (paralypc) ' 

Nilgiri Hills 


Kiilnttnnii/lia \V Knco r.1" \\T niiatc 

Memoirs of the Indian Museum. [Vol. Ill, 

Regd. No. 

malais, 4200 ft. | I 

iw«i » , , _ ,. , In rotten wood. | 

T. Bainbrigge Fletcher. 

.Madras Museum. 

This species is represented in the collection of the Madras Museum by two 
si^ecimens from the timber forest, Cochin State; and in that of the Bombay Natural 
History Society by a specitnen from Palur presented by G. I^und. Whether the latter 
record refers to the town of that name situated in Arcot or in Chingalpat it appears 
to be the most easterly yet made. 

Description. — Length 35'5-45 mm. Labriim transverse; usually asymmetrical, the 
left angle being slightly more prominent than the right; anterior margin slightly 
concave. Mandibles with upper tooth obsolete; anterior lower tooth longitudinally 
constricted before and behind, and bifid at the tip, that of the left mandible being 
larger and that of the right mandible smaller than the tooth in front of it. Mentum 
finely punctured in posterior angles and beside posterior end of broad median groove ; 
the rest coarsely punctured. Anterior margin of head usually almost imperceptibly 
asymmetrical, the left outer tubercle being more or less distinctly longer than the 
right; apex of both these tubercles simple in profile; marginal groove only present 
between the tubercles; supra-orbital ridges strongly peaked, broadly rounded behind 
the peaks, and slightly hollowed in front of them, the outer margin of the hollow 
extending on to the slightly prominent anterior angle of each side of the head ; 
anterior angles situated about as far from outer tubercles as these are from one 
another, each containing about 120°; external angle of canthus containing about 90°; 
parietal ridges broad, extending straight outwards or slightly forwards. Pronotum 
rectangular or with sides slightly diverging behind ; median and marginal grooves in- 
complete in front, latter not quite meeting former behind; scars small and unpunc- 
tured. Lateral plates of lower surface of prothorax smooth and hairless in front and 
behind, closely punctured and very hairy between. Scutellum smooth, with a few fine 

new ones, for the numbers under which specimens seen by .Stoliczka are entered in the register are ~' 
("named by Dr. vStoliczka and Mr. J. Wood-Mason"), '"j'* and '^' (both from "Dr. Stoliczka's Type 
collection"). Of these four numbers only one has been found. No. ^ bears a locality label in 
Stoliczka's writing which probably indicates its identity with No. -"p' which was presented by vStoliczka. 
No. ",'"," bears an identification label written by Stoliczka with a record that it was presented by the 
Madras Museum, and is presumably the specimen originally numbered ^'['". And No. ",'^', which bears 
Stoliczka's identification label but no record of the donor, is presumably the same as No. ~^. This 
last specimen has been listed in the new register with the preceding specimen, and both have been 
entered as presents from the Madras Museum. 
' See footnote on previous page. 

1914-] F- H. Gravely : .4;? Account of the Oriental Passa/idae. 215 

scattered punctures; mesotlioracic episterna punctured above and along anterior 
margin; mesosternum without scars, finely and more or less extensively stria to- 
punctate and sometimes hairy in front, punctured and hairy in lateral angles, 
otherwise smooth or transversely (more rarely longitudinally) wrinkled. Metasternum 
with central area more or less well, lateral areas ill, defined; latter very broad, often 
almost touching former ; central and posterior intermediate areas smooth and polished ; 
anterior intermediate and lateral areas punctured and hairy. Anterior margin of 
posterior part of hind coxae depressed and roughened, abdominal sterna and rest of hind 
coxae smooth. Elytra hairless, innermost groove of each almost entirely smooth, 
remaining grooves strongly punctured. 

Habits.— Br. Annandale informs me that this species is probably not gregarious. 
Mr. Bainbrigge Fletcher sent me specimens apparently obtained together with 
colonies of Basilianns, but he did not get many so probably they were living singly 
or in pairs with the other species. 

Subfamily ACERAIINAE. 
For the classification of this subfamily which is adopted here see Appendix 
I (below, p. 316). 

Genus TIBERIOIDES, Gravely. 
Tiberioides kuwerti (Arrow). 
PI. xi, fig 14. 
Reed No ' ( " " Sikkim " Stoliczka bequest, J. Wood 

^ ■ ■ / 9KU CAVi 9i7r, ( 

\^i n IfT I Mason, etc. 

,, ^-~ Peshoke, Darjeeling District, rt. 3000 ft. F. Haniiyngton. 

fiiM^i iii£^ Manipur H. H. Godwin- .\usten. 

Description. — Length 36-44 mm. Labrum as long as or somewhat longer than 
broad; sides straight, parallel; anterior margin concave; angles rounded, left one 
frequently somewhat sharper and a little further removed from deepest point of 
concavity than right one. Mandibles with upper tooth very feebly developed, 
anterior lower tooth consisting of a small anterior conical portion (that of the left 
side being truncate and that of the right pointed) and a still less prominent posterior 
portion. Anterior margin of median part of msntum with slight (often almost 
imperceptible) hollow in middle; lateral parts punctured evenly all over as a rule. 
Upper surface of head roughened, central tubercle extending back as a low mound 
between, and sometimes not clearly separated from, low rounded parietal ridges 
which extend outwards and are scarcely if at all crested. Inner tubercles almost 
equidistant from each other, from outer tubercles, and from central tubercle; ridge 
joining inner tubercles to each other and to outer tubercles forming single continuous 
curve when viewed from above. Outer tubercles with two widely separated rectangular 
apices one above the other. Supra-orbital ridges very broad and smoothly rounded 
behind, not peaked, hollowed out in front so that each forms two narrower divergent 
ridges of which the outer one extends to the anterior angle of the head. Anterior 

2i6 Memoirs of the Indian Museum. [Vol. Ill, 

angles of head each of about 120°; canthus rounded at its extremity. Pronotum 
with a fine marginal groove on each side which does not extend upwards much beyond 
the angles either before or behind ; median groove obsolete or absent ; scars usually 
small, almost or entirely unpunctured, other depressions sometimes present above, both 
in front of, and behind them. Lateral pia'es of under side of prothorax as in 
preceding species. Mesotliorax polished above, but dull below; scutellum smooth; 
episternum with normal oblique band of punctures. Mesosternum with or without 
very indistinct .scars. Metasternnm with central area distinct, vaguely grooved in 
the middle line, more or less distinctly in contact with smooth, strongly defined, 
oblique ridges bounding lateral areas ; lateral and anterior intermediate areas rugosely 
punctured and hairy; posterior intermediate areas polished, more or less scarred 
by large punctures in places especially on the inner side in front Posterior margin 
of hind coxae finely, often indistinctly, punctured and hairy. Second abdominal 
sternum irregularly marked with indistinct coarse punctures; succeeding sterna 
smooth. Elytra flattened above, hairless; lateral grooves much wider than ridges 
between them, their regularly arranged punctures drawn out into transverse furrows; 
dorsal grooves normal, two or three innermost on each side unpunctured at least 
in front. 

riberioides austeni, n. sp. 
PI. xi, figs. 15-15(7. 

Regd. No. '^' '^ {type) Dafla Expedition H. H. Godwin-Austen. 

"^ Dikranc; Valley ' H.H.Godwin-Austen. 

'"i't' Kalek, Abor Country, 3S00 ft. , 29-xii-ii S. VV. Kemp. 

Description. — Length 41-47 mm. This species differs from the last in the follow- 
ing characters only. Antennae more massive. Upper tooth of mandibles more 
strongly developed, especially that of the left side. Mentum with anterior margin of 
median area evenly convex or a little prominent' (never excavate) in the middle, 
extreme anterior angles of lateral areas smooth and unpunctured. Head with inner 
tubercles larger and thicker and directed more definitely forwards; ridge joining 
them to outer turbercles also stouter, not forming so continuous a curve with ridge 
joining them to one another ; distal margin of outer tubercles of Abor specimens slightly 
convex as seen from side instead of concave. Punctures of mesothoracic episternum 
extending further into upper front angle as in Pleurarius brachyphylltis ; mesosternum 
with very distinct scars which are broad in front and taper out behind. Hind coxae 
unpunctured; second abdominal sternum somewhat smoother; elytra with grooves 
uniformly narrow and punctured both above and at the sides. 

Habits. — Mr. Kemp informs me that the two specimens of this species that he 

' This label bears also the word " Nanang's." 

* This prominence may be indistinctly marked off by a vague groove behind (No. "-^'f shows this best 
among our specimens) and such specimens form a natural transition to the aberrant " Chilomaziis" 
borcalis. Arrow (1907, pp. 467-8), from the Naga Hills. 

I9I4-] F- H. Gravely : An Account of the Oriental Passalidae. 217 

found, were living deep down in a very hard dry log on a steep hill-face that had 
been cleared by the Abors for cultivation. 

Genus EPISPHENUS, Kaup. 
Episphcnus moorei, Kaup. 
PI. xi, fig. i6-i6(7. 
Regd. No. ?51- Ceylon H. Nevill. 

A specimen in the collection of the Bombay Natural History Society is labelled 
" Nitre Cave," but the whereabouts of the cave is not recorded.' This and No. '1*^ 
are the only two specimens of this species I have seen. 

Description .—Length 31-32 mm. Labrum broader than long, anterior margin 
straight or faintly concave, very slightly oblique, sides straight or slightly curved, 
angles rounded, the left one being somewhat sharper than the right. Mandibles with 
upper tooth distinct but very small; anterior lower tooth conical, simple on both 
sides, that of the left side a little larger than that of the right. Lower anterior 
margin of central part of mentum strongly convex , whole surface elevated in front 
in middle line, but not separated by groove from surrounding surface. Head smooth 
or with extremely fine microscopic punctures, except between outer tubercles, 
where it is marked with large, but somewhat indistinct punctures. Apex of outer 
tubercles as seen from side vertically truncate, but scarcely forked ; ridges connecting 
inner and outer tubercles defined on outer side by strong groove. Central tubercle 
extending backwards between parietal ridges, which extend outwards and, like the 
supra-orlaital ridges, are more distinctly crested than in the preceding species 
though less so than in E. comptoni, apical angles of supra-orbital ridges about 120°, 
ridges hollowed in front as in preceding genus, but less distinctly so, the inner wall 
of the hollow being almost obsolete though somewhat more distinct than in the 
following genus. Anterior angles of head slightly more than 120° ; canthus rounded 
at its extremity. Pronotum smooth, polished, marginal groove fine, incomplete 
across middle-line both before and behind, median groove complete or obsolete in 
front; .scars small, linear, indistinctly punctured, nearly vertical. Prothorax beneath 
as in preceding species SciUellum smooth and polished. Mesothoracic episterna 
punctured beside anterior margin below, and beside oblique dorsal margin, these two 
patches of punctures sometimes joined by an oblique sparsely punctured band. 
Mesosternum with anterior angle finely striatopunctate and clothed with short decum- 
bent hair, except on a wider or narrower area in the middle-line which is hairless, 
smooth and polished ; this smooth median area continuous behind with a highly 
polished keel, which in the Indian Museum specimen is surmounted by a fine Y-shaped 
crest, but in the specimen belonging to the Bombay Natural History Society is 
broader, flatter, and very faintly hollowed in the middle line ; keel (in both specimens) 
bordered on either side by the anterior extremities of a pair of broad, shallow, finely 

' Presumably the specimen is from Ceylon where, Mr. Green informs me, " the Nitre Cave district 
is away beyond Dumbara and varies in altitude from about 1800 to 3000 ft." 

2i8 Memoirs of the Indian Mitsciim. [Vor,. Ill, 

roughened scars ; surface of mesosternum bordering these scars on inner side viery 
highly polished, remaining triangular portion behind and between them finely 
roughened. Metasternum with central area clearly defined, widely separated from 
lateral areas, which are almost parallel-sided and rather narrow; lateral and anterior 
intermediate areas closely punctured and hairy ; posterior intermediate areas divided 
diagonally into an inner posterior very coarsely punctured part and an outer anterior 
smooth or very sparsely and more finely punctured part, hairless except along the 
closely but more finely punctured posterior margin. Posterior parts of hind coxae 
and scars of abdominal sterna closely and finely punctured. Elytra hairless, entirely 
smooth with exception of grooves, which are all of equal width and of which the outer 
ones are clearly punctured. 

Episphenus pearsoni, n. sp. 
PI. xi, fig 17. 

Regd. No. ™^ (type) Bulutota,' Ceylon (May 1907) Coloniho Museum. 

I have also examined two specimens from the same locality which remain in 
the Colombo Museum collection. 

Description. — Length 3i"5-33 mm. Differs from E. moorei in the following 
respects only : profile of outer tubercles rounded at apex ; ridges on head between 
inner and outer tubercles very fine, not bordered by definite grooves, while a pair 
of transverse grooves lying between the inner tubercles and the more distinctly 
peaked angles of the supra-orbital ridges are very distinct; lateral areas of metaster- 
num distinctly broader behind than in front ; outer part of posterior intermediate 
areas less sparsely punctured ; posterior parts of hind coxae and scars of abdominal 
sterna scarcely punctured. 

Episphenus comptoni, Kuwert. 

PI. xi, figs. i8-i9«. 

I. E. COMPTONI, Kuwert, s. .str. 

Regd. No. '— Hakgala,-^ Cent. Prov., Ceylon R. E. Green. 

■'^■'^ Ceylon British Museum. 

The Bombay Natural History vSociety's collection includes a sj^ecimen from 
Haputale, Ceylon. 

Description .^Length 39-42 mm. Anterior margin of labriim strongly concave, 
sides parallel or slightly convergent behind, angles round, left one usually somewhat 
more prominent and sharper than right. Mandibles with upper tooth obsolete; 
anterior lower tooth much reduced, consisting of a small conical projection on left 

I Mr. Green has obtained for me from the Surveyor-General the following information as to the 
whereabouts of this locality: "Bulutota is situated on the I'elmadulla Road. 10 miles beyond the 
Rakwana Rest House in Kolonna Koralc, Ratnapura District, Province of Sabaragarauava. The 
altitude is not recorded, but is I believe about 4000 ft." 

' Mr. Green informs me that the altitude of the Hakgala ]3otanical Gardens is 5500 ft. 

JQJ4.1 F. H. Gravely : A )i Account of the Oriental Passalidac. 219 

side and a larger blunter projection on right, from each of which a low ridge extends 
backwards toward middle lower tooth, this ridge elevated in about the middle of its 
length to form an additional very blunt tooth on left side but not on right. Mentum 
as described in definition of Chilomazus (p. 196). Head polished, punctured in front of 
and behind parietal ridges. Left outer tubercle slightly more prominent than right ; 
profile of former obliquely, of latter almost vertically truncate at apex; ridge 
connecting outer and inner tubercles not defined by a groove parallel to it. Parietal 
ridges sharply crested, their crests more or less distinctly continued on to sides of 
central horn; supra- orbital ridges also very sharply crested, crest continued along 
outer Hmb of anterior fork which descends abruptly at angle of somewhat over 120°, 
inner limb of fork rudimentary or absent. Anterior angles of head approximating 
to 120°, but not altogether constant; canthus rounded at its extremity. Frontal 
area somewhat variable in form, the frontal ridges being to a variable extent 
oblique and S-shaped, while the transverse ridge between the inner tubercles is either 
concave, straight or prominent in the middle. Prothovax as in Episphenus moorei, 
but frequently with a group of strong punctures at upper end of scar on each side 
of pronotum; lateral plates of lower surface not quite so perfectly smooth in front. 
Scutellum indistinctly punctured in anterior angles, often obscurely grooved in middle 
line in front, posterior angle smooth. Mesothoracic episterna punctured above and 
along extreme anterior margin ; below smooth, or obscurely punctured in posterior 
angles. Scars of mesosternum dull, the rest of this plate more or less polished, and as 
a rule finely punctured either in lateral angles, in middle or throughout ; other vari- 
able markings sometimes present. Metasternum as in Episphenus pearsoni, but with 
lateral areas as a rule still further' enlarged behind. Hind coxae and scars oi abdo- 
minal sterna almost smooth. Grooves of elytra normal , punctured ; shoulders slightly 

2. E. COMPTONI var. flachi (Kuwert). 

, ,, 191-. , . .^ . , ; PatipoUa, a.6oooft., Cent. ] 

Regd. No. ^ (13 sp,nt specimens). ^^^^ ^ .^^ ^^^^^^ ^_ Annandale. 

„ >i-H-^ (pinned). j wood (i3-i5-x-i9ii) / 

( Patipolla, a.6oooft., Cent. \ 

■ ..i7»-8o 9J85 I P>^ov-. Ceylon ; in rotten ^ ^ Gravely. 

' IS 18 j wood (first week in July, I 

I 1910) J 

9«^2 Ceylon H. Nevill. 

Mr. Green's collection contains specimens from Hakgalla and Namunakuli Hill. 

Description.— Length 32-37 mm. Not otherwise different from the typical 
form. In No. 'if= the process of the lower lip by which the genus is distinguished 
is paired instead of single. In the absence of any other distinguishing characters, 
however, I feel unable to regard this as more than an abnormality. 

Habits.— These insects live in galleries under the bark of fallen and decaying 
tree-trunks. Dr. Annandale tells me that the natives at Patipolla say that they 
Uve in pairs; but, as they are gregarious, satisfactory proof of this is not very easily 

220 Memoirs of the Iiulidii Mitscion. [Vol. Ill, 

obtained. He noticed that no larvae or pupae were to be seen at the time of year in 
which he was at Patipolla (middle of October), but that some of the adults were 
brown in colour and had apparently not been long hatched. 

3. E. COMPTONI, var. 
In a series of three specimens from Bulutota which are preserved in the Colombo 
Museum collection only one agrees in size with either of the preceding forms. This 
is equal to the largest specimen I have seen of E. comptoni var. flachi from Patipolla. 
The other two are very unequal in size, and the larger of them is nuich smaller 
than the smallest specimen of the series from Patipolla. Until a larger series is 
available it is impossible to be certain how these Ihilutota specimens ought to be 
regarded, but it is probable, I think, that the species E. comptoni will eventually be 
found to vary in size in the .same manner as Episphcnus indicus and certain other 
species (see below, p. 262), and that even var. flachi will prove to be unworthy of 
a distinct name '. 

Episphenus indicus, vStoliczka. 

PI. xi, figs. 20-20/>. 

Regd. No. ''^ "-f "-f- (all paralypes) S. India. F. Stoliczka, etc. 

'^-f-^ Trichinopoly Father Honore. 

!(,- N. Kanara H. I<. Andrewes'. 

.■•.7M7-i-.77o a»:2..5 „., . . ( Director , Forest School. Dclira 

13 la iMls,iris , r. u T \ 1 

\ Dun; H. L. Andrevves 

-5— Ootacamund Col. Beddome. 

^ Anamalais, S. India, en. 5000 \ 

ft.. 2I-i-I2 i 

' IF ~ui iF TiT ^T Anamalais, 4000 ft, 2 ;i- 1 2 

Anamalais, ca. 5000 ft , 2 i-i- 1 

A I • ,14. . I 111 rotten los^s. T. IJainbriege 

Anamalais, cfl. 5500 It., 21-1-12 \ ' ■^*' 

f Fletcher. 
Puddutottam, Anamalais, 

f ~i 4200 ft., 22-1-12, Pandal, 3000 ft., , 

j T. Bainbrigge Fletcher. 

Monica Estate, Anamalais, ) 

!^ \ T. Bainbriga-e Fletcher. 

4000 ft., 24-1-12 ) '^•^ 

Bainbrigge Fletcher. 







m luw 

, 19H li 



Bababudin Hills, Mysore, 1 

4000-5000 ft., 2-12-X-12 
High Range, Travancore 

4000-5000 ft., 2-12.X.12 i ^- «^'"briggc Fletcher. 

_ , Pusa Institute. 

May, 1891 I 

Collected by T. R. Bell. ^ This has since been confirmed: see below, p. 281. 

* Adults and larvae iu spirit. 

1914- J F- H. Gravely : An Account 0] the Oriental Passalidae. 221 

There are two specimens from Cuddapah (Madras Presidency), and two from 
Gudalur (Nilgiris), as well as several from the Anamalais, in the collection of the 
Bombay Natural History Society ; one fiom Naduvatum (Nilgiris), 7000 ft., in the 
collection at Dehra-Dun; and Mr. H. F. Andrewes' splendid series of this species 
includes specimens from the Nilgiris, the Anamalais, Madura, and N. Canara. I 
have examined altogether iig specimens in drawing up the following description of 
this variable species. 

Description. — Length 25-41 mm. Labnim hairy, about as long as broad, sides 
parallel or slightly convergent behind, angles rounded, the left usually somewhat 
narrower and more prominent than the right, anterior margin concave. Mandibles 
with anterior margin of upper tooth (which tooth is usually more or less distinctly bifid 
at its apex) distinct, vertical, more strongly developed on left side than on right; two 
upper terminal teeth strongly developed on both sides, the lowest of the three set 
further back, smaller than the other two especially on the right side, on left side 
often fused at base with anterior lower tooth ; right anterior lower tooth somewhat 
variable in size, always much smaller than anterior lower tooth of opposite side, 
sometimes absent, this variation not clearly correlated with size of specimen, and 
apparently inherent to some extent and not due entirely to friction. Mentiim with 
anterior margin sunk, very variable; hind wall of depression very sharply defined, 
occasionally straight or almost straight, more usually produced in the middle-line to 
form an angular prominence which is often joined by a median ridge to the anterior 
margin which may or may not be raised up to form a low rounded transverse ridge. 
Head more or less smooth and polished, sometimes punctured in front of outer ends 
of parietal ridges; parietal ridges crested, more or less arcuate, directed outwards; 
frontal area somewhat variable in form, more or less semi-lunar; inner tubercles 
distinct, but short and stout. I.eft outer tubercle directed inwards, outer margin 
f traight or very slightly convex ; ridge from left inner tubercle directed forwards 
or slightly outwards at base, but quickly curving inwards to run parallel to inner 
margin , distinct throughout whole or almost whole length of tubercle ; apex of tubercle 
bluntly rounded or truncate. Right outer tubercle acute as seen from above (in 
fresh specimens) and directed forwards, vertically truncate or imperfectly forked as 
seen from the side; supra-orbital ridges more or less crested behind, slightly peaked, 
outer wall of anterior excavation making with posterior part an angle of not less 
than 120°, often nearer 180° ; inner wall well-developed, indistinct, or absent. Anterior 
angles of head prominent, especially in small specimens, varying in size from about 
60° in the smallest to o-ver go° in the largest specimens, apt to be worn down to an 
angle of about 120° which is scarcely if at all prominent. Hollow between inner 
tubercles and supra-orbital ridges crossed by a transverse groove of variable depth and 
distinctness. Pronotum with marginal grooves incomplete across middle before and 
behind, more or less punctured especially in small specimens; scars small, incon- 
spicuous and little if at all punctured in large specimens, strongly punctured in small 
ones; sides of pronotum free from conspicuous punctures in large specimens, more or 
less punctured in anterior angles and in front of scars in smaller ones, more exten- 

222 MciHoirs ol the Indian Museum. [Vol. Ill, 

sively piitictured in the smallest of all; median groove often stronger in middle than 
at either end, probably always complete in unworn specimens. Prothorax beneath 
as in Episphenns comptoni; posterior area of prosternum very rarely bearing long 
erect hairs. Scutcllum either smooth or finely and indistinctly punctured laterally, 
pubescent striato-punctate area beside middle of anterior margin usually prolonged 
backwards as a more or less distinct double band of sparcer puncturing and pubescence. 
Mcsothoracic episterna polished and punctured above and in front, finely roughened 
across posterior angles. Surface of mcsosternum of somewhat variable texture; 
scars finely roughened, occasionally with a few large indistinct punctures, narrow or 
broad, well or ill defined. Central area of metasternum large and as a rule very 
clearly defined, with or without one or more depressions (varying in arrangement and 
distinctness) near hinder margin, and a large median circular depression in front; 
lateral areas narrow, either parallel-sided or very nearly so, not in contact with 
central area, closely punctured and hairy (usually very hairy especially in front, 
occasionally almost hairless); anterior intermediate areas very hairy, more coarsely 
punctured ; posterior intermediate areas strongly and coarsely punctured at least on 
inner side, hairless except along posterior margin where the surface resembles that 
of lateral areas. Posterior parts of hind coxae as a rule finely, scars of abdominal 
sterna more coarsely, rugose or pimctured, especially in small specimens. All grooves 
of elytra of normal width ; lateral grooves much more strongly punctured than dorsal 

Habits. — Mr. Bainbrigge Fletcher writes of the series of this and the following 
species that he collected in the Anamalai Hills : ' ' They were found in small colonies 
in rotten logs, stridulated loudly when disturbed and seemed to congregate together 
under dead wood, etc. on the ground when they were dislodged from their nidus. 
My impression was that the stridulation served as a means for rallying them together. 
In many cases, but not invariably, larvae were found in the logs together with the 
beetles." In another letter he says, "I have not noticed any particular difterences 
in the state of decay of logs inhabited by various Passalids. I should class them all 
as 'rotten.' Sometimes, whether owing to the sort of wood or exposure to sun or 
drought in the dry season, a log may be more or less tough, but I doubt whether j^ou 
can draw any hard and fast line between tough and soft. Most of the E. neelgherriensis 
from Ootacamund were found in logs which were more or less tough (perhaps owing 
partly to its being then very dry weather), but specimens of the same species from 
the Bababudins were from rotten logs so soft that we broke them up with our hands. 
In the Anamalais also, where I got E. indicus and P/eurariiis brachyphyllus, the 
logs were usually soft and quite rotten." 

Episphenus neelgherriensis (Percheron). 

PI. xi, figs. 21'Zla. 

Regd No. 5i^!«Z^" S.India { Col. Beddome, 

^ British Museum. 




F. H. Gravely : An Account of the Oriental Passalidae. 223 

T Baiiibrigge P'letcher. 

Bababudiu Hills 

High Range, Travancore 
( Coonoor, ca. 6000 ft. (Sims Park, 
\ Jungle, and Stillbrook Garden) 
[Ootacamund, 7500 ft. and 7S00 ft., 
\^ December, 191 2 ) 

, Auamalais (Kalyana, Pandal, 3000 , 
ft. ; Paralai Estate, 3600 ft.; Monica | 
/ Estate, 4000 ft.; Puddutottam, 4200 -T. Bain 

it.; ca. 5500 ft., in rotten wood; all 
I January 1912) 

Trivandruui Museum. 

R. B. Seymour Sewel!. 

Cul. Beddome. 

T. Bainbrigge Fletcher. 

brigge Fletcher. 

Description.— Length 24-29 mm. Differs from specimens of E. indiciis of same 
size in following characters only: lowest terminal and anterior lower teeth of left 
mandible always distinct at base; median keel of marginal depression of mentum when 
present usually broad and fiat, wider in front than behind, not sharply defined ; anterior 
angles of Aefli of about 120°, never prominent ; apical angle of supra-orbital ridges 
more pronounced ; lateral plates, of lower side of prothorax somewhat smoother and 
more highly polished towards the front; posterior plate of prosfernum usually hairy, 
often punctured. Puncturing of scUtellum variable in extent, but on a somewhat 
different plan from that usually found in E. indiciis, the strongest punctures being 
found in the posterior half of the plate which is often punctured almost all over 
metasterniim less markedly hairy than is usual in E. indicus. Posterior part of hind 
coxae smoother. 

Habits.— Wr. T. Bainbrigge Fletcher tells me that "Most [specimens found at 
Ootacamund in December, 191 2 J seemed to be adult beetles paired off (at least, two 
individuals together, presumably of opposite sexes) as if just ready to start new 
colonies. I only got one lot of larvae. The nights up there are frosty from Decem- 
ber to March and the weather dry, and perhaps that has some influence on the 
breeding season. Specimens found at lower elevations (4000-5000 ft.) where the 
conditions are more uniform as regards temperature and humidity do not seem to be 
seasonal, but apparently breed at any time, all stages being found." This species 
does not' seem to form colonies to the same extent as the preceding, judging from 
what Mr. Fletcher has sent me. Capt. Sewell obtained this species from under 
stones in Stillbrook Garden, Coonoor; those that he obtained from wood were all in 
extremely rotten wood from which the bark was easily removed. For further notes 
on this species see above (p. 221) under E. indicus. 

A series of small colonies (adults and larvae) each in a separate tube of spirit. 

1 1 1 

9S(ir-8 9471 94(iB 7I> 

' Sikkim ' 

224 Miinoirs 0/ the Indian Museum. [Vol. Ill, 


Ophrygonius cantori (Percheron), s. str. 

1. O. CANTORI (Percheron), s. str. 

PI. Xi, figs. 22-22(f. 

Regd. No. ^' Lainbatach, Jaunsar Forest Research Institute. 

( Stoliczka bequest, Asiatic 

1 Society's collection, etc. 

''j*^* Tista, Darjeeling District E. P. Stebbing. 

''""^•''^"" Sureil, Darjeeling District A. Alcock. 

rt.'. 9«8.i ,- r^ • 1- T-> .. • ^ ( E. A. D'Abreu. 

'■■i" ?^ kurseong, Darieeling District < ,, ^ „ . 

»« »s 5. J & ^ Museum Collector. 

'"'j;;' Gopaldhara, Rungbong Valley, H. Stevens (coll. \V. K. 

Darjeeling District Webb). 

" 'M Dikrans Valley, Dalla Hills H.H.Godwin-Austen. 

7JIT8 im\ i\u} p E. I. Company's Museum. 


I have also seen specimens from the British Museum collection front Gantok, 
Bhutan, and Manipur. 

The collection of the Imperial Forest Research Institute at Dehra Dun includes 
a specimen found boring in Sauer wood in the Teesta forest, and several froin 
Lambatach in Jaunsar, Dehra Dun, the latter being the most western Himalayan 
record I know for any Passalid, except one of the same species in the same collection 
from Tehri-Garhwal^a record which, I think, may be incorrect, as this has been 
proved to be the case for all other Passalids recorded from the same place that I 
have seen. 

Description. — Length 30-34 mm. Resembles both specimens of Episphcnus 
indicus of similar size and E. neelgheyriensis except in the following particulars. 
Anterior margin of lahrum straighter, sometimes with suggestion of slight median con- 
vexity such as is found in Ophrygonius inaequalis. Mandibles with upper tooth some- 
what less strongly developed, anterior lower tooth variable but more strongly deve- 
loped on an average than in Episphcnus indicus or E. neelgherriensis, never entirely 
absent, left lowest terminal and anterior lower teeth distinct at base as in E. 
neelgherriensis. Depression of anterior margin of mentum consisting of a pair 
of more or less separate concavities whose posterior wall is usually oblique, 
not vertical, the angle at which it meets the general upper surface being in con 
sequence blunter than in either of the two preceding species. Head punc- 
tured and somewhat hairy between central tubercle and supro-orbital ridges, 
both in front of parietal ridges and behind them; parietal ridges straighter, 
their outer extremities usually more or ] strongly directed backwards; left outer 
tubercle obliquely truncate, its ridge from left inner tubercle strongly marked and 
extending straight forwards to the acute anterior {i.e. outer) angle, thus giving -the 

1914.] F. H. Gravely : An Account of the Oriental Passalidae. 225 

whole tubercle an appearance of being pointed and directed more or less straight 
forwards; right outer tubercle of more or less similar (sometimes the same) shape, but 
shorter (although longer than in the two preceding species), apex simply pointed 
in profile, its ridge from inner tubercle directed slightly outwards; no transverse 
groove between inner tubercles and supra-orbital ridges ; ^^upra-orbital ridges with 
more or less peaked apical angle of about 120°, inner wall of anterior excavation 
very rarely suppressed; anterior angles of head containing about 120°, not pro- 
minent. Pronotum almost invariably punctured and hairy to some extent in the 
posterior angles and on the sides, prothorax beneath much as in Episphenus neelgher- 
riensis. Scutellum entirely smooth, hairless, unpunctured, and highly polished with 
exce]3tion of usual patch in middle of anterior margin ; mesothoracic episterna polished 
throughout, punctured above and close to anterior margin; mesostermim smooth and 
highly polished throughout or matt in the lateral angles ' , scars entirely absent or 
(rarely) indicated by a pair of slight depressions of the usual shape in the usual position 
Metastemum with central area very vaguely defined; posterior intermediate areas 
unpunctured except along posterior margin ; lateral areas broadened behind by the 
curvature of their inner borders. Abdominal sterna and posterior parts of hind coxae 
as in Episphenus neelgherriensis , often still smoother. Elytra with small tuft of 
hair at shoulders, grooves more weakly punctured throughout. 

2. O. CANTORi, subsp. coNVExiFRONS, Zang. 

Regd. No. ^^ Khasi Hills, Assam H. H. Godwin-Austen. 

"^^^^ Teriagbat, Khasi Hills H.H.Godwin-Austen. 

In the British Museum collection there are two specimens from the Ruby 
Mines District of Burma. 

Description. — Length 27-29 mm., otherwise as in 0. cantor i, s. str. The difference 
in size is an extremely small one, but appears to be constant in specimens from the 
Khasi Hills and Burma. One of the specimens from the Rubv Mines has the mentum 
sculptured ?lmost as in the following sub-species. 

3. O. CANTORI, subsp. DUNSiRiENSis, n. var. 

PI. xi, fig. 23. 

Regd. No. --ff (type) ''^ Dunsiri Valley, Assam H. H. Godwin-Austen. 

Description. — Length 28-31 mm. Resembles the two preceding forms in all respects 

except the following : posterior wall of depression of anterior margin of central part 

of mentum continued straight outwards for a short distance on to surface of lateral 

areas, then turned suddenly backwards, extending more or less distinctly to posterior 

lateral angles, this final portion of the wall being occasionally so formed as to produce 

a false superficial semblance of the scars found in the genus Macrolinus. 

' Apparently always polished throughout in Darjeeling specimens. 

226 Memoirs of the Indiau Museum. [Vol. Ill, 

Ophrygonius birmanicus, n. sp. 
Text-fig. 3A. 

Described from a single specimen from the Ruby Mines District of Burma, 
preserved in the British Museum collection. 

Description. —Length 35 mm. Differs from 0. caniori in the following characters 
onh'. Lamellae of antennie somewhat longer (but the first two not as long as the 
last four). Anterior lower tooth of both mandibles rather small but very sharply 
pointed. Left outer tubercle of head very slender, almost parallel-sided, sunply trun- 
cate distally, directed very distinctly inwards. Scars and lateral angles of meso- 
sternum matt. Posterior intermediate and lateral, areas of metasternum less distinct 
from one another behind, the puncturing of the latter extending a little distance 
over on to the surface of the former; posterior parts of hind coxae, and scars of 
abdominal sterna, scarcely as smooth. 

Text-i-igure 3. 

A. Ophrvgoniiis birmanicus , head x 4. 

B. Ol>hrygonms singapv.rae, head x 4. 

Ophrygonius singapurae, u. sp. 
Text-fig. 3B. 

Described from a single specimen from Singapore in the collection of the 
Konigliches Zoologisches Museum in Berlin. 

Description. —Length 31 mm. This interesting species is closely allied to the 
preceding, with whose characters it combines some of those of Ophrygonius inacqua- 
lis on the one hand, and of Aceraius wallacei on the other. The lamellae of the 
antennae are all very long and slender as in A . wxllacei. The lowest terminal tooth 
of the m.indibles is probably well-developed on both sides in fresh specimens, as is 
usual in this genus ; but that of the right side is very obtuse ( probably through 
friction) in the specimen before me ; the anterior lower tooth of both mandibles is 
very broad , and flattened dorso-ventrally, resembling that of the left mandible of 
most species of Aceraius, though the horizonal margin is scarcely grooved. The 
mentum resembles that of Ophrygonius inaequalis. The head bears distinct, finely 
keeled, frontal ridges (not grooves) as in 0. inaequalis; the left outer tubercle 
is intermediate in form between that of 0. cantori and 0. birmanicus ; the parietal 
ridges are somewhat short ; the supra-occipital ridge is united to the supraorbital 

igi^.] F. H. Gravely : An Account of the Oriental Passalidae. 227 

ridges as usual, but is also continued outwards behind them as in specimens of 
the sub-family Macrolininae— I know of no other species of Passalid in which this 
occurs. The sides of the pronotiim are closely punctured below the scars throughout 
their whole length ; the lateral plates of the ventral side of the prothorax are some- 
what rougher and more punctured in front than in 0. birmanicus, which this species 
resembles in all other respects. 

Ophrygonius inaequalis (Burmeister). 
PI. xii, fig. 24-24^?. 

/■ 9120 9]2-l^ 9:UI-3 -J 

Regd. No. ] \j5i3"34o ' [Malacca Stoliczka bequest. 

' ~r 'w- ^ 

*^- Perak Moti Ram. 

~ 967|ii «s«. siukep Island Moti Ram. 

Description. — Length 24-27 mm- lyamellae of antennae (very short : only four of 
them pubescent. Labrum hairy, but very sparsely punctured; sides slightly con- 
vergent behind ; angles rounded, left one sometimes very much drawn out ; 
anterior margin concave on both sides, convex in middle. Mandibles with upper 
tooth obsolete on both sides; anterior lower tooth acutely conical (very sharp 
when unworn), that of left side larger than that of right. Depression of anterior 
margin of median part of mentum extensive, middle part of its posterior wall 
bent forwards to form an angular prominence dividing depression into two more 
or less completely separated halves. Head polished, but somewhat roughened 
or punctured in front and at sides of (more rarely within) frontal area, and 
sometimes about the middle-line behind central tubercle; parietal ridges directed 
outwards, straight or somewhat S-shaped, usually defined in front by a groove; 
keels of parietal and frontal ridges (latter sometimes coalescent behind) strongly 
developed, continued to meet one another at apex of central tubercle; frontal 
ridges so widely arched (even angular in some specimens) as to be parallel or 
convergent in front ; inner tubercles indistinct, situated at base of inner side of 
outer tubercles; margin of head between outer tubercles vertical, slightly (often 
very indistinctly) convex in middle ; left outer tubercle very broad and more 
or less inwardly directed at base, then abruptly narrowed from the outer side 
and turned more forwards, bluntly rounded at the tip; ridge from left inner 
tubercle straight and directed vertically forwards at the base, curving inwards distally 
only where it has become faint; right outer tubercle somewhat long, pointed as seen 
both from above and in profile, apex directed forwards and a little outwards; supra- 
orbital ridges broadly rounded, not keeled behind, in front excavate, descending 
abruptly at an angle of about 120° which is slightly peaked, the two walls of the 
excavation keeled, outer one somewhat concave above forming with anterior angle 
of head, an upwardly-directed tooth ; anterior angle of head containing somewhat 
more than 120°, canthus vertically truncate at its extremity. Pronotum polished, 

228 Memoirs of the Indian Museum. [Vol. Ill, 

marginal groove incomplete across the middle both before and behind, its anterior 
extremities merged in a pair of somewhat triangular strongly pmictured areas, 
behind each ot which a more or less definite band of punctures extends back parallel 
to the sides, crossing in their course the upper parts of the two scars, which are 
also thickly punctured. Median groove very distinct, almost invariably complete. 
Punctures and hair on lateral plates of lower side of prothorax somewhat sparser 
than in 0. cantori ; posterior plate of prosternum with a few hair-bearing punctures, 
anterior plate unusually hairy. Scutellum striato-punctate and pubescent beside 
greater part of anterior margin, punctures continued backwards on either side of 
middle line about half way to posterior margin; otherwise smooth and polished. 
Mesosthoracic episterna polished and punctured above and along anterior margin, 
dull and unpunctured in posterior angles. Mesosternum striato-punctate in extreme 
anterior angle; scars dull, small, but continuous behind and on the inner side with a 
dull and finely punctured area occupying the whole of the lateral angles ; the rest 
highly polished. Metasterniim with lateral areas rough, slightly hairy in front, very 
narrow, parallel-sided or nearly so; central area extensive, badly defined; anterior 
intermediate areas somewhat hairy, but rather indistinctly punctured; posterior 
intermediate areas completely or very incompletely covered with sparsely or rather 
thickl}' scattered strong punctures, which are thickest close to the posterior margin. 
Posterior parts of hind coxae somewhat, scars of abdominal sterna strongly and 
extensively, roughened or finely punctured. Lateral grooves of elytra wide, with more 
or le.-^s transversely linear punctures ', dorsal grooves normal ; shoulders slightly hair3^ 

Genus ACERAIUS, Kaup. 

Aceraius wallacei (Kuwert). 

PI. xii, figs. 26-266. 

Rc-gd. No. "'Jy' Kuching, Sarawak Sarawak Museum. 

The Berlin collections include specimens from the Malay Peninsula and from 
Mt. Kina Balu; and the Hamburg collection specimens from Somgei Lalah. 

Description.— Length 32-37 mm. Labriini hairy, punctured, sides parallel, angles 
rounded, left angle more pointed but hardly more prominent than right. Antennae 
with all six lamellae pilose and very long and slender. Upper margin of mandibles 
behind upper tooth straight or slightly concave ; apex of right upper tooth acute ; 
left upper tooth acute or truncate, bent over inwards a little; horizontal groove 
of left anterior lower teeth very pronounced, dividing tooth into two parts of which 
the upper is often somewhat larger and blunter than the lower ; lowest terminal and 
anterior lower teeth of right side very small, both dorsal and ventral portions of 
right anterior lower tooth distinct. Median part of menium usually with sunken 
semi-circular area along anterior margin and three or four large punctures behind. 

See also below, p. 285, footuote. 

1914.] F- H. Gravely : An Account of the Oriental Passalidac. 229 

the sunken area sometimes indistinct or absent. Head above with frontal area more 
or less semi-circular, more rarely triangular; ridge joining inner tubercles usually 
slightly convex in middle, sometimes very strongly so ; left outer tubercle obliquely 
truncate, truncation straight or very slightly concave, both angles somewhat produced 
as a rule in fresh specimens; right outer tubercle not much shorter than left, but 
more nearly triangular, apex single in profile and very acute. Prothorax and meso- 
thorax as in .4. bomeanus except that the pronotum is less punctured at the sides while 
its median groove is obsolete, and that the mesosternuni is polished, throughout or 
nearly so, except in the long narrow scars. Metastevnum also as in A. borneanas 
except the posterior lateral areas in which a variable number of very coarse and often 
somewhat elongated punctures are more or less extensively developed from in front 
along the inner margin. Hind coxae and abdominal sterna as in A. bomeanus. Elytra 
not very hairy, the hair all short; tenth rib finely punctured and hairy close to 
shoulders, ninth rib with a few punctures scattered very sparsely along its whole 
length, seventh rib with or without even more sparsely scattered punctures. 

Aceraius perakensis, Kuwert. 

Described from three specimens from Taiping, 4000-4500 ft , and one from Mt. 
Singaleng ; all in the collections in Berlin. 

Description.— hength 38-5-42 mm. Labrtim as in preceding species; antennae as 
in following one ; left upper tooth very variable, left anterior lower tooth not so 
strongly divided as in preceding, m^n^tWes otherwise similar ; mentum without any 
anterior depression, but punctured, very sparsely or not at all in middle, not so 
closely as in following species. Upper surface of head somewhat as in preceding 
species, but with left outer tubercle stouter, ridge joining inner tubercles concave, 
inner side of right outer tubercle straighter and more oblique, apex of same tubercle 
truncate or faintly bifid in profile, canthus usually a little swollen at base; an- 
terior angles obtuse as in preceding species, not prominent. Thorax, abdomen and 
legs as in preceding species; elytra as in following. 

Aceraius mbschleri Kuwert. 
PI. xii, fig. 33. 
Regd. No. '^ Sinkep Island Moti Ram. 

I have also seen a specimen in the British Museum collection from the Malay 
Peninsula, and one from lyingga and another from Java in the Berhn collections. 

Description.— Length 36-42 mm. Difiers from the following species only in having 
the anterior angles of the head scarcely prominent on either side ' ; and in having the 
apex of the right outer tubercle less bent outwards as in A. perakensis. 

I This difference is noted by Kuwert in his brief diagnosis of 1891, but is omitted from the longer 
one of 1898, in which the two species are separated by the shape of the anterior margin of the labrum — 
a character which is always apt to vary somewhat in different specimens of a single species. 

230 Miinoirs o/ the Indian Museum. [Voi<. Ill, 

Aceraius illegalis, Kuwert, s. str. 
PI. xii, figs. 32-32(^ 

Regd. No. 'f^" •";;;' Johore J. MeMruiii ; Mpuftluisefl). 

I have also examined other specimens, inchiding one from Kuching and one from 
between Ulu Madahit and lyimbang which are i^reserved in the Sarawak Museum. 

Description.— liQngth. 39-43 mm. Labrum as in A. wallacei. Lamellae of 
antennae soniiewhat short and thick, first one pilose only round margin of anterior 
face. Mandibles as in A . wallacei. Mentum also as in that species but more closely 
punctured in middle. Anterior angles of head prominent, that of left side very 
slightly more so than that of right ; truncation of left outer tubercle straight or only 
slightly concave, vertical to the two parallel sides of the process, the outer side often 
with a thickening which is more or less rectangular; right outer tubercle rather long, 
with acute undivided apex bent slightly outwards. Prothorax as in .1. ivallacei. 
Scutellum with the punctured area of its anterior margin apt to be carried unusually 
far back along the middle line; mesothoracic episterna and sternum as in A. wallacei, 
except for presence of indistinct hair-bearing punctures in posterior angles of meso- 
thoracic episterna. Greater part of posterior intermediate areas of metasternum 
smooth and unpunctured, but with a few both of the coarse inner and fine posterior 
punctures always more or less distinctly present. Posterior parts of hind coxae dull, 
abdominal sterna polished, both smooth or very nearly so. Elytra somewhat sparsely 
covered with longer or shorter hair, seventh rib often, ninth almost always, rather 
sparsely punctured. 

Aceraius laevicollis (lUiger). 
PI. xii, figs. 2y-2ya. 

Regd. No. ?^ ?^ Johore J. Wood-Mason, etc. 

6«6.0 3046-50 _,. , Til ir i- n 

" TP ~T9~ Sinkep Island Moti Ram. 

>> ^^ Kuching, Sarawak, \ 

Borneo ) Sarawak Museum. 

>> ^ I.imbong, Borneo j 

The Sarawak Museum collection includes specimens from Busa and Serai ; the 
Berlin collections include specimens from Perak, Deli, Sol ok, Peinan, Nias, Lingga, 
Billiton, Java, Bandjermasin, Luzon and S. Palawan; and the Hamburg collection 
specimens from Deli, Serdang, Somgei Lalah, Tandjong and Banguey Island. 

Description. — Length 29-34 mm. Lamella of fifth joint of antennae long and pilose, 
the fourth joint sometimes furnished with a short lamella devoid of pilosity. Labriim 
as in preceding species. Mandibles also as in that species except that the upper 
margin of the left mandible behind the upper tooth is more or less convex and 
usually distinctly turned over inwards. Mentum more or less punctured, its anterior 
margin often sunk on each side of a median prominence which may be more or less 
distinctly grooved in front in the middle line. Head above remarkably variable • 


F. H. Gravely : An Account of the Oriental Passalidae. 


Anterior angles always distinctly prominent to greater or less extent, left one pro- 
jecting more or less markedly further than right. Angles of truncation of left outer 
tubercle less produced than in A. wallacei; the tubercle as a whole very variable, 
usually much broader than in that species, though not always, and as a rule 
thickened near base on outer side, this thickening when unworn and very well deve- 
loped rectangular as in fig. 27, but more often blunter and occasionally absent. Right 
outer tubercle normally short; very broadly and more or less obliquely truncate 
as seen from above, the truncation being sometimes slightly convex, sometimes 
straighter, and sometimes even sufficiently concave to produce a bidentate form, the 
outer and still more rarely the inner of the two denticles so formed, being occa- 
sionally produced into an acute, forwardly directed tooth. Frontal area more or less 
triangular. Parietal ridges perhaps inclined a little more backwards as a rule than 
in A. wallacei. Supra-orbital and supra-occipital ridges forming together an open 
semi-circle, the former approximately parallel to one another in front, rarely or never 
in the least degree convergent. Prothorax and scutellum as in A . wallacei] mesothoracic 
episterna with a patch of fine hair-carrying punctures in posterior angles as in .4 . illegalis ; 
surface of mesosternum very variable in texture, being sometimes extensively roughened 
and in one specimen' even coarsely granular in parts; scars variable in form and 
definition. Posterior intermediate areas of metasternum as a rule more completely 
and evenly punctured than is usual in the genus. Posterior parts of hind coxae 
finely granular throughout; abdominal sterna- as in preceding species. Sides of elytra 
pilose, densely near shoulders, more spacely further back where the pile is almost 
confined to the grooves, the ribs (except the anterior part of the tenth) being more 
or less completely devoid of punctures ; the ninth rib more persistently punctured 
than any other. 

Regd. No, 

Aceraius grandis (Burmeister). 
A. GRANDIS subsp. HiRSUTus, Kuwert 
PI. xii, figs. 28-29. 
. . Darjeeling 

9259-01 S5i5 

3032 3282 
19 19 

9455 328 t 
1 19 




Harmutti, Base of Dafla Hills 

Dikrang Valley 

Dunsiri Valley 

Duffla Expedition 

N.-E. Frontier 

1' Stoliczka bequest, 
; J. Wood-Mason. 

I Dr. J. Anderson, 
' E. T. Atkinson. 
I L. Maudelli. 

H. H. Godwin- Austen. 

That from Serai in the Sarawak Museum Collection. 

S. W. Kemp. 

Memoirs of the hid tan Museum. [Vol. Ill, 

Regd. No. ?^ KoImj, Abor Country, 400 ft. \ 

-^~ Janakmukh, Abor Country, 600 ft. 

"■^-■'(Spirit specimens) Rotung, Abor Country, 1400 ft. 
.. ^^ ~i^ Ujjper Rotung, Abor Country, 2000 

"Jf^ Kalek, Abor Country, 3800 ft. 

T Cachar j. Wood-Mason. 

/ 912a iUSl <J1S3 MW-SI X 

'• ' \: '• ' Uibsagar J Stoliczka bequest, 

rr^-T-.^;-' J - ^ S. E. Peal, Moti Ram. 

. Ut4-: 670 7114 7119 J 

\i s'iL"' "' (^'^^s'H'l's Shillong Museum. 

V 14 ' W • / 

'-^' .. ■• Tavoy .. .. .. .Museum Collector. 

.10401 ( Between Thinganuyinauug and ^ 

'^ " "\ Sukli, Dawna Hills, 900-2100 ft. ) "' ^'"^^'>'- 

•' "is"'' 15^' •• Sukli, Dawna Hills, c. 2100 ft. .. F. H. Gravely. 

Mr. vSteven's collection includes a specimen collected by Mr. W. H. Webb at 
Gopaldhara in the Rungbong Valley, Darjeeling District, and several from Silonbari 
at the base of the N. I^akhimpur Hills. The Berlin collections include specimens from 
Formosa (numerous localities), Hainan, S. Palawan, Tonkin, and Catchin Couri. The 
British Museum collection includes also .specimens from Sin Lum, and Cambodia. 

Description.— Length 33-49 mm. Labruni as in A. leavicollis. Lamellae of 
antennae much as in A. . leavicollis in small specimens,^ often proportionally shorter and 
thicker in large ones. Mandibles also much as in ^. leavicollis in small specimens, the 
recurved upper edge behind the left upper tooth often broader, especially in front in 
larger .specimens, sometimes forming a stout secondary tooth in the largest of all, its 
outer margin concave rather than convex. Mentum as in A . leavicollis. Anterior angles 
of head much more prominent than in A. leavicollis in most small, but very little more 
prominent in most of the largest specimens, almost always directed more or less inwards 
in the former, in which, consequently, the supra-orbital and supra-occipital ridges 
together appear to enclose more than half a circle; in the latter, the area enclosed by 
the supra-orbital and supra-occipital ridges usually resembles almost and often quite as 
perfect a semi-circle as in A. leavicollis; left outer tubercle somewhat less variable, 
usually rather long and .slender, parallel sided, and scarcely forked ; sometimes broader 
and shorter; rarely with even the faintest trace of an external enlargement ; right outer 
tubercle variable as in A. leavicollis, usually more pronouncedly bidentate with the 
two teeth somewhat nearer together ; frontal area often very short and broad , especially 

Of these specimens only the head and prothorax, in one case together with a single elytron, were 

In one si)ecimen from vSikkim (No. 'y,}') there are only five lamellae altogether, that of the sixth 
segment being suppressed or possibly reduced and fused with that of the fifth. I have seen no trace 
of a lamella on the fourth segment of any specimen. 

I9I4-] P- ^- Gravely : An Account of the Oriental Passalidae. 233 

in large specimens. Prothorax and mesothorax as in A. laevicollis, hut patch of fine 
punctures in posterior angles of raesothoracic episterna very variable and sometimes 
absent especially in small specimens. Posterior intermediate areas of metasternum 
marked on same plan as in rest of genus, but very variable; lateral areas very broad 
behind as in all the preceding species. Posterior parts of hind coxae often less densely 
granular than in A. laevicollis. Abdominal sterna as in that species. Seventh and 
ninth ribs of elytra almost always punctured throughout, usually densely in large and 
more sparsely in small specimens— in the smallest specimen I have seen there are 
very few punctures indeed on the ninth rib and even fewer on the seventh' ; sides of 
elytra clothed with a more or less short ' pile which is dense near the shoulders and 
sparse behind. 

Habits. —Mr. Kemp tells me that this form is gregarious. It was found to be 
very common in the Abor country, boring in tough wood under moderately thin bark. 

2. A. GRANDis (Burmeister, s. str.).' 

The Berlin collections include specimens from the Malay Peninsula, Bintang, 
Deh, Tengger Mountain, Bankalan, Bandjermasin, and N. Borneo. 

Description. — Length 39-52 mm. This form and the following variety are the 
representatives in the Sunda Islands and Malay Peninsula, of the preceding sub- 
species. They cannot be very sharply distinguished either from one another or from 
the northern race, as all three are highly variable. In the two southern forms, 
however, the length of the hair on the elytra is more variable, and usually much 
longer ; the seventh rib of each elytron is always unpunctured, whereas in the 
northern race it is almost invariably punctured, at least sparsely and often closely ; 
and the shape of the prolongation of the left anterior angle of the head seems to be 
more constant. In the present form this angle is always at least moderately long, in 
large as well as in small specimens, and is always curved inwards, as in the specimen 
of the northern race shown in fig. 28. 

3. A. GRANDIS var. rectidens, Kuwert.' 
PI. xii, fig. 30. 

The Sarawak Museum collection includes one specimen from Sarawak ; and the 
Berhn collections specimens from the Malay Peninsula, Solok, Tebing tinggi and Java. 

Description.— Length 47-54 mm. Resembles the preceding in all points except 
the following: upper tooth of left mandible always simple and acute, never bifid or 
with the upper margin convex and bent over inwards as it usually is in A. grandis, 
s. str. and sub-sp. hirsutns] prolongation of left anterior angle of head often moder- 
ately long, but as a rule shorter than in A. grandis s. str., directed forwards or slightly 
outwards, never curved inwards. 

' The seventh rib is unpunctured in most of the Hainan specimens, and in occasional ones from 
other locahties; the density of the puncturing is very variable even in specimens all of the same size. 

^ The hair is very short and close in all the Chinese, Himalayan and Assamese specimens I have 
seen, but is more variable in those from Tonkin and Burma. 

■^ See below, p. 322, last paragraph of footnote 2. 


Memoirs of the 1 iiduni Museum. 

[Vol.. Ill, 

Aceraius occulidens, Zang. 
Text-fig. 4A. 

I have examined two specimens, both preserved in the Deutsches Entomologisches 
Musemii. One (the type) is from Mt. Kina Bahi (c. 5000 ft.) in Borneo, the other is 
from DeH, Sumatra. 

Description. Lentith 48 mm. This s])ecies differs from .A. t^raiidis var. rectidens 
only in the presence of a stout upwardly directed tubercle on the canthus, just in 
front of the eve. 

Text-figure 4. 

A. Aceraius occulidens, head x 4. 

B. Aceraius triconiis, head x 4. 

C. Aceraius alutaceosternus, left mandible from the side 

D. Aceraius minor, head x 4. 

E. Aceraius acquideris, head x 4. 

Aceraius laniger, Zang. 

Two specimens of this species, both from Mt. Kina Balu, c. 5000 ft., are i)re- 
served in the collection of the Deutsches Entomologisches Museum. 

Description .—Length 47-52 mm. Differs from A . kuwerti in the following charac- 
ters only : right outer tubercle of head inclined to be more acute, apical angle of 
supra-orbital ridge incHned to be less so ; ridge separating lateral and intermediate 
areas of metasternum obtuse, rough ; tenth rib of elytra closely punctured and hairy 
at shouldefs as in A. occulidens and all preceding species, the hair on the elytra all 
shorter and closer. All these characters are apt to be variable in other species, and 
I doubt whether this form is really as distinct from the next as at present it seems 
to be. 

igi4.J F- H. Gravely : An Accoimt of the Oriental I'ussalidae. 235 

Aceraius kuwerti, Zang. 
PI. xii, fig. 31. 

The Berlin collections include a number of .specimens from Mt. Kina Balu, 
c. 5000 ft. 

Description. — Length 43-49 mm. This species resembles large specimens of 
A. grandis sub-sp. hirsutus in all characters except the following : upper tooth of left 
mandible somewhat more prominent, always bidentate in unworn specimens; anterior 
angles of head not prominent, symmetrical; apical angles of supra-orbital ridges 
inclined to be more prominent ; right outer tubercle usually more or less obsolete ; 
ridge separating lateral and intermediate areas always sharp , smooth and polished ; 
posterior parts of hind coxae apt to be less extensively punctured than is usual in 
A. grandis, sometimes finely rugulose or entirely smooth ; anterior partof tenth rib 
of elytra hairless and unpunctured. 

Aceraius tricornis, Zang. 

Text-fig. 4B, p. 234- 

There are examples of this species both in the British Museum and in the 
Deutches Entomologisches Museum ; all of them are from Mt. Kina Balu (c. 5000 ft.) 

Description. — Length 49-53 mm. Differs from the preceding species in the fol- 
lowing characters only : right outer tubercle of head absent ; inner tubercles minute, 
that of left side situated on inner margin of left outer tubercle, that of right side 
situated further back; apical angle of supra-orbital ridge still more prominent, and 
situated further forwards along inner wall of anterior truncation , where it forms 
a conspicuous tubercle directed upwards and a little inwards. 

Aceraius laevimargo, Zang. 

Described from specimens from Mt. Kina Balu, c. 5000 ft. in the collections of 
the British Museum and Deutsches Entomologisches Museum, and two from Sumatra 
in the latter collection. 

Description. — Length 34-39 mm. Differs from .-1. laniger and .4. k im e rt i'Ay 
in its smaller size and much greater variability.' Lamellae of antennae remarkably 
variable, the whole range of variation found in the genus being found in this single 
species; right outer tubercle of head quite as well developed as in ^. laniger, simply 
pointed in profile.^ Metasternum and e/y^m variable, resembling those of .4 . /^mge^ 
or A. kuwerti, or intermediate between the two; the seventh rib of the elytra some- 
times almost unpunctured. In all other characters this species resembles the two 
preceding. The scars on the mcsosternurn are almost always distinct. 

Aceraius pilifer (Percheron). 
PI. xii, fig 35. 
The specimens I have seen include insects from Sumatra in the British Museum 
collection; from Tjibodas, c. 5000 ft., and Preanger, 4-6000 ft. in Java, in the 

' The apex of the left upper tooth is probably always bifid in fresh specimens. 

^ But for size and this characteristic the species might easily be confused with .A. perakensis. 

236 Memoirs of the I ndiiin Museum. [Voi,. Ill, 

Hamburg and Berlin collections; and from Borneo in Mr. H. E. Andrews' collection 
and the Berlin collections. 

Description. — Length 28 32 mm. This species is very closely allied to the preced- 
ing, from which it differs chiefly in its smaller size. It also differs, however, in a few 
other characters ; and, although some of these are very variable in the preceding 
species, I prefer to keep the two distinct for the present as I have seen no true 
intermediates. Lamellae of .\ ntennae never very long and slender ; right outer 
tubercle of head somewhat more truncate in profile than in laevimargo in fresh 
specimens, the suture distinctly bent downwards; scars obsolete on mcsosternnm; 
anterior part of tenth x\h of elytra always punctured and hairy. 

A. alutaceosternus, Kuwert. 
PI. xii, figs. 34-34^'; text-fig. 4C, p. 234. 
Regd. No. ^' I'eiiaiig F. Stoliczka ? 

This specimen has been labelled Aceraius emarginatus by Stohczka, and I have 
little doubt that it is the specimen which he says he caught on Penang Hill (1871, 
p. 158). I have also seen a specimen from Taii)ing 4000-5000 ft., in the collection 
of the Deutches Entomologisches Museum. 

Description. — Length 35"5-36-5 mm. Lahrum as in A. laevicoUis. Mandibles also 
as in that species, but with convexit}' behind left upper tooth exceptionall}^ large. 
Lamella of fifth joint of antenna short and devoid of the close pubescence with which 
the five succeeding lamellae are clothed. Mentuni slightly prominent and more or 
less indistinctly grooved in the middle line in front. Anterior angles of head not pro- 
minent. Left outer tubercle rather long and slender, bifid at the end, the anterior 
angle somewhat longer than the inner one, the former directed somewhat upwards and 
the latter downwards; right outer tubercle longer than in .4 . borneanus, hnt shorter 
than in A.wallacei, triangular as seen from above, imperfectly forked at apex in 
profile. Prothorax much as in A. wallacei. Mesothorax as in A. illegalis except for 
the presence of a very small closely punctured area in the lateral angles of the 
scutellum, and the absence of punctures from the posterior angles of the episterna. 
Metasternum, hind coxae and abdominal sterna also much as in A. illegalis. Elytra 
with seventh rib sparsely, ninth somewhat more thickly punctured ' ; tenth rib 
with hair-bearing punctures in front ; hair long and not very thick. 

Aceraius himalayensis, n. sp. 

PI. xii, figs, ^d-^ba. 
Regd. No. '"f" ■■'^ Sikkim f Stoliczka bequest, J. Wood- 

\ Mason. 
"",'','■' ']1' (lypi:) Dikrang X'alley H H. Godwin- Austen, 

lir S Sirpo River, Nr. Ren'.<ing. \ m H C 

< Abor Country S 

't^ Naga Hills Robert, through J. Wood- 


' This, however, is no doubt a variable character as it has proved to be in other species: in 
Kuwert's type only the ninth rib was punctured. 

I9I4-1 F- H. Gravely : An Account of the Oriental Passalidae. 237 

I have also examined specimens from Pedong in the British Museum collection. 

Description. — Length 31-36 mm. Labrum, antennae and mandibles as in A . alutaceo- 
sternus, but convexity behind left upper tooth not unusually large. Mentum 
punctured in middle, a very short and more or less indistinct median groove sometimes 
discernible on anterior margin. Anterior angles of head not prominent lycft outer 
tubercle broad, its external margin sinuous but without distinctly rectangular enlarge- 
ment ; extremity truncate and somewhat concave, its anterior angle a little more 
produced than the inner one. Right outer tubercle rather short, apex simple or very 
imperfectly divided in profile. Frontal area very variable. Prothorax as in A. wallacei 
except that the imperfect median groove is more distinct. Scutellum polished, but 
more or less marked with indistinct punctures ; mesothoracic episterna and sternum as 
in A. alutaceosiernns ; posterior intermediate areas of metasternum with same scheme of 
markings as in other members of the genus,' the extent and distinctness of these markings 
very variable. Hind coxae and abdominal sterna much as in /I. wallacei^ the latter 
perhaps a little more distinctly punctured laterallJ^ Sides of etytra clothed with 
short hair, seventh rib usuall}^ more or less thickly, ninth always thickly punctured 
throughout, tenth with hair-bearing punctures in front. 

Aceraius assamensis, Kuwert. 
PI. xii, figs. 3>7-Z7(^- 

Regd. No. '^'y Cherra Punji H. H. Godwin-Austeu. 

,t 7^ Manipur ,, 

„ ''-—^ Assam British Museum. 

7070 7082 2859 p MacClclland, E. I. Com- 

pany s Museum, etc. 
Description. — Lewg^A 31-36 mm. Differs from A. himalayensis in the following 
respects only : mentum sparsely or not at all punctured in middle ; inner process of left 
outer tubercle somewhat less acute, but usually larger in proportion to anterior process, 
in consequence of which the tubercle as a whole seems even broader ; apex of right 
outer tubercle more or less distinctly bifid in profile, apex of upper portion always 
less than half as far from apex of lower portion as from right inner tubercle ; scutellum 
as in A. gr a ndis; posterior parts of Ani^i coxae, and scars of abdominal sterna less 
smooth ; seventh rib of elytra with or without punctures at anterior or posterior end 

Aceraius tavoyanus, n. nom. 
PI. xii, figs. 38-38((. 
This species is undoubtedly the same as that from Burma, erroneously described 
by Kuwert in 1898 under the name pilifer , Percheron. 

Regd. No. f, '^l,—^ Tavoy Museum Collector. 

There is a specimen from Siam in the British Museum. 

.e. coarsely punctured along inner more finely along posterior margin. 

238 Mcmoiis of the Indian Muscnm. | \'oi,. Ill, 

Description. ^Length 30-33 mm. Diflfers from A. assuntcnsis in the following 
characters only: left outer tubercle slender, its distal angles more produced; median 
groove of pronotum entirely obsolete as. in A. wallacci; posterior x^diXis oi hind coxae 
finely rugose ; abdominal sterna as in .4. hima'ayensis ; seventh rib of elj'tra punctured 
at posterior end only in the four specimens before me, as it was in Kuwert's 
specimens also. 

Aceraius helferi, Kuwert. 



/•5877 .nsst! 

PI. xii, figs. 39^39"- 
^ Teuasserim 

J. Wood-Mason. 



Upper Tenasseriin 

J. Wood-Mason. 

,282S.» istl 

r-nr ,„ 

/ '^ (adult and lar- 

I vae inspirit) 



' lietween Misty Hollow and Sukli, 
1 Dawna Hills, 2100-2500 ft. 

( Between Thingannyinaung and ; 
(. Sukli, Dawna Hills, 900-2100 ft. 

1 F. H. Cravely. 
^ F. H. (bravely. 




Bombay Natural 

There are specimens from Rangoon in the collection of the Bomljay Natural 
History Society; and specimens from Carin Ghecu 4000-5000 ft., Carin Asciuii Cheba 
3500-4000 ft., Carin Cheba 3000-3500 ft., and Mt. Mauson 2000-3000 ft. in the 
Berhn collections. 

Description. — Length 31-40 mm. Differs from .-1. favoyaniis in the following 
characters only: left outer tubercle, though not quite constant either in size or shape, 
nornaally with inner process more massive, sometimes itself truncate; right outer 
tubercle somewhat shorter, distance between apices of upper and lower portions of 
its extremity greater, always more instead of less than half as great as distance 
between apices of upper portion and right inner tubercle ; posterior parts of hind 
coxae usually smoother; seventh rib of elytra with or without punctures throughout, 
or more often at the posterior end only.' 

Habits. — This species lives in logs that have not decayed to any great extent. 
It does not appear to be gregarious, adults being found only singly or in pairs, some- 
times with larvae. 

Aceraius borneanus, Kaup. 

PI. xii, figs. 25-256. 
Regd. No. "f„' Borneo Sarawak Museum. 

The Berlin collections include specimens from Deli, Tebing tinggi, Peinan, 
Sipora, Hili Zabobo, Batavia, Bandjermasia, Tandjong, Kina Balu, c. 5000 ft., Sampit 

' Present throughout in all the three Rangoon specimens, but none of the f enasserim ones I have 
seen. The character seems to be so often variable, however, that I do not feel justified in establishing a 
separate variety for the former. All these punctures are usually, as Kuwert says, absent in the Teuas- 
serim form, but even in specimens found within a few hundred yards of such typical ones they may be 
p reseni at the posterior end exactly as in .1. lavoyanus. 

igi4.] F. H. Gravely : An AccouM of the Oriental Pinsalidac. 239 

and South Palawan. The Hamburg collection contains also specimens from Indra- 
pura, Langkat, and Banguey. The British Museum collection contains specimens 
from Perak. 

Description.— Length 2^-2^ mm. Anterior margin of labriim concave, left angle 
distinctly more prominent than right, sides parallel. Lamella of fifth joint of antennae 
usually rather long and often pilose. Mandibles with upper tooth well developed, 
denned on upper margin behind by a concavity which is much more marked on the 
left side than on the right; upper two terminal teeth large, the lowest one small on 
the left side, minute on the right; left anterior lower tooth rather large, triangular 
above and below, grooved horizontally between these two parallel surfaces ; its counter- 
part on the right side minute. Mentum punctured and hairy laterally ; middle portion 
smooth, with narrow depressed anterior border. Head above punctured and hairy in 
neighbourhood of long keeled parietal ridges, which extend vertically outwards from 
central tubercle, as is more or less invariably the case in all species of the genus I have 
seen, such variations as occur being always towards a backward direction; frontal 
grooves distinct, uniformly curved towards inner tubercles, or straight, or even 
curved outwards to terminate a little way behind them ; frontal area consequently 
variable in form; ridge joining inner tubercles almost straight, or faintly convex 
in middle; left outer tubercle long and slender, more or less inwardly directed, often 
much stouter at base than distally, variable in detail ; right outer tubercle moder- 
ately long, apex acute in fresh specimens as seen from above, truncate or imperfectly 
bifid in profile; anterior angles of head not prominent, containing at least 120°; 
apical angle of supra-orbital ridge .almost equally blunt as a whole, but strongly 
peaked. Pronotiim almost always with more or less distinct median groove, 
marginal grooves incomplete in middle before and behind; scars crescentic, they 
and the whole lateral border in front of them strongly and often closely punc- 
tured. Lateral plates of under side of prothorax sometimes slightly punctured and 
hairy in extreme anterior angles; then smooth for a space, behind which the 
surface is strongly punctured and hairy ; hairless behind the co.N;ae. ScHtclliim highly 
polished ; mesothoracic episterna punctured above and along anterior margin, somewhat 
dull in posterior angle; mesosternum finely roughened throughout or in part, scars 
extremely variable, never broad, sometimes long, sometimes rudimentary. Lateral 
areas of metasternum very sharply defined and narrow in front, very broad and 
indistinctly separated from intermediate areas -behind, not in contact with central 
area; anterior intermediate and lateral areas closely and rather finely punctured 
throughout, punctures becoming coarser and somewhat striate behind; posterior 
intermediate areas punctured along posterior border, but as a rul.e not further for- 
wards; central area smooth and poUshed, as is also ridge separating lateral from ante- 
rior intermediate areas. Posterior parts of hind coxae smooth or nearly so. Abdomi- 
nal s^ernrt indistinctly punctured in scars, the second with a few indistinct, but 
larger, hair-bearing punctures as well in fresh specimens. Elytra hairy, closely punc- 
tured at sides over whole surface from seventh rib onwards. 

240 Mcnwiy^ nf fhc Induin Mx^cum. [Vol.. Ill, 

Aceraius minor, n. sp. 

Text-fi,!>. 4I), p. 234. 

Described from five specimens from Taiping, 4oo()-50()() ft., in the Kgl Zool. 
Museum in Berlin. 

Description. — 2j 25-5 mm. Resembles A. borncanits except in the following 
respects. Upper tooth of mandibles obsolete, the upper margin practically- straight; 
lowest terminal tooth of left mandible well developed, equal to that of right ; ante- 
rior lower tooth of left mandible also well developed, but much smaller than that of 
right which, as is usual in this genus, is very large. Left outer tubercle of head shorter 
and somewhat stouter, inner angle of distal truncation scarcely developed, the 
tubercle as a whole consequently directed more forwards ; right outer tubercle some- 
what variable in both species, but as a rule rather better developed in this one 
than in the preceding. Abdominal sterna somewhat more distinctly punctured in the 
scars; elytra with hair-bearing punctures on the seventh and ninth ribs only, the 
whole of the eighth and tenth smooth and hairless. 

Aceraius aequidens, n. sp. 

Text-fig. 4E, p. 234. 

Described from one specimen from Kina Balu in the collection of the Deutsches 
Entomologisches Museum. 

Description. — Length 29 mm. Differs from the preceding in the following charac- 
ters only: anterior lower tooth of left mandible not very large, scarcely larger than 
that of right mandible ; men turn somewhat as in A . wallacei, anterior margin depressed 
on either side of the middle line only, in the single specimen that I have seen ; inner 
angle of distal truncation of left outer tubercle of head better developed ; right outer 
tubercle more as in A. borneanus; abdominal sterna also as in that species; seventh 
to tenth ribs (inclusive) of elytra with hair-bearing punctures, but with scarcely as 
many as in .1 . borneanns. 


Genus MACROLINUS, Kaup. 

The form of the frontal area of the head, the lengths of the lamellae of the 
antennae and the presence or absence of densely punctured areas on the pronotum, all 
of them characters which have been found apt to be of little value, and often actually 
misleading, for taxonomic purposes in other genera, have usually proved to be 
constant throughout each of our extensive series of specimens of different species 
of this genus, and to provide the best means of distinguisliiiig one species from 

I9I4-] F- H. Gravely : An Account of the Oriental Passalidae. 241 

Macrolinus nicobaricus, n. sp. 
PI. xiii, figs. 40-40^. 

9092-9 9101 9277 

r 1 1^1 

0279-81 51M 8130^2 

Regd.NoJ ' J 1* ■ Nicobar Islands r F. Stoliczka, 

TT -^{iyPe); 

\ F. A. de Roepstorff. 

«ip Rangoon' ? 

Description.— Length 30-36 mm. Labrum transverse; symmetrical or with left 
angle slightly more prominent than right; sides approximately parallel, angles 
rounded, anterior margin straight or slightly excavate. Lamellae of antennae of 
moderate length, the three proximal ones distinctly shorter and stouter than 
the three distal ones. Mandibles with upper tooth distinct ; ventral terminal tooth 
smaller and set further back than the other two; anterior lower tooth simple, 
conical, acute, that on the right side very little smaller than that on the left. 
Mentum punctured and sparsely hairy along posterior border between inner limits 
of scars, but not behind their posterior limits (i.e. not between either scar and 
posterior margin) ; this puncturing rarely extended forwards on to remaining part of 
surface of median area to any extent. Head closely and somewhat rugosely punctured, 
clothed with fine hair ; parietal ridges extending straight outwards or very slightly 
backwards from central tubercle, from which they are usually separated ; frontal ridges 
enclosing an acute angle, then curving rapidly outwards and then again forwards; 
inner tubercles joined to one another by a curved ridge whose concave side faces 
forwards; these tubercles joined to. outer tubercles by ridges enclosing a smooth and 
unpunctured area ; outer tubercles truncate or imperfectly forked at apex in profile ; 
anterior angles of head containing more than 120°; canthus rounded externally; 
posterior rounded portion of supra-orbital ridge meeting outer wall of hollow anterior 
portion in angle of not more than 120°, inner wall of anterior portion usually as 
distinct as outer wall. Pronotum with marginal grooves incomplete across the 
middle before and behind ; median groove absent or scarcely distinguishable ; a small 
depression present as a rule near anterior angles, usually containing (sometimes 
replaced by) two punctures (very seldom a small cluster) which often coalesce to 
form a short obHque groove; scars rounded, or linear and oblique, punctured, hairy; 
surface between scars and marginal grooves likewise punctured and hairy. Lateral 
plates of lower side of prothorax smooth towards inner side both in front of and 
behind coxae ; outer border punctured and hairy, especially behind. An indistinct 
median groove sometimes present on scutellum; mesothoracic episterna punctured 
except in posterior angle and along ventral border, punctured area bordered by a 
roughened band, the rest polished. Mesosternnm with usual anterior marginal striato- 
punctate area often followed by a few coarse punctures ; scars strongly impressed, 
their surface finely roughened, this roughness being continued more or less over 
the outer angles of the plate ; central area more or less poHshed with exception of a 

1 I find by looking up the register that this record is of doubtful authenticity ; it appears to be the 
only record of the species outside the Nicobars and is probably incorrect. 


Memoirs of the Indian Museum. 

[Vol. Ill, 

pair of circular areas which are finely rugose or punctured. Metasternum with lateral 
areas narrow, finely punctured and hairy throughout, not in contact with central area ; 
anterior intermediate areas punctured and hairy; posterior intermediate areas smooth 
and hairless. Posterior parts of hind coxae, and scars of abdominal sterna, finely 
punctured, either sparseh^ or closely. Elytra slightly hairy at the shoulders; very 
strongly punctured in the lateral grooves, more weakly in the dorsal ones of whicli 
the innermost are scarcely punctured at all. 

Macrolinus andamanensis (Stoliczka). 
PI. xiii, figs. 4i-4i(/. 
" — [paratypes) 

Regd. No. 

^' (paratypes) 

'■"^ (paratypes) 

UHiii-7o 4072- ^ 37i:i-; 
ft S III 

660^-8 6428-9 9860 
I 13 14 IS 


.i.ViKlainan Islands 

Stoliczka bequest, J. Wood- 
Mason, etc. 

Museum Collector. 

*i^°"- appear to be paratypes. Nos. ^\^^ and ^^f-^ are from " Dr Stoliczka's 
Type collection," and the rest were "named by Dr. Stoliczka and Mr. J. Wood- 
Mason " ; several of the latter series, including some from Wood-Mason's collection, 
bear labels written by Stoliczka. 

Description. — Length 31-38 mm. Differs from the preceding in the following 
characters only ; lamelliform processes of six distal antenna-joints somewhat longer 
and slenderer; ridge joining inner tubercles of head'- straight or slightly raised in 
middle line; frontal area in consequence of this and of a difference, less constant in 
both species, in the course of the frontal ridges (which in this spacies are usually curved 
so that they do not enclose an acute angle in front of the central tubercle), longer in 
proportion to its width, and semicircular rather than crescentic in form ; outer tubercles 
bifid in profile, upper process longer (often much longer) than lower, and separated from 
it by a distance nearly as great as that from inner tubercle ; anterior and posterior 
portions of supra-orbital ridges meeting in an angle of not less than 120°; mesosternum. 
more extensively polished, circular punctured areas mostly represented by smooth 
depressions or entirely absent. In the mesosternum of some specimens there is a 
ver>' strongly marked median groove whose surface is finely roughened, but more 
usually this is obscure or absent; I have not seen it in /?. nicoharicus, but it may 
very likely be found occasionally in that species also. 

' 1 find by looking up the register that this record is of douljtlul authenticity, so as it appears to 
be the only record of the species outside the Andamans it is probably incorrect- 

''■ No mention is made here of the central tubercle, as this is so very variable in both species, both in 
form and in the extent of its connection with the posterior ridges, that I can find no constant diflference 
between the two that is sufficiently definite to permit of description. No taxonomic importance is to be 
attached to the differences that appear in the figures. 


F. H. Gravely : An. Account of the Oriental Passalidae. 




706 ti(Oo-( 

98S9-90 986-1 

e:i9(l 6«2 6+53 

Macrolinus sikkimensis (Stoliczka). 
I. M. SIKKIMENSIS, Stoliczka, s. str. 
PI. xiii, figs. 42-42rt. 
'Sikkim" ' 
Khasi Hills 

Harmutti, base of Dafla Hills 
Dikrang Valley 
Dafla Expedition 
Camp 9, Dafla Expedition 
Kobe, Abor Country, 400 ft. 
Janakmukh, Abor Country, 600 ft. 
Rotung, Abor Country, 1400 ft. 
Naga Hills 

F. Stoliczka. 
Shillong Museum. 
J. Wood-Mason. 


, H. H. Godwin- Austen. 


vS. W. Kemp. 

Description. — Length 25-31 mm. Resembles the two preceding species except in the 
following respects. Parietal ridges of head on the whole more distinctly connected with 
central tubercle ; frontal ridges as in M. nicoharicus ; inner tubercles as in M. nicobari- 
cus, the ridge between them as in M. andamanensis ; outer margin of inner tubercles 
often produced backwards as more or less distinct ridge over anterior part of head ; 
posterior part of supra-orbital ridge meeting anterior part in angle of about 120°. 
Pronotum strongly punctured laterally behind anterior extremities of marginal groove, 
all along this groove at the sides, and usually near the scars also. Mesosternum usually 
more poHshed than in M. nicobaricus : with or without a single median roughened or 
finely punctured area of varying extent in place of the paired areas; a more or less 
obscure groove sometimes present in the middle line. Metastermim coarsely punctured 
in posterior intermediate areas, otherwise as in the preceding species. Elytra slightly 
hairy at shoulders, their dorsal grooves more or less distinctly punctured; lateral 
grooves very wide, with strong punctures elongated to form a series of transverse 

Habits. — Mr. Kemp did not find any large colonies of this species during the Abor 
Expedition. This agrees with the results of my own more scanty observations on the 
sub-species found in the Dawna Hills, and I am led to conclude that the species is 
not a markedly gregarious one. The sub-species tavoyanus seems to live in logs that 
have reached too advanced a state of decay to be suitable for Aceraius to inhabit; 
whether this is true of the northern form also has yet to be determined. 

2. M. SIKKIMENSIS, SUbsp. TAVOYANUS, n. subsp. 

Regd. No. 



• Tavoy 

Museum Collector. 

' Stoliczka tells us in his paper (1873, p. 162) that he collected this specimen "at about 1500 feet, 
some two miles east of Pankabari." Pankabari is many miles south of the Sikkim frontier of to-day. 


Memoirs of the Indian Mmeitm. 

[\ OL. Ill 

Regd. No. 


Between Misty Hollow and Sukli, 
Dawna Hills, 2100-2500 ft. 

I'. H Gravely. 



Between Thinganiiyinaung and Sukli, 
Dawna Hills, 900-2100 ft. 

F. H. (iravely 



Dawna Hills, Tenasserira 

F. H. Gravely. 

I have also seen specimens from Carin Cheba, 3(XJ0-J5()C) and 4<)oo-50()() ft., in 
the collection of the Deutsches Entomologisches Museum . 

Description.— Length 29-5-33-5 mm. Differs from .1/. sikkimensis, s. sty., only in 
having the mesosternal scars coarsely punctured partially or throughout. No trace of 
these punctures has been found in any Himalayan specimen with the exception of 
one from Kobo (regd. no. -rir") in which, although scarcely perceptible on one 
side, they are few but distinct on the other. This specimen may be regarded as transi- 
tional between the typical Himalayan form, and the form from the Dawna Hills which 
differs therefrom somewhat less markedly than does that from Tavoy. Probably, 
therefore, the change from the typical Himalayan form to the Tavoy form is a ejradual 
one and numbers of transitional forms may be expected to occur in the hills of 
north Burma. 

Macrolinus crenatipennis, Kuwert. 

Described from two specimens from Ceylon (one of them evidently the type, as 
it bears Kuwert's label), both in the collection of the Konigliches Entomologisches 
Museum in Berlin. 

Description.— Length 21-5 mm.' Differs from .1/. rc/nndi/rnns only in its smaller 
size, and in the extremely coansc puncturing of all the grooves of the t'/\'/;'(f , the 
dorsal grooves of .1/. crenatipennis being as coarsely punctured as the lateral ones of 
M. rotundi/rons. 

Macrolinus rotundifrons (Kaup). 
PI. xiii, lig. 43. 

Regd. No. -f,-^ Kandy, Ceylon \ ^- E- t^^een. 

< N. Anuandale. 

'-^■•■' Feradeniva, Ceylon \ ^- ^- ^reen. 

< F. H. Gravely. 
T'' Coylon F. H. Gravely. 

Mr. Green's collection includes also specimens from Galagedara, Central Province, 

Description.— Length 25-31 mm. Differs from M. sikkimensis in the following 
respects only. Median part of mentum unpunctured ; parietal ridges of head extend- 
ing straight outwards or slightly forwards, still more distinctly connected with 
central tubercle; ridge between inner tubercles strongly convex, its middle part 
sometimes almost confluent with ridge between outer tubercles. Pronotitm with 
strongly marked median groove; punctures along marginal grooves at sides confined 
to grooves themselves, although the other punctures are arranged as in T. sikkimensis. 

Kuwert says 26-27 mm. 

igi4.] F. H. Gravely : Aii Account of the Oriental Passalidae. 245 

Mesosternitm smooth and polished throughout, a few strong punctures sometimes 
present along inner margin of each scar. Elytra with lateral grooves of normal 
width, strongly punctured, but not transversely grooved. 

Habits. — This species is very common in the Royal Botanical Gardens at Pera- 
deniya, where it lives among the piles of dead leaves, etc., that are put to rot away 
on the river bank. 

Macrolinus waterhousei (Kaup). 
PI. xiii, fig. 44. 
Regd. No. '^' Bulutota, Ratnapura Dist., Ceylon Colombo Museum. 

Description. — Length 33-35 mm. Differs from M. rotundi/rons in the following 
characters only. Median part of mentum sparsely punctured. Ridge between inner 
tubercles of head scarcely convex, almost straight, so closely approximated to ridge 
between outer tubercles throughout its length as almost to conceal it (differing in 
this respect from M. andamanensis). Pronotiim unpunctured but (? always) with 
a more or less obscure depression in place of the strongly punctured area behind 
anterior extremities of marginal groove. Scars of mssosternum indistinct .or absent. 

Macrolinus weberi, Kaup. 

Described from a single specimen from the Philippines in the collection of the 
Konigliches Zoologisches Museum in Berlin. 

Description. — Length 25 mm. Differs from the following species only in having 
the first two lamellae of the antennae, somewhat shorter, and the anterior face of the 
central tubercle of the head almost vertical to the general surface. In the former 
respect, and in its locality, it agrees with Kaup's description of M. weberi, but the 
differences are very minute, and in the absence of other similar specimens I very 
much doubt whether it is really distinct. 

Macrolinus latipennis (Percheron). 

Regd. No. ~ Malacca ? 

'ip Johore J. Wood-Mason. 

There is one specimen from Borneo in the Sarawak Museum collection ; and the 
Berlin collections include specimens from Deli, Nias (Ombolata), Buitenzorg, S.-E. 
Borneo, and South Palawan. 

Description. — Length 22-5-27-5 mm. Labritm punctured and hairy, anterior 
margin straight, angles rounded. All six lamellae of antennae verj^ long and slender, 
their extremities arranged in a straight line when the antennae are furled. Upper 
tooth of mandibles small or obsolete ; lowest terminal tooth smaller than the two 
upper ones; anterior lower tooth conical, that of the right side slightly smaller and 
more acute than that of the left. Mentum smooth between the scars or with a few 
punctures near the middle-line. Head closely and finely punctured throughout above, 
or with the anterior half of an area in front of the short parietal ridges and outside 
the frontal ridges, and a small patch immediately behind the former, more or less 

-^4^ Monoirs of the Indian Museum. \\'Oi.. Ill, 

sparsely punctured or smooth ; the central tubercle is little or no steeper in front than 
behind, and the size of its apical angle is variable ; from the central tubercle a median 
keel extends forwards a short distance to the point of origin of the frontal ridges which, 
with the straight, convex or slightly W-shaped ridge joining the frontal tubercles, 
enclose a more or less semicircular area. Pronotnm with scarcely a trace of median 
groove ; marginal groove incomplete in middle before and behind ; a patch of punc- 
tures in anterior angles, scattered punctures present or absent between this and the 
curved punctured scars. Prothoiax beneath punctured and hairy at the sides Scutel- 
Itim smooth and polished ; mesothoracic episterna punctured above and along anterior 
margin ; mesostenium with a few hair-bearing punctures in front of an anterior polished 
area between the anterior ends of the scars, this smooth area usually continued 
backw^ards as an indistinct keel, or as a smooth and highly polished streak, to about 
the middle of a finely roughened area extending from the scars near their anterior 
extremities to the posterior margin; between this area and the posterior part of the 
scars the mesosternum is smooth and polished; the scars are long and almost always 
coarsely punctured, the punctures sometimes extending beyond the scars themselves 
into the lateral angles of the plate which they may completely fill. Metastenmm more 
or less densely punctured throughout the lateral and anterior intermediate areas, and 
often along the outer part of the posterior margin of the posterior intermediate areas 
and the ridges separating these from the lateral areas ; anterior and inner part of 
posterior intermediate areas rarely punctured ; the rest smooth and unpunctured ; 
lateral areas much broader behind than in front, but not in contact with central 
area. Hind coxae and abdominal sterna as in other members of the genus. Elytra 
slightly hairy at the shoulders; all grooves of equal width; lateral grooves little more 
coarsely punctured than those nearest the middle line. 



Parapelopides symmetricus, Zang. 

Text-fig. 5, A-B, p. 247. 

Described from specimens from Mt. Kina-Balu, c. 5000 ft., in the collections of 

the British Museum and the Deutsches Entomologisches Museum. 

Description.— Length 40-41-5 mm. Dift'ers from Trapezochilus nobilis in the fol- 
lowing characters only. Anterior lower tooth of right mandible small and obtuse, but 
distinct. Scars of mentum united to form a W-shaped figure, whose posterior trans- 
verse part is always distinct and deeply impressed ; anterior margin of mentum some- 
times with a slight concavity in the middle. Upper surface of head less strongly 
rugose ; outer tubercles narrower, their denticles ver>' obtuse, middle denticle rarely 
distinguishable. Only extreme lateral angles of scutellum unpunctured ; puncturing of 
posterior angles of mesothoracic episterna also more extensive. Whole of lateral 
margin of mesosternum bordered by a broad band of coarse punctures. Metastcrnum 
with at least a few punctures in the posterior intermediate areas. 

igi4.] F. H. Gravely : An Account of the Oriental Passalidae. 247 


Trapezochilus nobilis (Kmvert)'. 

PI. xiii, fig. 48; text-fig. 5, C-D. 

Regd. No. ~^' "!'* Tavoy Museum Collector. 

I have also seen a series of specimens from Taiping, 4000-5000 ft., in the collec- 
tion of the Konigliches Zoologisches Museum in Berlin ; and two specimens from 
Perak in the British Museum. 

Description. — Length 35-41 mm. Labrum hairy and punctured ; anterior margin 
straight or slightly concave, with a more or less indistinct denticle in the middle; 
angles rounded, symmetrical or very nearly so. Antennae with a more or less 
indistinct lamelliform process on the fifth joint, lamellae of subsequent joints of 
gradually increasing size, all short and stout. Upper tooth of mandibles usually 
rectangular sometimes blunter, minute; terminal teeth large, the two upper ones 
broadest in the vertical plane, the lowest one horizontal ; anterior lower tooth distinct 
and sharply pointed on the left side, obsolete or minute and fused with base of 


A. Parnpelopides symmeiricus, auterior margin of head x 4. 

B. ,, ,, right mandible x 4. 

C. Trapezochilus nobilis, anterior margin of head x 4. 
D ,, ,, right mandible x 4. 

E. Trapezochilus rcspectabilis, anterior margin of head x 4. 

lowest terminal tooth, on the right. Mentum punctured laterally, smooth between 
and behind the oblique groove-like scars. Head strongly rugose in front, smoother 
behind, the surface polished throughout. Outer tubercles somewhat variable, 
approximately symmetrical, each composed of three conical denticles of which the 
innermost is usually the largest ; the outermost is usually slenderer or quite short 
and blunt ; the middle one is small and set a little further back, its position is 
variable, and may be different on opposite sides of the same specimen, it is some- 
times obsolete on the right or on both sides. Anterior angles of head containing a 
little more than 120° ; canthus rounded externally. Parietal ridges curved slightly 
forwards. Supra-orbital ridges smoothly convex behind, excavate in front, their 
upper margin curved (not angular) in front when seen in profile. Pronotum 
smooth and polished, not obviously punctured except in the scars and margi- 
nal grooves; marginal grooves widely discontinuous in front, almost continuous 
behind ; median groove rudimentary or absent. Lateral plates 01 lower surface 

' This species has proved not to be distinct from the next ; and the names applied to them here 
must both be discarded in favour of dorsalis, Kaup (see below, pp. 297-8) 

248 Memoirs aj the Indian Museum. [Vol. Ill, 

of prothorax smooth and hairless in front, punctured and very hairy further 
back, often rugose behind the coxae. Scutellum traversed by a pair of broad and 
closely apposed hairy and finely punctured bands which are sometimes fused in the 
middle line ; mesothoracic episterna coarsely punctured above and in front , more finely 
in the posterior angles which are hairy ; a small smooth area between the two punc- 
tured ones. Mesosternum often more or less dull and rugose, usually with a patch 
of hair-bearing punctures in the lateral angles ; scars indistinct or absent. Metaster- 
num with lateral areas very broad behind, but not in contact with central area ; 
lateral and anterior intermediate areas hairy and rather finely punctured, posterior 
intermediate and central areas smooth and hairless. Posterior parts of hind coxae 
slightly roughened ; scars of abdominal sterna smooth or very nearly so. Elytra -with 
lateral grooves broader than dorsal ones, the punctures in the former forming rows of 
short transverse grooves. 

Trapezochilus respectabilis (Kuwert) '. 
Text-fig. 5 E, p. 247. 
Described from a series of specimens from Deli, vSumatra, in the collection of the 
Deutsches Entomologisches Museum. 

Text-fig. 6. 

Giiaphalncveiiiis sijjiph'x. n. sp., head x 4. 

Description. — Length 33-38 mm. Differs from preceding species in following 
characters only. Scars of mentum sometimes more or less distinctly united behind to 
form a W-shaped figure as in Parapelopides. Upper surface of head less strongly 
rugose; outer tubercles more prominent, their middle denticle usually obsolete, the 
inner one always broad and distally truncate in unworn specimens. Mesosternum 
rarely punctured in lateral angles. Posterior intermediate areas of met aster nam 
sparsely punctured in one specimen. 

Gnaphalocnemis simplex, n. s]). 
Text-fig. 6. 
Described from a .single sjiecimen from Perak in the British Mu.seum collection. 
Description. — Length 31 mm. Head, with its appendages, as in (l. monticulosus , 

' See foot-note on previous page. 

igi4.] F- H. Gravely : An Account of the Oriental Passalidae. 249 

except the anterior margin ; this is flat or slightly convex on both sides ; the left 
outer tubercle is simple and truncate ; the right one is similar but broader, its trun- 
cation somewhat concave ; two smaller pointed denticles are developed between this 
tubercle and the right anterior angle of the head. Prothorax, mesothorax and meta- 
thorax as in Trapezochilus nobilis. Posterior parts of hind coxae rough. Abdominal 
sterna poHshed. Grooves nearest inner margin of elytra scarcely punctured, next 
pair somewhat more distinctly punctured, puncturing of lateral grooves very strong 
— quite as strong as in G. hnrmeisteri. 

Gnaphalocnemis burmeisteri (Kaup). 
PI. xiii, fig. 49. 

Described from specimens from Deli, Pedong, Peinan, Soerian Plantation (nr. 
Solok), Battak Mountain, Kepahiang, Redjang-I.ebong, and Java (the type and two 
other specimens) in the Berlin collections ; specimens from Deli and Somgei Lalah 
in the Hamburg collection ; and a specimen from Sarawak in the Sarawak Museum 

Description.— Length 41-51 mm. Differs from the following species only in 
having grooves 5-7 of the elytra narrower, simply punctured without any defined 
flattened surface, transverse punctures, or longitudinal roughened lines. 

Gnaphalocnemis monticulosus (Smith). 
PI. xiii, fig. 49(7. 
I have not seen the specimen which Stoliczka says he received from Svimatra ; 
the specimen which he collected in' Penang, though not referred to in his paper, is 
evidently of this species and appears to have been identified by him. 

Regd. No. '■'■" Penang F- Stoliczka. 

I have also examined specimens from Kuching and Sadong belonging to the 
Sarawak Museum; specimens from Tengah Mountain, Deli, Tandjong Morawa, Solok, 
Medan, Sarawak and Mt. Kina Balu in the collections in Berlin; and a specimen 
from Lampung in the British Museum. 

Description.— Length 38-44 mm. Anterior margin of lahrum shghtly concave, 
sometimes almost straight, usually with a distinct denticle in the middle ; angles 
rounded, the left inclined to be slightly more prominent than the right. Upper 
margin of mandibles usually more or less convex or angular behind upper tooth; 
upper tooth distinct, more or less rectangular on left side, usually acute on right ; 
no trace of right anterior lower tooth present ; left anterior lower tooth and left lowest 
terminal tooth more or less fused at base, each distinct and well developed at apex. 
Mentum usually somewhat narrower and more convex between the scars than in 
Trapezochilus dorsalis (^nobilis). Head polished throughout, more or less rugose in 
front of parietal ridges. Outer tubercles complex, somewhat variable in form (compare 
figs. 49 and 49rt '). Anterior border excavate between left outer tubercle and front of 

I Prepared from specimens belonging to two different species ; but the structure and range of varia- 
tion of the outer tubercles appears to be the same in both, and also in the larger G. tridens (fig. 50). 

250 Memoirs of ihc Indian Museum. [Voi,. Ill, 

left supra-orbital ridge. Anterior angles of head containing about 120°, canthus 
rounded externallj' ; supra-orbital ridge as in Trapezochilus dorsalis ; parietal ridges 
directed slightly forwards. Prothorax and mesothorax much as in Trapezochilus 
dorsalis, but with the median groove of the pronotum often more distinct, and a pair 
of distinct large oval scars on the mesosternum. Metasternum with lateral areas very 
broad throughout, almost as broad in front as behind, but not in contact with central 
area ; central area unpunctured ; lateral areas finely punctured and hairy ; anterior 
intermediate areas hairy and a little more coarsely punctured in anterior angles, 
smooth behind ; posterior intermediate areas ver}' coarsely but often sparsely punc- 
tured, not hairy. Hind coxae usually smooth. Second abdominal sternum \X'-i\xa\\y 
punctured .li front of scars; abdominal sterna otherwise smooth. Elytra with grooves 
5-7 more or less broad, their flattened polished surface usually more or less completeh- 
bordered on either side by a roughened line, with which the punctures are often con- 
fluent, the punctures coarse and as a rule distinctly transverse. 

Gnaphalocnemis tridens (Wied.). 
PI. xiii, figs. 5()-50(?. 
Regd No. '^' Java J. Wood-Masou. 

I have also seen one specimen from the island of Madura which is the propert}^ 
of Mr. H. E. Andrewes; specimens from Sumatra, Tjibodas c. 5000 ft., Tengger 
Mountain, and Soekaboemi in the Berlin collections; and specimens from Preanger 
4-6000 ft., Buitenzorg, and East Java as well as Wiedemann's tj^pes in the Hamburg 

Description.— Length 45-56 mm. Differs from the preceding in the following 
characters only : upper tooth on both sides often more obtus3, the mandibles less 
(often not at all) convex above behind it ; surface of head less rugose ; parietal ridges 
usually extending somewhat more directly outwards; all but one or two of the punc- 
tures of posterior intermediate areas of metasternum usually confluent and concen- 
trated along the inner margin of this area ; posterior part of hind coxae more or less 
strongly punctured; grooves 5-7 and posterior part of groove 8 of elytra very broad, 
each with roughened margins and a flat polished surface marked by a single row of 

Genus GONATAS, Kaup. 

Gonatas germari, Kaup. 

PI. xiii, figs. 47-47(f. 

Regd. No. 'j:;'' Java W. S. Atkinson. 

Description. — Length 265 mm. Labrum hairy and punctured, anterior margin 
with a deep semi-circular excavation, angles rounded and prominent, sides straight 
and parallel. Antennal lamellae long, six in number. Left mandible somewhat larger 
than right ; upper tooth small on both mandibles, rectangular on left, more obtuse 
on right ; two upper terminal teeth partly fased, lower one distinct, meeting them in 
an angle of about 60°; anterior lower tooth rudimentary on left, large but blunt on 

I9t4.j F. H. GRAVEiyY : An Account of the Oriental Passalidae. 251 

right mandible. Mentum punctured laterally only. Head crossed by band of large 
sparse punctures immediately in front of parietal ridges ; parietal ridges extending 
directly outwards on either side of central tubercle ; frontal ridges extending at first 
more outwards than forwards from central tubercle, then bent abruptly towards inner 
tubercles; ridge between these slightly produced forwards in middle line, which is 
marked in frontal area by an indistinct ridge ; left outer tubercle broad, parallel-sided, 
truncate distally, the extremity being slightly concave as seen from above, straight and 
perpendicular from the side ; right outer tubercle of equal length but more slender, sides 
as seen from above abruptly converging near end to meet each other in a right angle, 
upper surface longer than lower, the process being obHquely truncate as seen from the 
side ; anterior angle of head containing about 100° ; canthus almost equilaterally 
quadrangular. Pronotum with strong median groove, marginal groove incomplete in 
middle in front, obsolete in middle behind. Prothorax below poHshed; hairy and 
indistinctly punctured close to margin in front, more distinctly punctured behind. 
Scutellum smooth and polished ; mesothoracic episterna polished throughout, punctured 
above and along anterior margin ; mesosternmn smooth and polished, with small deep 
scars. Metasternum finely punctured in lateral areas, which are very little wider 
behind than in front and are not in contact with central area ; more strongly punc- 
tured in anterior intermediate areas ; coarsely punctured in posterior intermediate 
areas. ' Hind coxae and abdominal sterna polished, almost unpunctured. Elytra finely 
punctured in dorsal grooves, more coarsely in lateral ones. 

Genus LEPTAULAX, Kaup. 
Leptaulax beccarii, Kuwert. 
Described from two specimens from Sumatra in the Berlin collections, one of 
them determined by Kuwert. 

Description. — Length 21 mm Differs from L. humerosus in the following char- 
acters only. PVontal area of head shorter and broader. Pronotum and abdominal 
sterna more as in Z,. dentatus, the latter not extensively punctured. Posterior parts 
of hind coxae smooth. Sculpturing of lateral grooves of elytra reduced (to a variable 
extent) in a somewhat different way, a polished tubercle being more or less distinctly 
developed from the middle part of each of the partly obsolete transverse ridges. 

Leptaulax humerosus, Kuwert. 
PI. xiii, fig. 51. 
Regd. No. ''^YJJ^' Ten miles south of Kuchiug, Sarawak C. W. Beebe. 

There is a single specimen from Kuching in the Sarawak Museum collection and 
another from the same locality in Mr. Andrewes' collection. I have also examined 
specimens from Perak, Deli, W. Sumatra, Ardjoeno, and Tengger Mountain in the 
collection of the Deutsches Entomologisches Museum. 

Description— Length 15-20 mm. Differs from the common allied species, L,. 

Mcmoits of the Indian Museum. 

[Vol. Ill, 

(ientatus, in the following characters only. Upper tooth of mandibles, usually weaker, 
anterior lower tooth simple. Inner tubercles of head perhaps a little inore widely 
separated ' ; frontal area almost always about as long as broad, semilunar or almost 
quadrangular in shape ; the two frontal ridges more or less evenly continuous with 
one another and joined to central tubercle by a single sharply keeled ridge— not 
curving round to meet in an acute angle close to apex of central tubercle as is usually 
the case in L. dentatus. Scars of mesosternum dull, their surface evenly continuous 
with a dull band extending along whole remaining length of outer margin. Lateral 
areas of metastcrnum much broader behind than in front, about equal in their widest 
part to minimum width of intermediate areas. Abdomen bordered beneath by a com- 
plete broad band of fine but dense puncturing which includes the whole of the 
terminal segment with the exception, in some specimens, ot a small smooth spot near 
the posterior margin. Depressed surface of two or three outermost grooves of elytra 
dull, the sculpturing somewhat indistinct and worn-looking, especially at the ends; 
the ninth and tenth ribs narrow throughout. 

Rcgd. No. 

It;50 i(!52 ViiV 

.w"7-»io:t ;t2Cii - 
i» i» 



Leptaulax dentatus (Fabr.). 

I. L. DENTATUS, Fabr. s. str. 

PI. xiii, figs. 52-52^^. 

Near Vizagapatam 

La-ai Valley 

Kobo, 400 it., Abor Country ^ 

Jauaktmikh, 600 ft., Abor Country 
Rotung, 1300-1400 ft., Abor Country 
Upper Rotuug, 2000 ft., Abor Country 
Renging to Rotung 261,0 ft. Aljor 

Kalek, 3800 ft., Abor Country 
N.-E. Frontier 

Museunt Collector". 
D. G. A. Pritchard and 
J. R. Waterfield. 

S. VV. Kemp. 

»' 19 

Sibsagar, Assam 

S. E. Peal and Museun 

" HI 

Dafla Expedition 

' U iT 19 

U 1!) 

Burroi, base of Da.<ia Hills 
Harmutti, base of Dafla Hills 

H. H. Godwin-Austen. 

(JH7-20 ««7-74 

u 19 

Dikrang Valley, Northern Frontier i 


9505 Mil 
" 1 1 » 

Dunsiri V^illey, Assam ^ 
Durrang, Assam ) 

eil5-8 963« 
" U IB 

I H. H. Godvviu-Austeu. 

' This remark, since it relates to a character which is somewhat variable in both forms, applies only 
to the species as a whole 

* Larvae, pupae, and adults in spirit. 


F. H. Gravely : An Account of the Oriental Passalidac. 


Regd. No. 

9135-6 9367-73 

, 91.1(1-63 9165-; 

vStoliczka bequest, 
Dr. Cameron. 

9375-7 9iS:-9 9491 

I 9169 -^' 

861-8711 3272^ 

Darjeeliiig (Sikkim in register) 


Stoiiczka bequest, etc. 

r Stoiiczka bequest, 
\ Asiatic Soc.'s collection, 
( E. T. Atkinson, ?. 
J. Wood-Masou. 

^ Stoiiczka Ijequcst, 
\ S. Kurz. 

F. H. Gravely. 

2913 872-96 89!, 
T 19 19 

5S-5-6 5879-85 


Kawkareik, Tenassenm, c. 300 ft. 

Misty Hollow, W. side of Dawna Hills, F. H. Gravely. 

Tenasserim, c. 2200 ft. 
Between Misty Hollow and Sukli, top of F. H. Gravely. 

Dawna Hills, 2100-2500 ft. 
Between Thingannyinaung and Sukli, F. H. Gravely. 

E. side of Dawna Hills, QOO-2100 ft. 
Tenasserim J- Wood-Mason. 

897 ■S99-'.'4t ( 
19 19 ' 

Upper Tenasserim 



Johore, Malay Peninsula 

Perak, Malay Peniusuk 

J. Wood-Mason. 

Museum Collector. 

( Stoiiczka bequest, 
\ Dr. W. Abbay. 
t ? 

C J. Wood-Mason. 

L ? INIuseum Collector. 
Moti Ram. 
H. E. Andrewes. 

I Specimens selected to show the full range of continuous variation from a series of 47 specimens 
taken from a single colony. No. ''J^" is from the same colony. 
* Adults and larvae in spirit. 

254 Memoirs of the Indian Museum. [Vol. Ill, 

I have also seeu specinieus from Buxa (Duars) in Mr. Audrewes' collection ; one 
from Silonbari at the base of the Lakhimpur hills in Mr. Stevens' collection ; and 
specimens in the Dehra Dmi collection from Kochugarh, Goalpara District, 14-V-06; 
Kapti, Chittagong Hill Tracts; and Chaduar, Assam, 9-iv-o6. In the last-named 
collection there is also a specimen said to be from Mussoorie, but as the label also bears 
the name Lucanus lunifer it has probably been transferred from another insect. 
This somewhat improbable locality must, therefore, be ignored. 

Description — Length 195-37 mm. Labriim punctured and hairy, anterior margin 
more or less concave, angles more or less rounded or acute ; sides strongly convex or 
almost straight, their general direction parallel, or slightly convergent behind. Man- 
dibles strongly angular on outer side; upper tooth well developed ; lowest terminal tooth 
somewhat smaller than and posterior to the other two; anterior lower tooth double, 
its anterior part long and sharply pointed in unworn specimens, the posterior part 
extending backwards as a narrow and more or less bilobed ridge, less completely 
separated from the former on the right side than on the left, and much more extensive 
in large than in small specimens— in the latter it may be obsolete. Mentum with some 
large punctures outside, and in front of, a pair of V-shaped, crescentic, or almost cir- 
cular scars. Head distinctly marked with round punctures which may be replaced 
in the anterior half by an obscure longitudinal rugosity ; frontal area very variable 
in form (compare figs. 526-c), usually traversed by a more or less distinct median 
ridge ; distance between two inner tubercles on an average about equal to that 
between these tubercles and the outer ones, the latter being situated about half way 
between the former and the anterior angles of the head; the exact proportions 
between these distances depends on the shape of the frontal area ; even when this is 
at its broadest, the distance between the inner tubercles is scarcely twice as great as 
that between the inner and outer tubercles ; median tubercle distinct, obscure, or 
obsolete ; end of canthus narrow, rounded or abruptly truncate, prominent in large 
specimens but as a rule not in small ones; parietal ridges short, not extending to the 
supra-orbital ridges. Pronotum smooth and highly polished above; thickly punc- 
tured at the sides, especially in the anterior angles and the neighbourhood of the scars ; 
median groove strongly developed. Lateral plates of lower side of prothorax more or 
less punctured in front of the coxae, unpunctured behind. Prosternum smooth and 
highly polished between and behind coxae, keel sometimes strongly grooved trans- 
versely in front, anterior median semicircular area dull and radially grooved, lateral 
areas finely punctured behind with a row of hair along their posterior margins. 
Scutellum with usual median hairy patch on anterior margin, otherwise smooth 
and highly polished ; mesothoracic episterna strongly punctured above and along 
anterior margin ; mesosternum smooth and highly polished, except in lateral angles 
which are dull, and in scars which are often finely roughened ; these scars do not 
extend to the posterior margin of the sternum; they are very sharply defined on 
the inner side, and often also (to a less degree) on the outer side, which does not 
always coincide with the outer margin of the sternum ; a few punctures sometimes 
present behind the .scars. Central area of metasternum polished, very sharply defined, 

igi4.] F. H. Gravely : An Account of the Oriental Passalidae. 255 

with or without a few symmetrically arranged depressions of greater or less distinct- 
ness, not in contact with lateral areas; lateral areas narrow, parallel sided or nearly 
so, more or less rugose ; intermediate areas strongly punctured. Posterior parts 
of hind coxae more or less roughened or punctured. Abdominal sterna finely punctured 
along anterior margin (often incompletely in the last two segments), and more 
strongly in a more or less triangular patch of very variable size (sometimes absent 
from last segment) on each side ; otherwise smooth and polished. Elytra polished 
throughout; the lateral grooves about as broad as the ridges between them, their 
punctures extended to form short transverse grooves ; the ninth rib broader than 
the eighth in part, the tenth broader throughout. 

Habits. — This species is gregarious, it hves under the bark of logs which have 
rotted till their outer parts have lost their strength. 

2. ly. dEntatus var. glabriventris. n. var. 

9 m 9JM0 9.il3-4 , 

„ , „ J 29U +nsn 8U4 f a , x, ( .StoHczka bcquest, J. Wood- 

Regd. No. < — -^ _ V Andaman Isles \ 1 'J 

3722-3 37-37 \ { Mason, ctc. 

\ 10 1(1 

'if Java 7 

^ H. E. Andrewes. 
■f-' Borneo ) 

— Philippine Isles East India Company's 


This variety is not a very distinct one, forms intermediate between it and 
the typical form being sometimes fo.und. It is represented in the Sarawak Museum 
collection by a specimen from Matang and another from Kuching — the former is 
of about the same size as the Andamans and Philippine specimens in the Indian 
Museum collection (245-29 mm.), but the latter is much smaller than any other 
specimen I have seen (21 mm.). Mr. Andrewes has sent me for examination specimens 
from Borneo and Java. 

Description. — Differs from the typical form only in having the frontal ridges of the 
head normally rising from a short anterior prolongation of the central tubercle ; and 
especially in the smaller size of the triangular punctured areas of the abdominal 
sterna, those of the last segment having almost always disappeared. 

Leptaulax cyclotaenius, Kuwert. 
I. Iv. CYCLOTAENIUS, Kuwert, s. str. 
PI. xiii, fig. 53. 
Regd. No. '^ ^^ Johore J. Wood-Mason, ? 

vSinkep I. Moti Ram. 


2613 „ 

,, -^ Penrissen 

Description. — Length 14-16 mm. A somewhat smaller and much flatter insect 
than the following subspecies, from which it further differs in the following points 
only: parietal ridges of head perhaps a little longer; mesosternnm finely roughened 




vSarawak Museum. 

256 Memoirs of the Indian Mnst'um. [Voi, III, 

along outer margin, otherwise highly polished with fine puncturing in middle and 
usually behind scars also; intermediate areas of metasternum as in L. dentatus. 


R^g<3-^'"- ^ Dikrang Valley ^ H. H Godwin- Austen. 

^■*"^-^"'« DaOa Expedition ^ 

■";';;•'* Upper Renging, 2150 ft. ) 

Description. — Length 15-17 mm. Differs from L. dentatus in the following respects 
only. Head and mandibles as in L. humerosns except that the parietal ridges are some- 
what longer and the frontal areas always semicircular. Mesosternum very variable ; 
rough or polished; in the latter case always,' in the former usually, with some large 
punctures. Central area of metasternum more or less strongly punctured; inter- 
mediate areas often somewhat broader in their narrowest part, with the outer margin 
frequently somewhat imperfectly defined. Posterior parts of hind coxae usually quite 
rough. Abdominal sterna usually more extensively punctured than in L. dentatus, 
sometimes the whole of the last two and the greater part of the rest punctured. All 
lateral ribs of elytra of about equal width. 

Habits. — Mr. Kemp informs me that he got this species from under very thin 
bark, tightly attached to the logs it covered. He only found it on the crest of the 
ridge at Renging. 

L. macassariensis, Schauf., subsp. anibarbis, Kuwert. 
Plate xiii, fig. 54. 
Regd. No. '^ Pcnrissen Sarawak Museum. 

Represented in the Sarawak Museum collection by two specimens from Penrissen, 
one of them from an altitude of 3200 ft. 

Description.— Length 22- s-2i mm. A convex and rather coarse-looking species, 
closely resembling in general appearance the much smaller form L. cyclofaeniiis 
subsp. himalayae, from which it differs in the following respects only. Angle of outer 
side of mandible obsolete. Parietal ridges of head move as in. L. cyclotaenius ; outer 
tubercles rudimentary, situated about twice as far from anterior angles of head as from 
the long inner tubercles. Mesosternum as in L. humerosns, but with the scars and 
dull lateral bands less sharply defined. Central area of metasternum unpunctured. 
Posterior parts of hind coxae polished. 

Leptaulax anipunctus (Zang). 
PL xiii, fig. 55. 

„3, CSansi Gorge, Chinese Frontier, Upper ^^ ^^, ^^^^^ 

^''^'^- ^'^ 1" ? Burma, f,oOO-8ooo ft. S ' ' 

Description.— Length 18 mm. Mandibles as in the following species.^ Head 
coarsely but not very distinctly striate in front, unpunctured except right at 
the back ; frontal area broad , the inner tubercles being about equ idi stant from one 
" I In the specimens I have seen, but not in those described by Kuwert. 

19I4-J F- ^ Gravely : An Account of the Oriental Passalidae. 257 

another and from anterior angles of head ; outer tubercles longer than inner , and 
situated only about \ of the way from them to anterior angles ; parietal ridges short 
as in L. dentatus'. Pronotum rather sparsely punctured in anterior angles and beside 
lateral margin, more closely in scars. Scidelliim and mesothoracic epislerna as in 
L. dentatus and L hicolor \ mesosternum polished, the surface of the scars uneven, 
their inner walls straight, extending the whole length of the plate. Central area of 
metasternum with finely punctured depression in front", and a few large punc ures 
arranged in a pair of lateral groups " rather more than half way back ; lateral areas 
linear, slightly roughened ; intermediate areas smooth except for some punctures near 
inner margin of posterior division Hind coxae and abdominal sterna smooth and 
highly polished. Elytra much as in /.. hicolor, the grooves a little broader, however, 
and the punctures correspondingly coarser, but scarcely transverse. 

Leptaulax bicolor (Fabricius). 

I. L. BICOLOR (Fabricius), s. str. 

PI. xiii, fig. 56. 

i. 4f' Naga Hills ? 

'^ Janakmukh, 600 ft., Abor Country S. W. Kemp. 

'-i^'- Dafla Expedition H. H. Qodwiu-Austen. 

^"'' (Dikrang Valley, Northern frontier 1 .^ tt r^ j • ^ . 

i» ( . , H. H. Godwin- Austen. 

( of Assam ) 

"^' Dunsiri Valley, 500 ft, Assam ? 

Dunsiri Valley, Assam 

9187 9 1K9-9 8 919.)-9 

1 1 ' 1 
9200-;i 9206-10 9212- 

9221- 3 9225-8 9230-1 

1 1 1 
92:U-9 9303-1 1129' 

19" J 


Nilgiri Hills 

Between Sukli and Misty Hollow, 

H. H. Godwin-Austen. 
Stoliczka bequest, etc. 

IStoliczka bequest, 
J. Wood-Mason. 

H. E. Andrewes. 

Dawna Hills, Tenasserim, 2100- , F. H. Gravely 

2500 ft. 

Kuching , Sarawak 

Mt. Kina-Balu, N. Borneo 

Sarawak Museuu 
H. E. Andrewes. 

' Zang placed this species in his genus Leptaulacides , which is characterized by the possession of 
long parietal ridges, and in the type specimens there is less of a break than in ours, between these 
ridges and the supraorbital ridges. In our specimen the break, on one side especially, is of a somewhat 
uncertain character, and I hesitated for some time before definitely describing the parietal ridges as 
short. Subsequent observations of additional specimens have convinced me that the nature of these 
lidges is variable in all species which fall, in the arrangement adopted here, between (but not includ- 
ing) the two dominant species L. dentatus and L. hicolor. 

* These are probably subject to particularly great variation as in other species. 

258 Memoirs of the Indian Museum. {Vox.. Ill, 

This form is represented in the Sarawak Museum collection by specimens from 
Kuching, Matang Rd., 4th mile Rock Rd., Mt. Kinabalu 5000-6000 ft., and Lingga; 
in Mr. Andrevves' collection by specimens from Karkur Ghat (1500 ft., May igii) and 
Nadgam (2500 ft., Oct. 1910) in the Nilgiris, and from Kuching and Mt. Kinabalu in 
Borneo and from Java; and in Mr. Stevens' collection by a specimen from Silonbari 
at the base of the N. L,akhimpur hills in Assam. 

Description.— Length 16-25 mm. Lahrum and mcntum mucli as in the previous 
species of the genus. Mandibles externally angular , upper tooth well developed , though 
less prominent than in L. dentatus ; in front of it a characteristic oblique ridge which 
forms an equally prominent tooth in large specimens but is often imperfect or absent 
in very small ones; anterior lower tooth simple on right side, simple or faintly 
lobed on left. Head more or less clearly punctured in depressions ; frontal area 
usually traversed by a more or less distinct median ridge, more or less longitudi- 
nally rugose, variable in form but normally much broader than in L. dentatus, 
the inner tubercles being at least twice as far from each other as from the smaller 
outer tubercles ; frontal area connected to central tubercle by narrow median ridge ; 
parietal ridges long, extending as far as margin of supra-orbital ridges, low but sharply 
crested along their anterior margin ; canthus less prominent than in L. dentatus. 
Pronotum with strong median groove ; in large specimens strongly and somewhat 
closely punctured in and around scars, very sparsely near angles; in somewhat 
smaller specimens the puncturing of the latter areas is denser, and in smaller ones 
still the pronotum is equally densely punctured along the whole of the sides. 
Prothorax beneath as in L. detitatus. Scutcllum and mesothoracic episterna also as in that 
species. Mesosternum with inner wall of scars normally short and straight (rarely longer 
and more or less curved), the surface of the plate polished, becoming often more or less 
roughened or rugulose (in small specimens especially) and sometimes dull in the 
lateral angles. Metasternum with central area sharply defined, in most large specimens 
highly polished as in L. dentatus , hut in smaller ones usually roughened at least in front 
by fine close puncturing of varying distinctness, a few large scattered punctures some- 
times present ; anterior intermediate area finely punctured ; a broad unpunctured 
band extending from anterior intermediate area between central and lateral areas along 
whole outer border of posterior intermediate area , the inner angle only of which area 
is punctured in large specimens and that sparsely ; lateral areas narrow, linear, rough. 
Hind coxae quite smooth and polished, punctured, or rough. Abdominal sterna with 
lateral punctured areas well developed, otherwise smooth or more or less exten- 
sively and distinctly punctured, the whole of the last segment and the greater part 
of the last but one being densely punctured in some specimens, especially small ones. 
Lateral grooves of elytra not wider than the ridges between them, the strong punc- 
tures with which they are marked not distinctly drawn out to form transverse 

Habits.— Mt. H. L. Andrewes tells me that the much-flattened examples of this 
species which he sent to Mr. H. E. Andrewes from the Nilgiris were obtained in 
Karkur Ghat and adds, " It is gregarious and seems, as its shape would lead me to 

F. H. Gravely : An Account of the Oriental Passulidae. 



suppose, to keep between the bark and the wood of decaying trees instead of workin, 

into the rotten wood as the other Passalidae I know do." 

2. L. BicoLOR var. viciNUS (Perch.). 
PI. xiii, fig. 56f?. 

Regd. No. 


Ganiaduwa, Ceylon 
Dimbula, Ceylon 

I. M. Mackwood. 

Colombo Museum. 

Museum Collector. 

H334-9 9SH-51 0.i5:i-5 

awM 4037:8 40% I Andamans 

1 5 

3721 3724-G 

1(1 111 


8730-i 1 137-s 

Jg SilSl 



•S339-ti; 1H99 
1 1 


> Nicobars 

519 B*2r, 1139 
B 14 19 



Java \ 
Bat] an 1 

iil7 6399 n41 

Sinkep Island 


Philippine Isles 

( Stoliczka bequest, 
( J. Wood-Mason, etc. 

( Stoliczka bequest , 
\j. Wood- Mason, etc 

H. E. Andrewes. 

Moti Ram. 

East India Co.'s Museum. 

Sarawak Museum. 

=^ Kuching, Sarawak 

--f Borneo ) 

This form is represented in the Sarawak Museum collection by specimens from 
Kuching and Matang Road, in the Colombo Museum collection by specimens from 
Ga.maduwa and Dimbulla, in Mr. Green's collection by specimens from Pundaloya, 
and in Mr. Andrewes' collection by specimens from Java and Batjan. 

Description.— Length 13-25 mm. Differs from the typical form in the following 
respects only. Frontal area of head as a rule somewhat more nearly rectangular in 
shape. Pronotum less strongly punctured ; large specimens with a few sparce punc- 
tures in anterior angles, a few set very close together in scars and, as a rule, a few 
more scattered near outer margin ; small specimens more densely punctured, especially 
in neighbourhood of scars.' Mesosternm-n always highly polished, its scars with 
inner wall usually longer and more strongly arched. Puncturing of intermediate areas 
of metastemum sometimes less distinct or more irregular ; central area always highly 
polished. Lateral punctured areas of abdominal sterna more or less obsolete. 

' The puncturing of the pronotum in small specimens is very like that found in larger specimens 
of the typical form. I have not seen any with the uniformly dense marginal puncturing found in most 
small specimens of the typical form. 


Memoirs 0/ the Indian Museum. 

[\OL. Ill, 

Leptaulax roepstorfi, Kuwert. 

PI. xiii, fig. 57. 

This species and the next are the most strongly flattened forms known to me. 

Regd. No. 

Yambung, iioo ft., Abor Country 
Rotung, 1400 ft., Abor Country 
Misty Hollow, W. side of Dawna 

Hills, Tenasserim, c. 2200 ft. 

S. W. Kemp. 

N. Aiiuandale. 

Museum Collector 

UlS 3;:« 37S-,-t 


J. Wood-Mason, etc. 

Description. — Length 15-18 mm. Mandibles sometimes withont ridge between 
upper tooth and first terminal tooth ; otherwise as in L. hicolor. Head proportionally 
broader than in that species, with inner tubercles more widely separated from outer 
tubercles and from anterior angles ; keels of parietal ridges less overhanging. Prono- 
tum very much flattened, with deeply impressed median groove; very sparsely punc- 
tured at sides, punctures usually thickening a little in anterior angles ; prothorax beneath, 
scntellum and mesothoracic epistcrna as in all preceding species of the genus ; mesa- 
stermim as in L. anipunctus, the horizontal central area occasionally marked with 
obscure fine punctures. Metastemum entirely unpunctured, often more or less 
roughened in lateral and anterior intermediate areas, more rarely in posterior 
intermediate areas also; lateral areas linear, not in contact with central area. 
Posterior parts of hind coxae never altogether smooth and polished, mostly rough. 
Abdominal sterna with usual aiiterior marginal band and lateral patches of punctures 
of variable extent ; terminal sternum smooth or more or less covered with more or 
less indistinct punctures. Lateral grooves of elytra not broader than ridges between 
them, their strong punctures not drawn out transversely. 

Habits. — -Mr. Kemp tells me that this species lives, in company with flattened 
Staphylinids and many other kinds of beetles, in deep fissures in very hard jack-fruit 

Leptaulax planus (Illiger). 

Regd No. 

' According to Registei 

PI. xiii, fig. 58. 

Tenasserim ' 

Museum Collector. 


Museum Collector. 

Johore, Malay Peninsula 

J. Wood-Mason, etc. 



Hills between Tenasserim ant 

1 Si; 


about lat. 13 45', long. 98°. 


F. H. Gravely : An Account of the Oriental Passalidac. 261 

Regd. No. -^j^ 

Borneo Sarawak Museum. 

6398 8400 Sinkep Island Museum Collector. 

iJii' Mujang, Sarawak, Borneo C. W. Beebe. 

Specimens are preserved in the Sarawak Museum from Kuching, Matang Road, 
Sadong and Paku. 

Description.— Length 11-14 mm. Differs from L. roepstorfi in the following 
characters only. Mandibles with a minute denticle above at base of upper terminal 
tooth in unworn specimens. Frontal area of head usually longer in the middle line and 
more rounded behind. Pronotum somewhat more thickly and uniformly punctured at 
sides. Mesosternum with inner wall of scars more or less incomplete behind ; central 
area more often, and sometimes also more distinctly, roughened or finely punctured; 
central area of metasterniim normally marked thus ; intermediate areas of metasternum 
strongly punctured throughout. Hind coxae polished. Abdominal sterna uniformly 
marked with large shallow punctures quite unHke those found in any other species of 
the genus that is known to me. 

Trichostigmus thoreyi, Kaup. 

Described from a single specimen from Middle Luzon, in the Hamburg Museum 

Description.— Length 24 ram. Head and its appendages much as in the specimen 
of Leptaulax bicolor shown on pi. xiii, fig. 58 : the middle lower tooth of both man- 
dibles is, however, somewhat more acute ; the median tubercle of the anterior margin 
of the head is obsolete ; the frontal area is wider behind, consequently appearing larger 
and almost rectangular ; and on the mentum the scars are more sharply bent than is 
usual in the genus Leptaulax. Pronotum with strongly developed median and 
marginal grooves, former scarcely complete in front, latter markedly incomplete 
across middle before and behind ; a few punctures devoid of hairs present in anterior 
angles, in anterior extremities of two halves of marginal groove, and above scars ; 
scars and rest of marginal groove thickly set with hair-bearing punctures. Lateral 
plates of prothorax smooth in front and behind, somewhat sparsely punctured and 
hairy between ends of coxal cavities and lateral margins. Scutellum with a few 
distinct punctures near middle. Mesothoracic episterna punctured above and along 
almost whole length of anterior margin. Mesosternum smooth except in scars which 
are rugose. Lateral areas of metasternum narrow, linear, finely roughened, widely 
separated from central area; posterior intermediate areas punctured beside inner 
margin, smooth laterally and behind. Hind coxae and abdominal strrna almost entirely 
smooth. Elytra with lateral grooves more strongly punctured than dorsal, punctures 
of last four grooves somewhat confused with somewhat smaller hair-bearing punctures 
with which the ridges between them are thickly covered. 

262 Memoirs of the Indian Museum. |\'()i. Ill, 


One of the most striking features of the species of Oriental Passalidae, as these 
have been defined above, is that a few of them are remarkably variable in size, 
whereas most are not. The two most variable species are found in the Leptaula- 
cinae. Thej' are Leptanlax bieolor (length 13-25 mm.), and Leptaulax dentatiis (i9'5-37 
mm.). Of both of them the largest specimens I have seen are about twice 
as long as the smallest. Next to these come three species of Aceraiinae, Episphcnus 
indicHS {25-41 mm.), Episphcnus comptoni (27"5-42 mm.) and Aceraius grandis 
(northern race, which I have been better able to study than southern one, 33-49 mm., 
total range of species, including southern forms, 33-54 mm.), of each of which the 
largest specimens are a little more than half as long again as the smallest. Of the 
remaining species of Oriental Passalidae none appear to vary in size in any such 
striking manner; the most variable, as far as my own observations go, are Aceraius 
helferi (30-40 mm.), Leptaulax humerosus {15-20), Leptaulax planus (11-14 mm.), 
Pleurarius brachyphyllus (35'5-45 mm.) and Ophrygonius canton {s. lat., 27 34 mm.), 
of all of which the largest specimens are somewhat more than one and a quarter times 
as long as the smallest; then come Ceracupes austeni (20-25 mm.) and Tiberioides 
kuwerti (36-44 mm.), in both of which the largest specimens are just one and a 
quarter times as long as the smallest ; these are followed by others little less variable 
than themselves; and so on through all the rest. 

Leptaulacides bieolor, Leptaulax dentatus, Episphenus indicus, Episp'ienus comp- 
toni, and Aceraius grandis are all common species, so it is natural to suppose that 
their range of variation may be more completely known than that of other .species, 
most of which seem to be less abundant. But if this was the real and only reason 
for their apparently exceptional variability, we ought to find isolated examples of 
unusual dimensions belonging to at least a few of the many rarer species, and such 
are not found ; and the comparative uniformity in length of all specimens of the long 
series I have seen of Episphenus neelgherricnsis, is sufficient to indicate that even species 
closely allied to those of exceptional variability may be remarkably constant. Tiberius 
andamanensis and sikkimensis, too, are represented in our collection by uniform series 
of sufficient size to render it practically certain that these species do not vary in 
length to any great extent. 

I have examined over a hundred specimens of each of the five variable species, 
except Episphenus comptoni, of which I have seen only thirty-eight'; so it is not 
surprising that the continuity of the variation in length of the specimens before 
me, is much more broken in this species than in any of the others. But for the 
presence among these of three .specimens from Bulutota, of which one is 275, one 
30, and one ^y mm. long, all the evidence would point to the species being divisible 
into two groups distinguished from one another by size alone, in view of which I have 
thought it best provisionally to describe these groups above under separate names, 
since names already in use can be applied to both of them. To my mind, however, 

' I have since seen a number of additional specimens. 

IC)14] F- H : -4;/ Account of the Oriental Passalidac. 263 

the Bulutota specimens indicate with sufficient clearness that a larger series would 
prove the range of variation to be perfectly continuous, as it is in the other four 
variable forms ; for which reason I have described them above without the introduction 
of any new name. This view, as will be seen later, is further supported by the fact 
that Episphenus comptoni is a gregarious species, is the only asymmetrical member 
of the Aceraiinae found in Ceylon, and is also the commonest member of that sub- 
family living in the island . 

The specimens from which my descriptions of Episphenus indiciis and Acerains 
grandis sub-sp. hirsiitus were drawn up show, as a whole, no breaks in the continuity 
of the range of variation in length. Moreover they seem to indicate that different 
colonies differ from one another in average size, and that the lengths of different 
members of any one colony do not usually differ very greatly one from another. 
The evidence is most definite in the case of Aceraius graniis. Out of a series of 
38 specimens of this species known to have been taken from a single colony at 
Janakmukh, 37 show a continuous range of length from 36 to 39 mm., the remain- 
ing specimen being 33 mm. long ; and again, the specimens from Kalek form a single 
short series of individuals all between 38 and 40 mm. in length. On the other hand, 
two specimens, presumably of opposite sex, taken from a single burrow at Rotung 
were 34 and 41 mm. long respectively. Such facts suggest that although large and 
small specimens may be quite well able to unite to found a new colony, their off- 
spring tend to be to of a moderately uniform size ; and that the majority of speci- 
mens in each colony are the descendants of the original founders of that colony, 
which do not leave their home until they are compelled to do so, when they may 
either found new colonies or enter the ranks of an old one from whose average size 
they may differ very considerably. 

Such evidence as is available regarding the variable species of lyCptaulacinae all 
points in the same direction. Leptaulax dentatus, s. str., is represented in our collection 
by over three hundred specimens, whose range of variation in length is unbroken; 
but when these are arranged according the localities from which they come, it is found 
that those from each well-represented locality fall into several separate series, each of 
which probably includes only specimens from colonies that are not represented in 
any of the other groups, unless it be by one or two specimens that have come into the 
colony from outside. I have not attempted to make any elaborate measurements 
with which to support these facts ; for length measurements of dried Passalids are 
only approximately comparable one with another, on account of the variable extent 
to which the back of the head and front of the metathorax are telescoped into the 
prothorax ; and breadth measurements are so variable in proportion to length that 
it would be still more misleading if I were to use them as an index of length. The 
following measurements of the extreme members of each of the several unbroken 
series, into which the specimens from each of our three best represented localities 
appear to the unaided eye to fall, will I think give a better idea than anything else 

I Since the above was written I have obtained direct confirmation of the continuity of the variation 
found in this species. See below, p. 282, footnote. 

264 Memoirs of the Indian Museum. [Vol. III. 

of what has ])een found. Out of 46 specimens labelled " Upper Tenasserini " 17 are 
2I-5-24-5 mm. long ; 2 are 26-5 and 28 mm. long respectively ; and 27 are 30-37 mm. 
long. Out of 45 specimens labelled " Tenasserim " 21 are 21-24-5 mm. long, i is 
275 mm. long; and 23 are 30-35 mm. long. And out of 43 specimens labelled 
" Sikkim " 2 are 195 and 20 mm. long respectively, 8 are 21-23 mm. long, and 33 are 
24-5-31 mm. long. On the other hand, 47 specimens known to have been collected 
from a single colony at Kawkareik are all 29-335 mm. long, and form a single 
unbroken series. 

Leptaulucides bicolor is unusually variable in the extent to which the body is 
flattened, as well as in total length, and in the extent to which certain plates are 
punctured. For this reason, and partly, perhaps, because the specimens I have seen 
belong in almost equal numbers to two dififerent forms of the species, which must, at 
least provisionally, be treated separately, these specimens have proved not to be nearly 
enough to form a satisfactory basis for any full account of the manner in which different 
colonies vary one from another. It appears, however, that size is not a character 
that is at all likely to prove of any great importance for the sub-division of the 
species into sub-species or varieties. 

Members of a single colony in this, as in other variable species, probably differ 
comparatively little one from another in .size, which suggests that they may be com- 
paratively uniform in structure also. Any investigations on a collection in which 
different individuals, instead of different colonies, have to be taken as imits, must be 
conducted with great caution ; for the presence in collections of a number of short 
series from different colonies, is likely give a false appearance of the constancy of a 
number of more or less distinct types ; and I am confident that this fact more than 
any other has led, on account of the remarkable variability of the common species 
L. bicolor and L. dentatus, to the appalling multiplication of so-called species of the 
genus Leptaulax. The colony ought, in my opinion, to replace the individual as the 
unit for taxouomic work in all these variable species, and it is partly in the hope that 
collectors will turn their attention to collecting separate colonies, and partly to avoid 
adding to what I am convinced is largely a meaningless multiplication of names, that 
I put forward here, on the somewhat inconclusive evidence at present before me, the 
hypothesis that members of any one Passalid colony are much more uniform than 
the species as a whole when this is markedly variable. 

In connection with this hypothesis it would be interesting to know how long logs 
of different kinds can remain suitable for the habitation of a single species of Passalid 
under the normal conditions of different jungles, and the length of time occupied by 
the life cycle of different species of Passahdae in various localities, together with any 
facts relating to the period or periods of reproduction of individual pairs. For if logs 
rot more rapidly than colonies can be formed, it would follow that colonies are 
produced by the attraction of suitable food ; in which case the members of each would 
be unlikely to have a common descent, and it would no longer seem so probable that 
members of a single colony should be more uniform than the species to which it 

I9I4-1 F- H. Gravely ; An Account of the Oriental PassaUdae. 265 

In the Passalidae, as in the L,ucanidae, variation in certain structural characters 
is found to be more or less closely correlated with variation in size. The mandibles 
of a Passalid never attain anything approaching the remarkable development of 
those of a male Lucanid except in the genus Ceracupes, the species of which are not 
known to vary greatly in size ; yet in two of the most variable Passalids — A ceraius 
^raridis sub-sp. hirsutus and Leptaulax dentatus (? s. str. only) — their dentition is less 
complex in small than in large specimens. In the former the upper tooth of the left 
mandible is often double in very large specimens (fig. 28a), but not in medium sized 
(fig. 28), or small ones ; and in the latter, the anterior lower tooth of the left mandible 
is tridentate in large specimens (fig. 52) and bidentate (sometimes scarcely more than 
unidentate) in small ones (fig. 521^). 

The processes of the anterior margin of the head on the other hand tend to be 
more acute, and often proportionally longer, in small specimens than in large ones, 
when any such differences are found. This is best seen in the processes developed 
from the anterior angles of the head in Episphenus indicus and Aceraius grandis 
(? sub-sp. hirsutus only) as will be seen on comparing fig. 20 with fig. 20b and 
fig. 28 with fig. 28a. A somewhat similar phenomenon is found among Lucanidae in 
the genus Aegus. In large specimens of certain species of this genus (e.g. A . kandyensis 
and A. roepstoyffi) the front of the head bears two median protuberances, one above 
the other; while in smaller specimens of the same species, although the uppermost of 
these disappears, the lower one is represented by a pair or minute tubercles, situated 
side by side and more distinctly separated one from another the smaller the specimen is. 

The relative extent of the punctured areas of large and small specimens presents, 
however, a much better instance of the similarity between the variations in structure 
associated with variations in size, found in the two families. In both families it 
seems to be always the small specimens that are most extensively punctured, in 
species in which any such difference occurs. In the lyucanidae (e.g. certain 
Himalayan species of Dorcinae) the difference is most marked in the elytra, and after 
that in the pronotum and sometimes the abdominal sterna. In the Passalidae the 
elytra are always furrowed, and such slight variations as occur in the extent to which 
the furrows are punctured, do not appear to be in any way correlated with variations 
in the size of the insect, nor does the puncturing of the lateral ribs of the elytra of 
Aceraius grandis appear to be so correlated ; but in Episphenus indicus Sind Leptaulax 
bicolor (s. str. and to a less extent var. vicinus as well) , the pronotum is more extensively 
punctured in small than in large specimens ; and in Leptaulax bicolor the punctured 
areas of the abdominal sterna are commonly more extensive in small specimens than 
in large ones. 


The principal difficulty in drawing up this list has been the working out of the 
synonymy which it has involved. An attempt to discuss the distribution of Oriental 

266 Memoirs of the Indian Museum. [Vol. Ill, 

Passalidae, which does not include an attempt to deal with the synonymy of Kuwert's 
multiplicity of names, would be of little value, since for most species he gives only a 
single record ; but his descriptions are so inadequate that no synonymy drawn up — 
as this is— without reference to a complete set of his types is likely to be wholly free 
from error. As, however, the large series of specimens that I have been able to 
examine, especially of the genera Episphenus , Aceraius smd Leptaulax^ besides showing 
how unexpectedly variable in size, and in some respects also in structure, certain 
species of these genera may be, have given me an exceptionally good opportunity of 
acquainting myself with the range of variation which individual species normally 
exhibit, I have ventured to apply the knowledge thus acquired to the synonymy of 
species which I have not seen. It should be noted that the collection in the Konigliches 
Zoologisches Museum in Berlin shows that Kuwert was often himself unable to recog- 
nize his species a second time, and has sometimes applied the same name to quite 
distinct species, as well as describing specimens of a single species under a number 
of different names. An examination of Kuwert's cotypes, such as I have made in a 
number of instances, is not , therefore, so safe a guide in dealing with his synonymy 
as it ought to be. 

I have tried to include in this part of the paper every published reference to 
every described species of every genus at present knowai from the Oriental Region 
together Avith China and Japan. This has involved the inclusion of many species with 
which I am not directly acquainted. Consequently the completeness of the synonymy 
indicated probably varies considerably in different genera. The names of genera and 
species which I have not myself examined are marked with an asterisk, as also are 
references to works w'hich I have not been able to consult. References to works which 
do not contain descriptions or original notes on the particular species mentioned are 
enclosed in brackets — references to many of the new species in Kuwert's catalogue of 
1891 are included among these as many of the new names there published were nomina 
nuda, and remained such till the publication of his dichotomous tables in 1896-1898. 
The word ' ' part ' ' in square brackets in such cases indicates that two or more distinct 
species have been included in the author's synonymy; but when used in connection 
with a published description it implies that two or more species have been confused 
by the author himself when drawing up his description, no notice being taken of his 
synonymy. Apart from this, square brackets always indicate that a species is not 
known from the Oriental Region, China or Japan. A cross (f) in front of a specific 
name, implies that I have seen specimens to which this name has been applied by the 
author mentioned in the reference accompanying the name so marked. 

The names of forms which I regard as distinct varieties or sub-species are 
included in the synonymy of the species to which I consider them to belong, as at 
present any attempt to recognize them separately could only lead to confusion. 
All PassaHds recorded from " Sikkim " belong to species found in " Sikkim ' ' , 
which is now better known as the " Darjeeling District" ; that term is employed 
throughout this list in place of the term "Sikkim." And it has been found con- 
venient to group together all localities in the Eastern Himalayas and their foothills, 

1914.1 F H. Gravely : An Accottnt of the Oriental Passalidae. 267 

restricting the terms Bengal and Assam to the parts situated south of these 
slopes. "Malacca" has so often been used to signify the whole of the Malay 
Peninsula that it is unsafe to attach any more precise significance to it when dealing 
with old records; "Malacca" has therefore been replaced by the term "Malay 
Peninsula " here, a term used to include islands situated close to the mainland, such 
as Singapore. 


Members of this subfamily occur in China and Japan and throughout almost the 
the whole of the Indo- Australian region ; but only one specimen has yet been recorded 
from the Indian Peninsula, and none from Ceylon. 

Genus COMACUPES, Kaup, 1871. 
= Aulacocyclus [part J, Kaup, 1868. 
This genus occurs in the Malay Peninsula, Sunda Islands, Philippines and 
Celebes. " Comacupes" felderi, Stoliczka (1873, p. 152) probably belongs to the 
genus Tristorthus, so is omitted here. 

Comacupes foveicollis, Kuwert. 

Comacupes foveicollis, Kuwert, 1891, p. 171. 
Comacupes foveicollis, KvLweit,i8g7, -p. 284. 
Comacupes t minor. Heller, 1896, pp. 3-4, pl- i- figs- ii-ii«- 
The hairless abdomen and practically unpunctured mesosternum suggest that 
this species may be more closely alUed to the Australian genera Tristorthus and Aula- 
cocyclus than to any member of the otherwise purely Oriental genus Comacupes. The 
mentum is, however, more or less keeled in front, so I retain it in the last-named genus. 

In the ten specimens from Sarawak in the British Museum the central tubercle 
is even less distinctly bifid than in the type of C. minor, Heller, from Celebes; so the 
latter probably represents a distinct local race. 

Localities:— ^ q FOVEICOLLIS , s. str, 

Borneo : Sarawak — Baram River. 

2. Sub-sp. MINOR, Heller. 
Celebes: N. coast — Buol. 

Comacupes basalis (Smith). 

Passalus -fbasalis, Smith, 1832, p. 18, pl. i, tig. 5. 

Aulacocyclus comatus, Kaup, 1868a, p. 9. 
[Aulacocyclus comatus + Leptaulax basalis, Gemminger and Harold, 1868, pp. 968-9). 

Comacupes basalis, Kaup, 1871, p. 19, pl. iii, figs. 3(?-3^- 
(Comacupes basalis, Wytsmann, 1884, p. 329). 
(Comacupes Basalis, Kuwert, 1891, p. 171). 

Comacupes basalis, Kuwert, 1896, pp. 284-5. 
(Comacupes basalis, Zang, 19036, pp. 418-9). 

Locality: — 

Philippine Islands. 

268 Memoirs of the Indian Museum. (Vol. Ill, 

Comacupes cavicornis, Kaup. 

Aiiliicocycliis cavicornii, Burmeister iii Kaup', + lucvkuinis, Kaup, i868rt, p. 6. 
{Aiilacocyclns cavicornis + laevicoinis , Gemminger aiul Harold, 1868, pp. 698-9). 

Comacupes cavicornis, Kaup. 1871, p. 19. 
{Aulacocyclus ^ parryi, Stoliczka, 1873, p. 150). 
(Comacupes cavicornis, VVytsman, 1884, p. 329) 
(Comacufes Cavicornis [part], Kuwert, 1891, p. 171). 

Comacupes ^westermanni, Kuwert, 1897, p. 284. 

Comacupes cavicornis, Zang, 3905c, p. 226. 

Comacupes \ cavicornis + vat. -fborneensis + var. j; lacvicornis., (iravely, above, pp 204-205, pi xi. 
figs. 1-26. 

All the specimens identified by Stoliczka as A idacocycins parryi have proved to 
be in reality Comacupes cavicornis var. laevicornis. Zang has pointed out that Kuwert's 
Comacupes wester manni, not his Tristorthus cavicornis, is identical with Kaup's Coma- 
cupes cavicornis. 
Localities : — 

Malay Peninsula: Penang; Johore (? all var. laevicornis). 
Sumatran Islands; Sumatra ; Siukep Island (? all typical form). 
Java (? another form).' 
Borneo Sarawak: — Kuching (? all var. borneensis). 

Comacupes stoliczkae, Gravely. 

Comacupes cylindraccus (part), Kaup, 1871, p. 18, pi. iii, upper unit of fig 4. 

Comacupes ^cylindraceus, Stoliczka, 1873, p. 151. 
?{Comacitpes cylindracetts , Kuwert, 1891, p. 171). 
? Comacupes cylindraceus [part] Kuwert, 1897, P- 285. 

Comacupes ^stoliczkae. Gravely, above, p. 206, pi. xi, figs. 3 jk. 

This species was described by Kaup as a form of C. cylindraceus. As pointed out 
above (pp. 205-6) Kaup's two forms of C. cylindraceus are quite distinct; and it 
is probably to the other that the name was first applied. 
Localities : — 

Malay Peninsula : Johore ; ? Singapore. 
Sumatran Islands: Sinkep Island. 
Kuwert definitely recorded this species from Singapore in 189 1 ; but later he 
appears to have doubted the correctness, either of his identification of the insect, or 
of the label indicating the locality from which it came. 

Comacupes masoni, Stoliczka. 

Comacupes ■fmasoni, vStoliczka, 1873, pp. 151- 2. 
Comacupes Masoni, Kuwert, 1891, p. 171. 

' Kaup appears to have described this species under an unpublished name attached by Burmeister 
to the type specimen in Germar's collection. 

* This record is based on a specimen in the British Museum. 

igi4.] F. H. GKAVEI.V : Aii Account of the Oriental Passalidac. 269 

Comacupes fcylindraceus, [part], Kuwert, 1S97, p. 285. 
Comacupes jniasoni, Gravely, above, p. 207, pi. xi, figs. 4-4ii. 

Zang has suggested that Kuwert's C. masoni of 1897 is not StoHczka's, and that 
the latter is identical with Kuwert's C. angusticornis ; an examination of the type 
of C. masoni, and of specimens determined by Kuwert, has confirmed the former 
suggestion but not the latter. 
Locality : — 

Malay Peninsula : J oho re ; Perak ' ; Tengah Mountain. ' 
N. Sumatra : Silulangit.^ 

Kuwert's 1891 record "Lahore " is doubtless a mistake, as he appears to have 
been unacquainted with the species at that time except through StoHczka's account 
of it. 

Comacupes cylindraceus (Perty). 

Passcdus cylindraceus, Perty, 1831, p. 36, fig. 3. 

Passalus cylindraceus, Percheron, 1841, pp. 40-1, pi. Ixxix, fig. 4. 

Passalus punciifrons , Hope, 1845, p. 29. 

Passalus cylindraceus, Burmei',,ter, 1847, pp. 516-7. 
(Aulacocycltts cylindraceus [part], Kaup, i86S«, p. 4). 
(Aulacocyclus cylindraceus [part], Gemminger and Harold, 1868, p. 968). 

Comacupes cylindraceus [part], Kaup, 1871, p. 18, pl. iii, lower unit of fig 4. 
{Comacupes cylindraceus [part], Wytsmann, 1884, p. 329). 

Comacupes ^ angusticornis + ^ cylindraceus [part] + masoni, Kuwert, 1897, p. 285. 
(Comacupes cylindraceus, Zang, 190^^, p. 419). 
(Comacupes f Masoni, Zang, 1905c, p. 231). 

Comacupes f cormocerus, Zang ; 1905^, pp. 233-5. 

Comacupes f punciifrons, Arrow, 1907, p. 444. 

Comacupes ^ cylindruceus. Gravely, above, p. 207, pl xi, figs. 5 6(i, text-fig 2. 

Other references to the name cylindraceus will be found under C. stoliczkae 
(previous page). 
Localities : — 

Malay Peninsula: S. Perak (Telom, 4000 ft.), Larut Hihs, 3300-4300 ft., 

Sumatran Islands : Sinkep Island. 
Sumatra — Padang ; St. Rambe. 

Borneo: Sarawak — Kina-Balu, ca. 5000 ft.; Kuching; Trusan ; Penrissen 4200- 
4500 ft. ; Klingkang. 

1 These records are based on specimens in Berlin which I did not see in time to include any account 
of them in part 4 of this paper. 

^ There is a single specimen from this locality in the British Museum ; it is less distinct from 
C. cylindraceus than the specimens I have seen ixom. the Malay Peninsula, the anterior part of the 
central tubercle, though much elevated, being angular beneath; it mav perhaps belong to a distinct 
local race. 

270 Mcniuirs 0/ the Indian Museum. [Vol,. Ill, 

Genus TAENIOCERUS, Kaup, 1871. 
= Aiil(icocyclns [part], Kaup, 1868; Tacniocerus [partj, Kaup, 1871. 
This genus includes species found in all parts of the Oriental Region except the 
Indian Peninsula and Ceylon. 

T. platypus, Kaup. 
Aulacocychis platypus, Kaup, 18680, p. 5. 
(Aulacocyclns platypus, Gemmiuger and Harold, i8ij8, p. Qbg.) 
Taeniocerus phdypus, Kaup, 1871, p. 21. 
Aulacocychis platypus, Lewis, 1883, p. 341. 
{Taeniocerus platypus, Wytsmau, 1884, p. 329.) 
{Taeniocerus Phitypus, Kuwert, 1891, p. 170.) 
Taeniocerus platypus, Kuwert, 1897, p. 275. 

Locahties ' : — 

Malay Peninsula : Perak ; Naungcliik (Bukit Besar, 2500 ft.).^ 
Sumatra: Deli. 
Borneo: Sarawak. 

Taeniocerus bicanthatus (Percheron). 

Passalus bicanthatus, Guerin-Meneville' in Percheron, 1841, pp. 41-2, pi. Ixxix, tig. 5. 
?Passalus bicanthatus, Burmeister, 1847, p. 516. 
{Passalus bicanthatus. Smith, 1852, p. 17.) 

Aulacocyclns bicanthatus, Kaup, 1868a, p. 6. 
(Aulacocychis bicanthatus, Gemininger and Harold, i8b8, p. 968). 

Taeniocerus bicanthatus, Kaup, 1871.. p. 21, pi. iii, fig. 6. 

Taeniocerus ^bicanthatus, Stoliczka, 1873, p. 152. 
(Taeniocerus bicanthatus, Wytsmaa, 1884, p. 329.) 
(Taeniocerus Bicanthatus, Kuwert, 1891, p. 170.) 

Taeniocerus ^bicanthatus i-a.b. perturbans + imitator , Kuwert. 1897, p. 276. 

Taeniocerus bicanthatus, Zang, 1905^, p. 105. 

Taeniocerus ^bicanthatus, Gravely, above, pp. 208-209, pi. xi, figs. j-jb. 

Burmeister 's description of this species is not altogether in agreement with my 
observations. Neither in this, nor in either of the other known species of Taeniocerus, 

' Borneo is the locality given by Kaup, who mentions 19 specimens in the British Museum. Lewis, 
however, speaks of "A. platypus, Kaup, from Tibet, of which there is a series in the National 
Collection" of Britain. The other localities given here are those of specimens in Berlin, which I did not 
see in time to include any account of them in part 4 of this paper. 

' This record is based on a pair of specimens in the Oxford Museum. 

•' Percheron gives Guerin-Meneville as the author of this and other names, without giving a reference. 
In the case of Passalus (Episphenus) neelgherriensis, where he gives as leference " Rev. Zool.", a search 
through all possible volumes of that publication has failed to reveal any mention of the species ; and as 
Guerin-Meueville himself wrote (1843, p. 50) two years after the publication of Percheron's paper 
"Cette espC-ce a ete dccouverte par M. Ad. Delessert sur le plateau des Neelgheries. Nous I'avons 
communique a M. Percheron, qui I'a fait eutrer en notre nom, dans sa Moiiographie . ..." it is 
evident that Percheron was the first to publish both the name and the description. Consequently in all 
such cases I have attributed the species to him, giving in the synonymy, however, also the name of 
the author to whom he attributed it. 

I9I4-] F- H. Gravely : An Account of the Oriental Passa/idae. 271 

is the raesosternum hairy ; whereas this is the case in every species of Comacupcs 
known to me. Consequently, I should have had no hesitation in regarding the speci- 
men described by Burmeister from Germar's collection, as a member of the latter 
genus, had not his description of the head appeared to me to be rather that of the 
head of the former genus. Until the specimen is re-examined its identity must remain 
uncertain. Passalids often break in two between the pro- and mesothorax, and it 
is quite possible that the specimen is really a composite one, the anterior part of a 
Tacniocerus having been gummed on to the posterior part of a Comacupes. Concern- 
ing Zang's suggestion that variations in the form of the central tubercle indicate a 
splitting into local races, see above, p. 208 footnote. 

I_,ocalities : — 

Malay Peninsula : Johore. 
Sumatra: N. E. Sumatra. 

E. Sumatra — I^angkat. 
Bornean Islands : Borneo. 

Smith's record of this species from India is too vague to be of any value, since 
the term may easily have been used to include the whole of " Further India " as well 
as the Indian Peninsula. 

Taeniocerus pygmaeus, Kaup. 

Aulacocyclus pygmaeus, Kaup, iS68a' p. 5. 
(Aulacocydus pygmaeus, Gemminger and Harold, 1868, p. 969.) 

Taeniocerus pygmaeus, Kaup, 1871, pp. 20-21. 
{Taeniocerus ^pygmaeus, Stoliczka, 1873, p. 152.) 
(Taeniocerus pygmaeus, Wytsmau, 1884, p. 329.) 
(Taeniocerus Pygmaeus, Kuwert, 1891, p. 170.) 

Taeniocerus "f parvus -i- pygmaeus, Kuwert, 1897, p 27b. 
{Taeniocerus pygmaeus, Zang, 19036, p. 419.) 

Taeniocerus ^pygmaeus, Gravely, above, pp. 209-210, pi. .xi, figs. 8-Sa. 

The specimens of this species that I have seen show quite clearly that the three 
characters by which Kuwert separates his T. parvus from Kaup's T. pygmaeus are 
all of them variable. Indeed, the hook-shaped and antler-shaped pronotal scars, by 
which they are primarily distinguished in his table, may occur on opposite sides of 
the same individual. 

Localities : — 
Malay Peninsula. 
Borneo : Sarawak — Kuching. 

' This record is based on speciineas in Berlin, not seen in time for description in part 4 of this 

272 Memoirs ni the Indian Museum. [Vol. Ill, 

Taeniocerus bicuspis, Kaup. 

? Passaliis biliastatus 9 , Percherou, 1844, p. 13. 

Aulacocyclus bici(sf>is, Kaup, iS68rt, p. 5. 
(Aiilacocycliis bicitspis, Gemminger and Harold, 1868, p. 968.) 

Taeniocerus bicuspis, Kaup, 1871, pp. 2t-2. 

Taeniocerus ^bicuspis, Stoliczka, 1873, p. 152. 
(Taeniocerus bicuspis, Wytsmati, 1884, p. 329.) 
(Taeniocerus Bicuspis, Kuwert, 1891, p. 170 ) 

Taeniocerus bicuspis, Kuwert, 1897, pp. 276-7. 

Taeniocerus "fbicuspis. Gravely, above, pp. 210-211, pi. xi, figs. 9-9rt. 

Localities : — 

E. Himalayas: Darjeeling Dist. — Sureil; Choongtang; Upper Tista, 4000 ft.; 

Gopaldhara, Rungbong Valle}-. 
Dafla Hills— Harmutti, base of Dafla Hills; Dikraiig Valley. 
Abor Country — Kobo, 400 ft. ; Rotung, i.:|oo ft. ; Upper 
Rotung, 2000 ft. ; Upper Renging. 
Assam : Cachar. 

X. Lakhimpur — vSilonbiri (base of hills). 

Malay Peninsula. 

Genus AULACOCYCLUS, Kaup (1868) 1871. 
= Aulacocyclus [part], Kaup, 1868; incl. Taeniocerus [part] Kaup, 1871. 

I have not attempted to do more than accept the opinions of previous authors 
in the following list of the species of this genus. Blackburn (i8g6, p. 234) has 
already pointed out the difficulty of dealing with it. He suggests that several of the 
names accepted below are really synonymous. I am inclined to agree with him; but 
I cannot help thinking, in addition, that the characters to which he has been driven 
to attach the greatest taxonomic importance, are likely to prove to have been 
misleading, when a larger number of specimens are compared together. 

The genus is centred in the parts of the Indo- Australian Region east of Wallace's 
Line, but occurs in the Philippines and Sunda Islands also. Only one specimen has 
yet been found on the mainland of Asia ; it is from the Indian Peninsula. 

I Aulacocyclus deyrollei, Kaup. | 

Aulacocyclus Deyrollei, Kaup, 1868a, p. 7. 
(Aulacocyclus Deyrollei, Gemminger and Harold, 1868, p. 968.) 

Taeniocerus Deyrollei, Kaup, 1871, j). 20. 
(Taeniocerus Deyrollei, Wytsman, 1884, p. 329.) 

' These records are based 011 specimens in Berlin, not seen in ti.ue for description in part 4 of 
this paper. 

igi4.] F. H. (iRAVELY : An Accouni of the Oricutal Passalidae. 273 

(Tueitiocerus Deyrollci. Kuwert, 1891, p. 170.) 
Taeniocenis deyroUei, Kuwert, 1897, p. 275. 
Taeniocents deyroUei, Arrow, 1907, p. 447. 

Locality : — 

Australia: N. yueensland — Port Denison^ 

[Aulacocyclus masters!, MacIyeay.J 

* Taeniocerus mastersi, MacLeay, 1871, p. 174. 
(Taeniocents Mastersii, Wytsraan, 1884, p. 329.) 
{Taeniocerus Mastersii, Kuwert, 189:, p. 170.) 
Taeniocerus mastersi, Kuwert, 1897, p. 275. 

It is possible, I think, that this species and the last may prove to be identical. 

Locality : — 

AustraUa : Queensland — Oayndah '. 

[* Aulacocyclus teres (Percheron).] 

I'assuliis teres, Percheron, 1841, pp. 39-40. 

Passalus teres, Burmeister, 1847, p. 515. 

Passalus teres, vSmith, 1852, p. 19. 

Passalus teres, Redtenbacher, 1867, p. 50. 

Aulacocyclus teres, Kaup, 1868a, p. 9. 
(Aulacocychis teres, Gemminger and Harold, 1868, p. 969.) 

Aulacocyclus teres, Kaup, 1871, pp. 15-16, pi. iii, figs. i-i«. 
[Aulacocyclus teres, Wytsman, 1884, p. 328.) 
(Aulacocyclus Teres, Kuwert, 1891, p. 170.) 

Aulacocyclus teres, Kuwert, 1897, p. 277. 

Locality : — 

Australia: ? Sidney. 

Redtenbacher' s Sidney specimen very possibly has the same sort of history as 
his Mastochilus poliins, concerning which see Stoliczka, 1873, p. 156. 

[* Aulacocyclus tereoides, Kuwert]. 

Aulacocyclus Tereoidts, Kuwert, 1891, p. 170. 
Aulacocyclus tereoides, Kuwert, 1897, p. 277. 

Locality: — 

[* Aulacocyclus aliicornis, Kuwert.] 

Atdacocyclus alicornis + ah. sulcatipons, Kuwert, 1897, pp. 277-9. 
Aulacocyclus alicornis ab. sulcatipons, Zang, 1905c, p. 230. 

' Not St. Denis, Reunion Islands (see Arrow loc. cit.). 

* Kuwert's 1891 catalogue gives not Queensland but N.S. Wales as the locality of this species, 
perhaps because it was originally described in the " Transactions of the Entomological Society " of that 
country. This is not repeated in his later work. 

274 Memoirs of the Indian .^lusenni. [VOL. Ill, 

Locality : — 

I * Aulacocyclus foveipunctatus, Kuwert.] 

Aitlucocycliis fovcipiiiniiilns, Kuwert, 1S97, p. zyS. 

Locality : — 

[* Aulacocyclus, rotundatoclypeatus, Kuwert. 

Aulacocyclus Rotiindatoclyfyeatiis. Kuwert, 1891, p. 170. 
Anlacocycltis rotundcilodypealiis , Kuwert, 1897, pp. 278 and 2i>z. 

Locality : — 

Australia: Cape York. 

[* Aulacocyclus arcuatoclypeatus, Kuwert. J 

Aulacocyclus Arcualoclypealus , Kuwert, 1891, p. 170. 
Aulacocyclus arcualoclypeatus, Kuwert, 1897, pp. 278 and 282. 
Locality : — 
New Guinea : Ply River. 

* Aulacocyclus fratricornis, Kuwert.] 

Aulacocyclus Fratricornis, Kuwert, 1891, p. 170. 
Aulacocyclus fratricornis, Kuwert, 1897, pp. 279 and 282. 

Locality : — 
? South Sea Islands or Australia. 

Aulacocyclus rosenbergii, Kaup. 

Aulacocyclus Rosmbirgii ■(- Perchsroni, Kiup, i868(i, pp. 7 and 8. 
(AtUacocyclus Rosenbergi + Percheroni, Geminiuger and Harold, 1868, p 969.) 

Aulacocyclus Rosenbergii, Kaup, 1871 p. iG. 
{Aulacocyclus Rosenbergii, Wytsman, 1884, p. 328.) 
{Aulacocyclus Rosenbergii, Kuwert, 1891, p. 170.) 

Aulacocyclus Rosenbergii -^^ Percheroni, Blackburn, 1896, p. 234. 

Aulacocyclus rosenbergi Kuwert, 1897, pp. 279 and 282. 
{Aulacocyclus Rosenbergii, Zang. 19036, p. 419.) 

Localities : — 

Sumatran Islands : Nias. 
Australia : Brisbane '. 

Zoogeographical considerations suggest that Kaup, in his effort to reduce the 
number of species of Aulacocyclus to five, regarded two distinct forms as one when he 
merged A. percheroni in A. rosenbergii. This course has, however, been followed by 
all subsequent authors, except perhaps Blackburn, most of them omitting any refer- 
ence to the locality of the type specimens of A . percheroni. 

' Kaup gives "Trisbane," Ijut Geniniinsier and Harold are presumably right in regarding thi-^ 
as a misprint for Brisbane. 

igi4.] F. H. Gravely : An Account of the Oriental Passalidae. 275 

[* Aulacocyclus kaupi, MacLeay.] 

*Aulacocydus kaupi, MacL,eay, 1871, p. 173. 
{Aulacocyclus Kaupi, Wytsman, 18S4, p. 328.) 
{Aulacocyclus Kaupi, Kuwert, 1891, p 170.) 

Aulacocyclus Kaupi, Blackburn, 1896, p. 234. 

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

lyocality : — 

Australia: Queensland — Gayndah. 

[* Aulacocyclus edentulus (MacLeay).] 

Passaius edentulus, MacLeay. 1826, p. 439. 
*Passalus furcicornis, Boisduval, 1835, p. 242. 

Passaius cylindraceus, Percheroii, 1835, pp. 103-4, P'- vii, fig- 8. 
* Passaius furcicornis, Dejean,' 1837, p. 195. 

Passaius edentulus, Percheron, 1841 , p. 39. 

Passaius edentulus. Burmeister, 1847, pp. 515-6. 
{Passaius edentulus. Smith, 1852, p. 20.) 

Passaius furcicornis, Montrouzier, 1855, p. 29. 

Passaius turcicornis, Montrouzier, i860, p. 288. 

Passaius furcicornis, Fauvel, 1862, p. 135. 
{Aulacocyclus edentulus, Kaup, 1868a, p. 8.) 
{Aulacocyclus edentulus, Gemminger and Harold, 1868, p. 969.) 

Aulacocyclus eientul us, Kaup, 1871, pp. 14-15. 
{Aulacocyclus edentulus, Wytsman, 1884, p. 328.) 
{Aulacocyclus Edentulus, Kuwert, 1891, p. 170.) 

Aulacocyclus edentulus , Blackburn, i8g6, pp. 233-4. 

Aulacocyclus edentulus, Kuwert, 1897, pp 280 and 282. 
Localities : — 

Australia: Port Jackson. 
New Caledonia : Balade. 

[* Aulacocyclus errans, Blackburn.] 

Aulacocyclus errans, Blackburn, 1896, pi. 233. 
Locality : — 

[* Aulacocyclus collaris, Blackburn.] 

Aulacocyclus collaris, Blackburn, 1896, pp. 233-4. 

Locality : — 

Aulacocyclus andrewesi, Gravely. 

Aulacocyclus ■^ andrewesi. Gravely, above, pp. 2TI-212, pi. xi, figs. lo-ioa. 
Locality : — 

India: Anamalai Hills, ca. 3500-4000 ft. 

Gemminger and Harold give d'Urville as the author of this name in Dejean 's catalogue. 

276 Memoirs of flic Indian Museum. [Vol. Ill, 

I* Aulacocyclus glabriusculus, Kuwert.) 

Aulacocyclus Glabriusculus, Kuwert, 1891, p. 170. 
Aulacocyclus glahrinsctdus, Kuwert, 1897, pp. 280 and 282. 
Locality: — 
Aru Islands. 

♦Aulacocyclus diUtus, Kuwert. 
Aulacocyclus DiLitus, Kuwert. 1891, ji. 170. 
Aulacocyclm (Hiatus, Kuwert. 1897, pp. 281 and 282. 
Locality: — 

Philippine Islands. 

Aulacocyclus parryi, Kaup. 

Aulacocyclus Parryi, Kaup, i868((, p. 8. 
(Aulacocyclus Parryi, Gemminger and Harold. 1808, p. gbq.) 

Aulacocyclus Parryi, Kaup, 1871, p. r6. 

Aulacocyclus parryi, Kirsch, iSjja, p. 139. 
(Aulacocyclus Parryi, Wytsman, 1884, p. 328.) 

Aulacocyclus Parreyi, Kuwert, 1891, p. 170. 

Aulacocyclus parryi, Kuwert, 1897, p. 281. 

Aulacocyclus parreyi, Kuwert, 1847, p. 282. 

Localities : — 

Moluccas: Cerani. 

New Guinea : Rubi. 
.? Australia. 

Kuwert in 1897 omits ail reference to his Javanese record of 1891, so it may be 
doubted whether this species is really Oriental.' 

L* Aulacocyclus perlatus, Kaup. J 

Aulacocyclus perlatus, Kaup, i8oS«, p. 7. 
(Aulacocyclus perlatus, Gemminger and Harold, 1868, p. 909. ) 
Aulacocyclus perlatus, Kaup, 1871. p. 15. 
(Aulacocyclus perlatus, Wytsman, 1884, p. 328). 
Aulacocyclus Parreyi var. perlatus, Kuwert, 1891, p. 170. 
Aulacocyclus perlatus, Kuwert, 1897, pp. 281 and 282. 
Localities : — 
New Guinea. 

The Austrahan record rests solely on the authority of Kuwert's iSgi list; it is 
omitted in his subsequent paper. 

' There are specimens in the .Stuttgart Museum collection from Java, which perhaps belong to 
this species. 

iqi4.| F. H. Gravely . An Account of the Oriental Passu/idae. 277 

Aulacocyclus aruensis, Kuwert. 

Aulacocydus Parreyi var. tirtiensis, Kuwert, 1891, p 170. 
Aulacocyclus ^(iriiensi<;. Kuwert, 1897, p. 282. 

Localities : — 

Borneo: Sarawak.' 
Aru Islands. 

I* Aulacocyclus celebensis, Heller.] 

Aulacocyclus celebensis. Heller, 1898, pp. 22-3. 

Locality : — 

Central Celebes: Luhn-Djaladja. 

I* A. variolosus, Kuwert.] 

Aulacocyclus variolosus, Kuweit, 1897, p. 282. 
New Guinea. 

I* A. pugnax (Fauvel) ] 

*Coinacupes pugnax, Fauvel 1903, p. 359. 
Aulacocyclus pugutix, Zang, i905(', p. 22S. 

Locality : — 
New Caledonia. 

Genus ERACUPES, Kaup, 1871. 
= Aulacocyclus [part], Kaup, 1868. 
The genus Ceraciipes is only known from the K. Himalayas, Tibet, Tonkin, 
Upper Burma and Formosa. 

Ceracupes fronticornis (Westwood). 

Passalus fronticoniis, Westwood, 1842, pp. 124-5. 
Passalus bihastaius d" , Percheron, 1844, pp. 12-13, pi. cxxxv, fig. 3. 
* Passalus jronticornis, Hope, 1845. 
Passalus bihastaius, Burmeister, 1S47, p. 517 
Passalus bihzstatus , vSmith, 1852, p. 17. 
(Aulacocyclus bihastaius, Kaup, i868rt, p. b.) 
(Aulacocyclus bihastaius. Gemminger and Harold, 1868, p. 968.) 
Ceracupes fronticornis, Kaup, 1871, p. 17, pi. iii, fig. 2 za 
Ceracupes fronticornis, .Stoliczka, 1873, p. 151. 
(Ceracupes fronticornis, Wytsmau, 1884, pp. 328-9.) 
(Ceracupes Fronticornis, Kuwert, 1891, p. iji.) 
Ceracupes fronticornis, Kuwert. 1897, p. 271, 

' Kuwert has determined as A. aruensis a speciinen preserved in the Konis^liches Zoolos;isches 
Museum in Berlin, collected by Wallace in Sarawak, 

278 Memoirs of the Indian Museum. | Vol. Ill, 

Ceracupes ffronticornis [ ? part], Zang, igo^a, p. 100. ' 
CeratocHpes fronticornis, Arrow, 1907, p. 444. 
Ceracupes f fronlicornis, Gravely, above, p. 212, pi. xi, lig. 12. 
Zang refers Fruhstorfer's Nigidius-like Passalid from Ke-Lung in Formosa 
(Fruhstorfer, 1902, p. 28) to this species; but Heller has since described a Formosan 
form under a separate specific name, and it is therefore probable that it is really to 
this species that Fruhstorfer's specimens belong 
Localities: — 

K. Himalayas: Nepal. 

Darjeeling District — Gopaldhara, Rungbong \'alley. 
Tonkin : Mt. Mauson 2000-3000 ft. ; Thau Moi. 
Burma : Ruby Mines '. 

Bhamo^Sin Lum ". 
Stoliczka remarks: " It must come from the Chinese portion of eastern Tibet, for 
western Tibet has no forests." 

The specimen from Than Moi (whose altitude above sea level is not, Dr. Horn 
informs me, very great) is in the Deutsches Eutomolgisches Museum. Its horn is 
more slender and more strongly widened and turned up at the end, and its elytra 
are less strongly punctured than in the specimens I have seen from higher altitudes 
in the Darjeeling District and Mt. Mauson; but I do not feel justified in describing 
it as a new species or variety without seeing longer series of specimens from all three 

Ceracupes arrowi, Heller. 

* ? Fruhstorfer, 1902, p. 28. 
? Ceracupes fronlicornis [part], Zang, 19051?, p. 100. 

Ceracupes Arrowi, Heller, 1911, pp. 256-7, text-fig. 

Ceracupes ■\ arrowi, Gravely, 1914a, p. 31. 
Localities: — 

Fomosa: Kosempo ; Hoozan;? Ke-Lung. 

Ceracupes austeni, Stoliczka. 

Ceracupes ^austeni, Stoliczka, 1873, p. 151. 
{Ceracupes Ausfeni, Wytsman, 1884, p. 329.) 
{Ceracupes Austeni, Kuwert, 1891, p. 171 ) 

Ceracupes austeni, Kuwert, 1897, p. 274 

Ceracupes f austeni. Gravely, above, pp. 212-213, pi. xi, figs. it-iiK 

Localities: — 

E. Himalayas: Dafla Expedition, Camp 9; Dikrang Valley. 
Naga Hills: Manipur and an unrecorded locality at an altitude of about 6000 
ft. above sea level. 

' Fruhstorfer has apparently referred to the species in a price-list under the name Ceratocupes 
cornuttts (see Zang, loo. cit.). 

^ These records are based on specimens in the British Museum. 

igi_^.] F. H. Gravely : An Account of the Oriental Passalidae. 279 

Stoliczka recorded this species from the " Naga Hills, North-Easteru Districts of 
Bengal ' ' : and Wystman and Kuwert have both been content to quote Bengal as its 
locality. The Dafla and Naga Hills are situated on the Northern and Eastern 
Frontiers of Eastern Assam, and are nowhere near Bengal in its modern restricted 


Genus CYLINDROCAULUS, Fairmaire, 1880. 

Only one species of this genus is yet known ; it comes from China. 
*C. bucerus, Fairmaire. 
* Cylindrucaulus bucerus, Faiimaire, 1880, p. 164. 
Cylindrocaulus bucerus, Fairmaire, 1887, pp. 99-100. 
(Cylindrocaulus Bucerus, Kuwert, 1891, p. 173.) 
Cylindrocaulus bucerus, Kuwert, 1897, p. 304. 
Cylindrocaulus bucerus, Zang, 1905c, pp. 228-9. 
Cylindrocaulus bucerus, .\rrow, 1907, p- 446- 
Locality: — 

Western China: Sz-Tschwan— Moupin. 

Genus AURITULUS, Zang, 1905. 
This genus is only known from Japan. 

Auritulus patalis (Lewis). 

Aulacocyclus -f patalis, Lewis, 1883, p. 341, pi. xiv, figs. 6-7. 
Auritulus patalis, Zang, 1905c, pp. 328-9. 

.4 uriku 


us-fpataiis, Arrow, 1907, p. 446. 

three or four decaying hard-wood trees of three feet girth, which 
had been blown down, and had remained five or six years under shade of 
a dense forest above Yuyama, on the western side of Ichibosayama. 

Of this sub-family only one genus of two species is known. 

Genus PLEURARIUS, Kaup, 1886. 
This genus is recorded from Sumatra and the Indian Peninsula. 

*Pleurarius pilipes, Kaup. 

Pleurarius pilipes, Kaup, 18686, pp. 1-2. 
(Fleurarius pilipes, G niininger and Harold, 1868, p. 970.) 
Pleurarius pilipes, Kaup, 1871, p. 28, pi. iv, fig. i. 
(Pleurarius pilipes, Wytsman, 1884, p. 330) 
(Pleurarius Pilipes, Kuwert, 1891, p. 179.) 
Pleurarius pilipes, Kuwert, 1898, p. 162. 
Locality: — 

Sumatra : Gambong District— Fort Mangala. 

28o Memoirs of the l)idiii)i Museum. [Voi,. Ill, 

Pleurarius brachyphyllus, Stoliczka. 

IHeururius -^ brachyphyllus, Stoliczka. 1873, pp. 152-3. 
(Pleurarius brachyphyllus, Wytsman, 1884, p. 330.) 
{Pleurarius Brachyphyllus, Kuwert, 1891, p. 179.) 
Pleurarius brachyphyllus, Kuwert, 1898, p. 162. 

Pleurarius f brachyphyllus. Gravely, above, pp. 213-214, pi. xi, tigs. 13-1311. 
Localities: — 
Travancore: Kulattupuzha (W. base of W. Ohats). 
Cochin : timber forest. 
Madras Presidencj' : Malabar. 

Nilgiri Hills. 

Anamalai Hills — Puddutottain, 4200 ft., and other un- 
recorded localities up to 5500 tt. 
S. Arcot or Chengalput — Palur. 

Subfamily .1 CERA UNA E. 

This sub-family occurs throughout the Oriental Region to which it is probably 

Genus TIBERIOIDES, Gravely, 1913 

= Acerams jpartj Kaup, r86S ; [part] Kaup, 1869; Tihcrius [part J 
Kuwert, 1896-8. 

The genus Tiber ioides is only known from the E. Himalayas and Naga Hills. 
Tiberioides kuwerti (Arrow). 

Aceraius cancrus, Kaup, i868(j, p 29. 

Aceraius cancrus. Kaup, 18686, p. 4. 
(Aceraius cancrus, Gemminger, and Harold, 1868, p. 972.) 

Basilianus cancrus, Kaup, 1871, pp. 56-7. 

Basilianus \ cancrus, Stoliczka, 1873, p, 160. 
(Basilianus cancrus, Wytsman, 1884, P- 336.) 
(Tiberius Cancrus, Kuwert, 1891, p. 164.) 

Tiberius ^cancrus, Kuwert, 1898, p. 188. 

Tiberius kuwerti. Arrow, 1906, p. 446. 

Tiberioides ^kuwerti. Gravely, above, pp. 215-216, pi. .xi, fig. 14. 

Arrow has pointed out that Kaup, who has been followed by other authors, was 
mistaken when he identified a species of the genus Basilianus as defined by him, with 
Percheron's Passalus cancrus. 
Localities: — 

E. Himalayas: Nepal. 

Darjeeling District— Peshoke. 
Assam : Naga Hills — Manipur. 

ig 14.] F. H. Gravely : An Account of the Orienfal Passalidac. 281 

Tiberioides austeni, Gravely. 

Tiberioides f austeni, Gravely, above, pp. 216-217 pi. xi, figs. 15-153. 

Localities: — 

E. Himalayas : Dafla Hills — Dikrang Valle5\ 

Abor Country—Kalek, 3800 ft. 

Tiberioides borealis (Arrow). 

Chilomazus '\ borealis, Arrow, 1906, pp. 467-S. 
Locality : — 

Assam: Naga Hills. 

Gemis EPISPHENUS, Kaup, 1871. 

Incl. Acerciius [part], Kaup, 1868; Laches ( = Chilomazus, Zang, 1905) [part] + 

Basilianus [part], Kaup, 1871.' 

This genus is confined to the Indian Peninsula and Ceylon. 
Episphenus moorei, Kaup. 

Episphenus Moorei, Kaup, 1871, p. 45. 
{Episphenus Moorei, Wystman, 1884, p 334.) 
(Episphenus Moorei, Kuwert, 1891, p. 165.) 

Episphenus ■f moorei, Kuwert, 1898, p. 189 

Episphenus f moorei, Zang. r905rt, p. 163. 

Episphenus f moorei + f pearsnni, Gravely, above, pp. 217-218, pi. xi, bgs. 16-17. 

Localities : — 

Ceylon: Central Province — Nitre Cave district, c. 1800-3000 ft.; PatipoUa ; 
Ratnapura District — Bulutota. 

The series of specimens in Berlin and the British Museum prove conclusivelv 
that the characters by which I supposed E. pearsoni to be distinguished from 
E. moorei (see above, loc. cit.) have no real taxonomic value. 

Episphenus comptoni (Kaup). 

Aceraius Comptoni, Kaup, 1868a, p. 28. 
Aceraius Comptoni, Kaup, 18686, pp. 3-4. 
{Aceraius Comptoni, Gemminger and Harold. 1868, p. 972.) 
Laches Comptonii, Kaup, 1871, pp. 49-50, pi. iv, figs. 5-51?. 
(Laches Comptonii, Wytsman, 1884, p. 335.) 

Laches Comptonii + Flachii + Frustorferi + Puella, Kuwert, r'^91, p. 167. 
Laches parallelogrammifrons + fruhstorferi, + puella + flachi + sociits + comptoni. Kuwert, 

1898, pp. 338-9. 
Laches -^ Comptoni, Zang, 1905a, p. 163. 
Episphenus f comptoni + var. f flachi. Gravely, above, pp. 218-219, pi. xi, figs. 18-193. 

See Appendix I, p. 316. 

282 Memoirs of till' I ndiiiu Museum. \\o\.. Ill, 

Zang has shown that Kuwert's species are all of them identical with C. cumptoni , 
Kaup ; and the specimens I have seen confirm this . 

Localities: — 

Ceylon: Central Province — Hakgalla; PatipoUa c. 6000 ft. ; Namunakuli Hill 
Uva — Haputale. 
Ratnapura District -Bulutota. 

Episphenus indicus (StoHczka). 

Acerains Cantori [part] + lu-cli^hiirinisis f? part], Kaup. i868rt, pp. 28 and 30. 
Acerains Canton [part] + neelgherriensis [? part], Kaup, 18686, p. 4. 
{Aceraius Cantori [part] + neelgherriensis, [part] Gemminger and Harold, 186S, p. 972.) 
Basilianiis neelgherriensis [? part], Kaup, 187 1, pp. 55-11. 
Basilia)ius 1[ indicus, Stoliczka, 1873, pp. 159-160. 
(Basilianus neelgherriensis [part] ^indicus, Wvtsman, 1884, p. 33O.) 
(Basilianus Indicus + Certns, Kuwert, 1891, p. 164.) 

Basilianus indicus + stoliczkae + neelgherriensis. Kuwert, 1898, pj). 340-341. 
Basilianus indicus, Zang, 1905c, p. 223. 

Basilianus ■\ certns + stoliczkae + 'f intliciis , ZRi\g., igo6a, pp. 180-3. 
Episphenus t indicus, Gravely, above, pp. 220-222, pi. xi, figs. 20-206. 
The great range of variation found in this species appears to have been fully 
recognized by Kaup, for it is evident from the measurements he gives for his Basili- 
anus neelgherriensis that he must have referred to that species specimens of the present 
one. The localities which he quoted for his B. cantori in 1868, show that he confused 
certain specimens of Episphenus indicus with that species also. Kuwert's B. sto/iczkae 
and B. indicus cannot be recognized as distinct species; and his B. neelgherriensis is 
doubtless a worn specimen of the same. Kuwert's B certus of 1891 appears to be 
identical with Episphenus indicus, while that of his i8q8 paper is identical with E. 
neelgherriensis ; that they are not identical one with another has already been indicated 
by Zang in his synonymy of iqo6. 
Localities : — 

Bombay Presidency : N. Kanara. 
Mysore: Bababudin Hills, 4000-5000 ft. 
? Cochin. 
Travancore : High Range. 

Madras Presidency : Anamalai Hills— Puddutottam, 4200 ft.; Monica Estate. 
4000 ft. ; other unrecorded locaHties up to 5500 ft. 

' While this paper was still in the press I was able myself to examine Zatig's series iu Berlin. It 
contains a s[-ecinien 38 mm. long, so the only distinction found between /:. comptoni var. flachi (32-37 mm.) 
and £. comptoni, s. str. (39-42 mm.) breaks down as was expected (see above, p. 220). In this series the 
three first lamellae of the antennae are markedly shorter than the last three in small specimens and the 
two first than the last four in large ones. This character is not, however, quite perfectly correlated with 
size; and further, the series includes a specimen with antennae exactly intermediate between the two 
extreme forms. The degree of asymmetry found in different specimens varies greatly, and may be very 
much greater than is shown in my figure. 

I9I4-] F- H. Gravely : An Account of the Oriental Passulidac. 283 

Nilgiri Hills — Ootacamund ; Gudalur ; Naduvatum. 

Madura — Dindigal. 



Episphenus neelgherriensis (Percherou). 

Passalns neelgherriensis, Guerin-Meneville in Percherou,' 1841. p. 4, pi. Ixxvii, fig. I. 

Passalus neelgherriensis, Burmeister 1847, p. 469. 
{Passaltis neelgherriensis, Smith, 1852, p. 6.) 
? Aceraius neelgherriensis [part], Kaup, i868«, p. 30. 

Aceraius'\ infantilis + f puerilis + ? neelgherriensis [part , Kaup, 18686, pp. 4 and b. 
(Aceraius infantilis + puerilis + melgherriensis [part], Gemminger and Harold, 1868, p. 972.) 

Laches ■\ infantilis + f puerilis + ? Basilianus neelgherriensis [part], Kaup, 1871, pp. 48 and 55-6. 
(Laches infantilis ^- puerilis ^Basilianus neelgherriensis [part], Wytsnian, 1884, pp. 335-6-) 
(Basilianus Neelgherriensis + Laches Puerilis + Epilaches Infantilis. Kuwert, 1891, pp. 164 and 167.) 

Analaches f filius + infantilis + Laches puerilis + Basilianus inaequalis + certus, Kuwert, 1898, 
PP- 336, 338 and 341. 

Basilianus ^binominis, Zang, 1905a, pp. 243-4. 

Basilianus ^neelgherriensis, Zang, 1905c, p. 223. 

Basilianus ^neelgherriensis + ^binominis, Zang, 1906, pp. 179-183. 

Episphenus "f neelgherriensis, Gravely, above pp. 222-223, pi. xi, ligs. 2i-2ii(. 

The measurements given by Kaup indicate that he has confused with this species 
some large form (doubtless E. indicus). Although E. neelgherriensis is less variable 
in size than the preceding, it is sufficiently variable in other respects to have received 
several distinct generic and specific names, based on characters which are proved by 
the series before me to be of no taxonomic value. 

The name Analaches filius, Kuwert 1898, should read Epilaches filius ; for the 
generic names Analaches and Epilaches, Kuwert 1891, have been transposed by their 
author in his 1898 paper, as Zang has already pointed out (1905, p. 24). Epilaches 
filius, Kuwert, of which I have seen specimens determined by Kuwert himself, is 
identical with Episphenus neelgherriensis ; and so are Epilaches infantilis (Kaup) and 
Lacltes puerilis, Kaup, of which I have seen the types. Of two specimens of the 
former species from Mniszech's collection one proved to be Episphenus neelgherriensis 
and the other a species of Cetejus. 

L,ocalities : — 

Madras Presidency : Coorg. 


Nilgiri Hills— Coonoor, c. 6000 ft.' ; Ootacamvmd, 7800 ft. 

Annamalai Hills— various localities ranging from about 

3000-5500 ft. ' 
Madura — Dindigal. 

See above, p. 270, footnote 3. 

For more detailed reference to localities round Coonoor and in the Annanialais see above, p. 223. 

284 Memoirs of the Indian Museum. [Vol. Ill, 

Mysore : Bababudin Hills, 4000-5000 ft. 


Travancore — High Range. 

Kuwert's record of " Busilianus inacqualis " from Malacca doubtless refers to the 
true maequalis, and not to the species which he described under that name from speci- 
mens which clearly must have come from Dindigal. The localities given for the types 
of Laches infantilis and puerilis, Kaup — Vanicoro and Aru — can hardly be correct. 

Genus OPHRYGONIUS, Zang, 1904. 
Incl. Basilianiis [part| Kaup, 1871.' 
This genus probably occurs throughout the Oriental Region east of the mouths 
of the Ganges and Brahmaputra, but it has not yet been recorded with certainty from 
the Piiilippines. It is not known from the Andamans or Nicobars. 

Ophrygonius cantori, Percheron. 

Piissutus cantori, Percheron, 1844, pp. 3-4, pi. cx.Kxiv, fig. 2. 

Passaliis Cantori, Biirmeister, 1847, pp. 46S-9. 
{Passalus Canlori, Smith, 1852, p. 7.) 

Aceraius Cantori [part], Kaup, 1868a, p. 28. 

Aceraius Cantori [.' part], Kaup, 1868/;, p. 4. 
(Aceraius Cantori, Geraminger and Harold, 1868, p. 972.) 

Basilianus Cantori [part], Kaup, 1871, p. 57. 

Basilianus f Cantoris, Stoliczka, 1873, p. 159. 
{Basilianus Cantori [part], Wytsman, 1884, p. 336.) 
(Basilianus Cantori, Kuwert, 1891, p. 164.) 

Basilianus cantori + ^ intcrrogationus, Kuwert, 1898, pp. 340-r. 

Basilianus ^ convexifrons , Zang, 1904a, p. 698 footnote. 
(Basilianus cantori. Arrow, 1907, p. 445.) 

Basilianus ^Cantori + ^ convexifrons, Zang, 1903, pp. 178-183. 

Ophrygonius 1; canlori + subspp. -^ convexifrons + ^ ditnsiriensis. Gravely, above, pp. 224-225, 
pi. xi, figs. 22-22«. 

Kaup records this species from Malabar, Malacca, and Cambodia, and as 
it has otherwise never been recorded from any locality outside the E. Himalayas, 
Assam and Burma the probability is very strong that he has confused several species 
under one name. 0. convexifrons (Zang) at present seems to be a distinct local race, 
although it differs from the typical form of 0. canton in size only; the convexity of 
the frontal area, by which Zang distinguishes it, is a variable character found in 
some specimens of both forms. 
Localities : — 

I. O. CANTORI, Percheron, s. str. 
E. Hinralayas: Darjeeling District — Tista ; Sureil ; Kurseong ; Gopaldhara, 
Rungbong Valley. 

See Appendix I, p. 3i(), below. 

1914-] F. H. Gravely : An Account of the Oriental Passalidae. 285 

Sikkim — Gantok . 

Dafla Hills— Dikrang Valley. 
Naga Hills : Manipur. 

2. Subsp. CONVEXIFRONS, Zang. • • 
Assam: Khasi Hills— Shillong ; Teriaghat. 
Burma: Ruby Mines; Sin Lum, Bhamo, 6000 ft. 

3. Subsp. dunsiricnsis, Gravely. 
Assam : Dunsiri Valley. 

Ophrygonius birmanicus, Gravely. 

? Basilianm canlori [part], Kaup, 1871, p. 57. 
Ophrvgonius ^birmanicus, Gravely, above, p. 226, text-fig. 3A. 
The specimen from Cambodia which Kaup identified as 0. cantor i, may very 
possibly have belonged to this species; but his specimen from the Malay Peninsula 
can scarcely, I think, have done so, in view of the great difference that exists 
between the Passahd f luiia of this peninsula and that of the rest of the continent 
of Asia. 

Localities : — 

Burma : Ruby Mines. 
? Cambodia. 
?? Malay Peninsula. 

Ophrygonius singapurae, Gravely. 

?? Basilianus canton [part], Kaup, 1871, p. 57. 

Ophrygonius -f singapurae, Gravely, above, pp. 226-227, text-fig. 3B. 
It is diffic alt to identify with this species the specimen from the Malay Penin- 
sula which Kaup referred to 0. cantori ; for in his description of that species he notes 
particularly the shortness of the antennal lamellae. No other species of Ophrygonius 
is yet, however, known from the Malay Peninsula except 0. inaequalis, which he him- 
self recognizes as distinct. 
Locality : — 

Malay Peninsula : Singapore. 

Ophrygonius inaequalis (Burmeister). 

Passalus inaequalis, Burmeister, 1847, p. 468. 

Passalus ^oroleius ', Smith, 1852, pp. 17-18, pi. I, fig. 4- 

I The type specimen of Opltrygonius oroleius has elytra with strongly punctured lateral grooves ; 
but the puncture? are round, rather than transversely linear as .in the typical 0. inaequalis from the 
Sunda Islands. The British Museum collection contains a series of exactly similar specimens from 
Penang, and also two or three specimens transitional between the two forms; so o^'oZems ought perhaps 
to be recognized as a variety. 

2^6 Memoirs of the Indian Museum. [Vot.. Ill, 

(Leplaulax orolims + Aceraius inaequalis, Geiniuir.^cr ami Harold, i86S, pp. 970 and 972.) 

Leplaulax orolieus, Kaup, 1869, p. 40. 

Basilianus inaequalis. Kaup, iiS7i, p. 56. 

Basilianus ^inaequalis, Stoliczka, 1873, p. 159. 
(Basilianus inaequalis, VVytsman, 1884, p. 336.) 
(Basilianus inaequalis, Kuvvert, 1891, p. 164.) 

Basilianus ^sinkepicus, Kuvvert, 1898, pp. 339-340. 
(Basilianus inaequalis, Zang, 19036, pp. 418-9.) 

Ophrygonius ^quadrifer, Zaiig. 1904a, pp. 697-700, text-fia;. 3. 

Ophryoniwi ^inaequalis, Zau^, 1905a, p. 192. 

Basilianus sinkepicm, Zaiig, 1906a, pp. 177-1.S3. 

Ophrygonius ^inaequalis, Gravely, above, pp. 227-22S, pi. xii, Ggs. 24-2411. 

Kuwert gives 36 mm. as the leagth of his Basilianus sinkepicus ; but I have seen 
specimens identified by himself, and found the biggest of them to be only 27 mm. 

I/OCalities: — 

Malay Peninsula: Penang ; Perak; Johore; Singapore. 
Sumatran Islands: Sinkep Island. 
Sumatra: Deli. 
? Philippines. 

The sub-family to which this species belongs is so abundant in the Oriental 
Region right up to Wallace's Line and seems to stop so abruptly there, that I am 
incHned to regard Burmeister's " MoUucca" record as a mis]:)rint for " Malacca " (see 
also Kaup, 1871, p. 56). 

Genus ACERAIUS, Kaup (1868) 1871. 
= Ocythoe [partj, Castelnau, 1850; incl. Hdcrochilus, Kuwert, i8()6 = Rhipsaspis, 

Zang, 1905. 

Kuwert 's classification of this genus is extremely unsatisfactory, for it is based 
primarily on variable characters. The order in which the species are arranged here is 
based primarily on the form of the upper margin of the left mandible which , though 
not always quite constant, is the group character that I have been able to 

The genus is common throughout the Oriental Region east of the mouths of the 
Ganges and Brahmaputra, and is probably not found elsewhere. It is doubtful 
whether it occurs in the Andamans or Nicobars. 

Aceraius wallacei (Kuwert). 

(Heterochilus crinilTts, Kuwert. 1^91, p. 166.) 

Heterochilus ^wallacei + crinilus -t- oculitesselatus, Kuwert, 1898, pp. 3,;4-5. 
Heterochilus ^wallacei. Zang, 1905a, p. 167. 
Aceraius ^wallacei. Gravely, above, pp. 228-229, pi. xii, ligs. 26-26/'. 

igi_l ] F. H. Gravely : An Account of the Oyiental Passalidae. 287 

Localities : — 

Malay Peninsula. 
Sumatran Islands : Nias. 

Svimatra — Somgei Lalah in Indragiri. 
Borneo: Sarawak — Kuching. 

British N. Borneo— Mt. Kina-Balu, c. 5000 ft. 

Aceraius perakensis, Kuwert. 

Aceniius perakensis, Kuwert, 1898, p. 308. 
Aceraim f perakensis, Gravely, above, p. 229. 
lyocalities : — 

Malay Peninsula; S. Perak (Telom, 4000 ft.); Taiping 4000-4500 ft.'; Larut 

Hills, 3300-4300 ft.' 
vSumatra : Mt. Singaleng. 

Aceraius mbschleri, Kuwert. 
Aceraius moschleri, Kuwert, 1891, p. 163. 
Aceraius moschleri, Kuwert, i8g8, p. 344. 
Aceraius ^mdschb.-ri. Gravely, above, pp. 229-230, pi. xu, tig. 33. 

Localities : — 

Malay Peninsula : Singapore. 

Sumatran Islands : Linga ; Sinkep ; Sumatra ; Nias. 


Aceraius illegalius, Kuwert. 

Aceraius illegalis, Kuwert, i8qi. p. •[63. 
Aceraius illegalis, Kuwert, 1893, p. 345- 
Aceraius ^illegalis. Gravely, above, p. 230, pi xii, figs. 32-.^^"- 

Localities : — 

Malay Peninsula : Johore ; Penang. 

Borneo : Kuching ; between Ulu Madahit and Limbang. 

Aceraius laevicollis (lUiger). 

Passalus -f laevicollis . Illiger, 1800, p. 103. 
Passaius emarginatus, Weber, 1801, pp. 81-2. 
Passalus emarginatus , Fabricius, 1801, pp. 255-6. 
*Passalus laevicollis, Schoaherr, 1806-17, I (3) p. 333- 
*Passalus laevicollis. Dejeau, 1837, pp 194 and 501. 
Passalus emarginatus, Smith, 1852, p. 6. 

Aceraius emarginatus [part], Gemminger and Harold, 1868, p. 972. 
Aceraius f emarginatus [part], Stoliczka, 1873, p. 158. 
(Aceraius emarginatus [part], Wytsmau, 1884, p 336)- 
Aceraius Meyeri, Kuwert, 1891, p 163 
Aceraius f meyeri + f pontifex f -fluzonicus, Kuwert, 1898, p. 345- 

Aceraius ^laevicollis, Gravely, above, pp. 230-23:, pi. xii, figs 2y-2ya. 

~ ' Dr. Aunandale informs tne that there is no hill oFa^tliing like this altitude in the immediate 
neighbourhood of the town of Taiping. 

' This record is based on specimens in the British Museum. 

288 Memoirs of the Indian Museum. [Voi,. Ill, 

.The species is a very variable one and in each of three, at least, of the series 
before me (from Borneo, Sinkep Island, and Johore) I find forms identical with 
Kuwert's species meyeri, pontifex, and luzonicus together with all intermediate forms. 
Localities: — 

Malay Peninsula: Perak, Johore. 

Sumatran Islands: Sumatra (Deli, Somgei Lalah, vSerdang, Peinan and Solok); 

Nias ; Ivingga ; Sinkep Island ; Billiton. 

Borneo: Kuching ; Limbong ; Busa ; Serai; Tandjong; Bandjermasin. 
Banguey Island. 

Philippines: Luzon; S.Palawan. 
I have also seen two specimens in the Deutsches Entomologisches Museum col- 
lection labelled "Assam," and one in the British Museum labelled " Nepal," but 
these records are scarcely credible, as the species appears to be confined to the Malay 
Peninsula, the Sunda Islands, and the Philippines. 

Aceraius grandis (Burmeister). . 

Passalus emarginatus, Percheron, 1835, pp. 21-2, pi. ii, fia;. i. 

Passalus emarginatus, Percheron, 1841, p. 3. 

Passalus grandis, Burmeister, 1847, p. 463. 

Ocythoe emarginatus, Castelnau, 1850, p. 179. 

Passalus grandis. Smith, 1852, p. 5. 

Aceraius grandis, Kaup, i868rt, p. 27. 

Aceraius grandis, Kaup, 18686, p. 3. 
(Aceraius grandis, Gemminger and Harold, i868. p. qyz). 

Aceraius ^grandi-, Stoliczka, 1873, p 158. 
{Aceraius grandis, Wytsman, 1884, p. 336). 

Aceraius Addendus i- Rectidens -i- Hirsutus ^ Ceylonicns, Kuwert, 1891, p. 163. 

Aceraius addendus + f grandis + fminutifrons + prosternisulcntns + ceylnniciis + fhirsiitii'i f ^rectidens 
+ t'««g«"5 'r chinensis ■{■ molossHS , Kuwert, 1898, pp. 342-4. 

Aceraius \grandis, Zang, 19046, pp. 184-5. 

Aceraius ^hirsutus, Zang, i()0$a, -p 104. 

Aceraeus ^falawanus, Zang, 1905^, pp. 236-238. 

Aceraius ^grandis. Gravely, 1914a, p. 31. 

Aceraius ^ gravdis + vzii. ^ rectidens ■¥ ?,\\h. sp. -f hirsutus. firavely. abo\-e. pp. 2.',i-2,',3, pi. xii, 
figs. 28-30. 

Zang (1904ft) has shown that A. addendus and minuiifrons are identical with .4. 
^mn:/«s, at the same time suggesting that .4. rectidens, mapius and molussus might 
also prove to be identical with this species. The three last named are probably all 
identical with the var. rectidens described abo\-e, which alone of them seems to me to 
be sufficiently distinct for recognition, 

Kuwert's description of the elytra of A. chinensis makes it clear that this 
form is not identical with the north Indian race hirsutus ; and it seems improb- 
able that A. grandis, s. str., can occur in S. China, in view of the fact that hirsutus 
is known to occur in Tonkin and Hainan, and so presumably occurs all along the 

igi4.] F- H. Gravely : An Account of the Oriental Passalidae. 289 

hill ranges of Yunnan and the southern Chinese frontier. Unless there has been some 
mistake about the locality record , therefore, A . chincnsis must provisionally be regarded 
a distinct race. 

Kuwert's hirsntus and ceylonicus are undoubtedly the same, as Zang has already 
suggested (1905^, p. 104). Specimens of this northern race with the process of the 
left anterior angle of the head fully developed, and directed forwards or a little 
outwards, are not common; but they do occur in the series before me, and they are 
not sharply separated from others in which the process is directed inwards ; so I am 
unable to recognize them even as a definite variety. 

Localities : — 

I. Sub sp. HiRSUTUS, Kuwert. 

E. Himalayas: DarjeeHng District ^Gopaldhara, Rungbong Valley. 

Dafia Hills— Dikrang Valley; Harmutti (base of hills). 
Abor Country— Kobo, 400 ft. ; Janakmukh, 600 ft. ; Rotung 
1400 ft.; Upper Rotung, 2000 ft.; Kalek 
3800 ft. 
N. Lakhimpur— Silonbari (base of hills). 
Assam : Sibsagar ; Khasi Hills ; Dunsiri Valley ; Cachar. 
Upper Burma: Cachin Cauri. 

Bhamo— Sin Lum, 6000 ft. 
Lower Burma : Amherst District— SukU, E. side of Dawna Hills, 2100 ft. 


Tonkin: Mt. Mauson 2-3000 ft. 
Hainan Island. 

Formosa : Kosempo ; Polisha ; Le-hi-ku ; Chip-Chip ; Lake Candidius ; Fuhosho ; 

Hoozan; Sokutsu (Banshoryo Distr.) ; Kankau ; Taihorinsho ; Suisharyo ; 


Philppine Islands : South Palawan. 

Kuwert's Ceylon record is hardly credible, as Zang has already pointed out 

(1905a, p. 104). The genus Aceraius seems to occur neither in Ceylon nor in the 

Indian Peninsula. 

2. A. GRANDLS, Burmeister, s. str.' 

Malay Peninsula. 

Bintang Island. 

Sumatra : Deli. 

Java: Tengger Mountain, Bankalan. 

Borneo : Bandjermasin ; N. Borneo. 
Kuwert records A. prosternisulcatns (=A. grandis, s. str.) from the Moluccas; but 
as the Aceraiinae seem to be strictly an Oriental subfamily it is probable that 
this record is based on a misreading of the word Malacca. 

See below, p. 322, last paragraph of footnote 2. 

290 Memoirs of the Indian Museum. [Vol. Ill, 

3. Var. RECTiDENS, Kuwert.' 
Malay Peninsula. 
Sumatra: Solok ; IVhiiiK tiiigjj;!. 
Borneo: Sarawak — Kucliing. 

? Southern China. 

4. Subsp. CHiNENSis, Kmvert. 

Aceraius occulidens, Zang. 

Aceraitis 1[ occulidens, Zang, 1905^1, pp. 190-1. 
Aceraius ^occulidens, Gravely, above, p. 234, text-lig. 4A. 
Locality : — 

Borneo: Mt. Kina-Balu. 

Acerais laniger, Zang. 
Aceraius ^laniger, Zang, 1905a, p. iqi-2. 
Aceraius ^laniger. Gravely, above, p. 234. 

Locality : — 

Borneo: Mt. Kina-Balu, c. 5000 ft. 

Aceraius kuwerti, Zang. 

? Aceraius emarginatus [part], Kaup, r86S(i, p. 27. 
? Aceraius emarginatus [part], Kaup, 18686, p. j. 
? Aceraius emarginatus [part], Kaup, 187 [, p. 53. 

Aceraius ^Kuwerti, Zang, 1903a, p. 339. 

Aceraius Kuwerti, Zang, 1905a, p. 189. 

Aceraius ^kuwerti, Gravely, above, p. 235, pi. xi, fig. 31. 

Specimens of each species in which the anterior angles of the head are not pro- 
minent appear to be without exception of comparatively uniform size. With the 
exception of A. tricornis, whose head is markedly peculiar, and .-1. laniger, which 
seems to be very rare, this is the only one of these species yet described which is 
large enough to be identified with the biggest of the specimens to which Kaup applied 
the name emarginatus. His other .specimens must have belonged to different smaller 
species, including perhaps .-1 . alutaceosternus. 

Localities : — 
?Java {A. emarginatus, Kaup). 
Borneo: Mount Kina-Balu, c. 5000 ft. 

Aceraius tricornis, Zang. 

.Aceraitis ■'^tricornis, Zang, 1903a, p. 339. 
Aceraius tricornis, Zang, 1905a, p. 189. 
Aceraius ^tricornis. Gravely, above, p. 235, text-fig. 4B. 
Locality : — 

Borneo: Mount Kina-Balu, c. 5000 ft. 

See below, p. 322, last paragraph of footnote 2. 

I9I4-] F- H. Gravely: An Account of the Oriental Passalidae. 291 

Aceraius laevimargo, Zang. 

Passaliis emarginalus, Burmeister, 1847, pp. 463-4. 
(Passalus cinxrginatiis, [part], Gemtnitiger and Harold, 1868, p. 9S2.) 
(Aceraius emarginatus [part], Wytsman, 1884, p. 336.) 

Aceraius ^laevimargo, Zaug, 1905^, pp. 244-5. 

Aceraius \ laevint'irgo , Gravely, above, p. 235. 

Localities : — 
Borneo: Mt. Kina-Balu, c. 5000 ft. 

Aceraius pilifer (Percheron). 

Passalus piUfer, Percheron, 1835, pp. 23-4, pi. ii, fig. 2. 

Passalus pili/er, Percheron, 1841, p. 3. 

{Aceraius emarginatus [part], Gemminger and Harold, 1868, p. 972.) 
{Aceraius emarginatus [part], Wytsman, 1884, p. 336.) 
(Aceraius pilifer, Kuwert, 1891, p. 166.) 

Aceraius emarginatus 4- f reticulaticollis + f boriiganus -1- i nicobaricus , Kuwert, 1898, pp. 346-8. 

Aceraius \ pilifer, Gravely, above, pp. 235-23'), pi. xii, fig. 35. 

The name nikobaricus Redtenbacher (1867, p. 94) ought I think to be allowed 
to drop altogether, so I insert it here for reference. Redtenbacher states that 
in his insect the head is symmetrical, and his description, until he comes to the 
elytra, seems to me to be that of Tiberius nicobaricus and not that of an Aceraius at 
all. When he comes to the elytra", however, there can be no doubt that he is 
dealing with an Aceraius as stated by Stoliczka (1873, p. 158), followed by Kuwert. 
The head and pro- and meso-thorax of a PassaHd often get detached from the re- 
mainder of the body, and there can I think be little doubt that his type was a 
mixture of two different insects. Kuwert appears to have examined a complete insect 
of the genus Aceraius which he supposed came from the Nicobars ; but in view of 
Kuwert 's frequent inaccuracies and of the fact that no other specimen of the genus 
ever seems to have been recorded either from the Andamans or from the Nicobars 
some confirmation of the locality record is desirable. 
Localities : — 
?? Nicobars. 

Java: Tjibodas, ca. 5000 ft. ; Preanger, 4-6000 ft, 

Aceraius alutaceosternus, Kuwert. 

■ } Aceraius imarginatus [part], Kaup, i868((, p. zj. 
} Aceraius emarginatus [part], Kaup, 18686, p. 3. 
? Aceraius emarginatus [part], Kaup, 1871, p. 53. 

Aceraius f emarginatus [part], Stoliczka, 1871, p. 158. 

Aceraius alutaceosternus, Kuwert, 1898, pp. 347-8. 

Aceraius ^alutaceosternus. Gravely, above, p. 236, text-fig. 4E, pi. xii, figs. 34-34a. 

292 Mt'moirs of the hidian Miiscitni. [Vol. Ill, 

Localities : — 

Malay Peninsula : Peuanj^ Hill ; Taiping, 4000-5000 ft. ; Larut Hills, 3300- 
4300 ft.' 

Aceraius himalayensis, Gravely. 

Aceraiui ■femurginutiis IpartJ, Stoliczka, 1873, p. 158. 

Aceraius ^ himalayi-nsis. Gravely, above, pp. 236-237, pi. xii, tigs. 36-36<j. 

Localities : — 

H. Himalayas: Darjeeling District. 

Dafla Hills— Dikrang Valley. 
Sikkim : Pedong. 

Abor country — Sirpo Valley, nr. Reuging. 
Assam : Naga Hills. 

Aceraius assamensis, Kuwert. 

Aceraius ■femarginaliis [part], Stoliczka. 1S75, p. 158. 
Aceraius assumensis, Kuwert, i8()8, p. 347. 

Aceraius f assamensis, Gravely, above, p. 237, pi. xii, tigs. 37 371'. 
Localities : — 
Assam : Khasi Hills — Cherra Punji. 
Naga Hills — Mauipur. 

Aceraius helferi, Kuwert. 

Aceraius Helferi, Kuwert, 1891, p. 163. 
Aceraius pilifer + helferi, Kuwert, 1898, pp. 346-7. 

Aceraius ■ftavoyanus^ + f helferi, Gravely, above, pp. 237-238, pi. xii, tigs. 38-39(i, 
Localities : — 
Upper Burma : Cariu Ghecu, 4000-5000 ft. ; Carin Asciuii Cheba 3500-4000 ft. 

Carin Cheba 3000-3500 ft. ; Ruby Mines; Sin Lum, Bhamo booo ft. 
Lower Burma : Rangoon. 

Amherst District of Tenasserim. — Dawna Hills between Misty 

Hollow and Thingannyinaung, goo-2500 ft. 
Tavoy District of Tenasserim. 
Tonkin: Mt. Mauson, 2000-3000 ft. 

Aceraius borneanus, Kaup. 

Aceraius ^borneanus, Kaup, 1871, p. 52. 
Aceraius kaupii, Kirsch in Kaup, 1871, p. 52. 
Aceraius perchcronii , Kaup, 1871, p. 53. 

' This record is based oij a specimen in the British Museum. 

* The series of specimens in the British Museum has convinced me of the identity of tavoyanus and 
helferi. It includes a number of specimens from Sin Luni and several from the Ruby Mines as well as 
some from various places previously recorded. 

igi4.] F. H. Gravely : An Account of the Oriental Passalidae. 293 

Aceraius incidens, Kirsch, 1877, p. 28. 
(Aceraius emarginatus [part], Wytstnan, 1884, p. 336.) 
(Aceraius Percheroni + Incidens + Kaiipi, Kuwert, i8gi, p. 164.) 

Aceriaius -fpercheroni ■(- f incidens + f Katipi + fwawf^s, Kuwert, 1898, pp. 348-9. 

Aceraius "fKaupii, Zang, iqo5rt, p. 112. 

Aceraius laevicoUis, Zang, igo5c, p. 225. 

Aceraius -f borneanus, Gravely, above, pp. 238-239, pi xii, figs. 25-25Z> 

The only two differences Kaup was able to find between A. percheronii and 
A. kaupii can be accounted for by supposing the latter to have been described 
originally from rubbed specimens — -especially as the denticle in the middle of the 
truncation of the left outer tubercle varies greatly in size in the specimens before me. 
That this difference in freshness is the only real difference between the two, seems to 
be indicated by the fact that in Kuwert 's description of the specimens that he identi- 
fied as A. kaupii, there is no reference to the left, but only to the right, outer 
tubercle being shorter than in .4. incidens, the only difference given between A. incidens 
and A. percheronii being one of no value whatever. A. nanus, Kuwert, also appears 
to be identical with the present species. 
Ivocalities : — 

Malay Peninsula : Perak. 

Sumatran Islands : Sumatra — Deli; Indrapura ; Tebing tiuggi ; Peinau; Lang- 
Nias — Hili Zobobo. 
Mentawei — Sipora. 
Java: Batavia. 
Borneo: Kina-Balu, c. 5000 ft.; Sampit; Tandjong; Bandjermasin ; Banguey 

Phillippines : vSouth Palawan. 

Aceraius minor, Gravely. 

Aceraius I minor. Gravely, above, p. 240, text-fig. 4D. 
Locality : — 

Malay Peninsula: Taiping, 4000-5000 ft. ; S. Perak (Telom, 4000 ft.). 

Aceraius aequidens, Gravely. 

Aceraius f aequidens. Gravely, above, p. 240, text-fig. 4E. 

Locahty : — 

Borneo: Kina-Balu. 

This sub-family is known from Celebes, and from all parts of the Oriental Region 
except the Indian Pensinsula. Only one genus can at present be recognized. 

Genus MACROLINUS, Kaup, 1868. 
Incl. Aceraius [part], Kaup, 1868; Basilianus [part], Kaup, 1871 ; Tiberius [part], 

Kuwert 1891. 

294 Memoirs of tlic Indian Museum. \\OL. Ill 

IMacrolinus duivenbodei, Kaup.J 

Macrolinus ^ Diiivtnbodei, Kaup, i868«, p. 19. 
{Macrolinus Dinvfttbodei, Gemminger and Harold, itl68, p. (jjo.) 
Macrolinus Duivenbodei, Kaup, 1871, p. 43, pi. iv, tig. 6 
(Macrolinus Duivenbodei, Wytsmau, 1884, p. 334.) 
(Macrolinus Duivenbodei, Kuvvert, 1891. p. 165.) 
Macrolinus duivenbodei, Kuwert, 1898, p. 184. 
Locality : — 

Celebes: Meuado. 

IMacrolinus urus, Heller. J 

(Macrolinus i urus, Heller, i.SqS, pp. 23-4, pi. i, fig. 26.) 

Locality : — 

Celebes; Mt. Bonthain, 3(X)()-7ooo ft. 

Macrolinus nicobaricus, Gravely. 

Basilianus ^ andamanensis [part], Stoliczka, 1873, pp. 160-1. 
Tiberius Andamanensis [part], Kuwert, 1891, p. 164. 

Macrolinus ^nicobaricus, Gravely above, pp 241-242, pi. xiii, figs. 40-40(7. 
Localities ' : — 
?? Burma : Rangoon. 

Macrolinus andamanensis (Stoliczka). 

Basilianus -f andanidnensis [part], .Stoliczka, 1873, pp. 160-1. 
(Basilianus andamanensis, Wytsuiaii, 1884, p. 336.) 
(Tiberius andamanensis [part], Kuwert, i8gi, p. 164.) 

Tiberius andamanensis, Kuwert, 1S98, p. 188. 
(Tiberius andamanensis, Zaiig, i^o^a, p. 163.) 

Macrolinus ^andamanensis. Gravely, above, p. 242, pi. xiii. tigs. 4i-4i(r. 
Localities : — 
?? Lower Burma : Amherst District — Moulmein. 

Macrolinus sikkimensis (Stoliczka). 

Basilianus^ sikkimensis,Sio\\i:7.ka., 1873, pp. 161-2. 
(Basilianus sikkimensis, Wytsman, 1884, P- .336.) 
(Tiberius sikkimensis, Kuwert, 1891, p. 164.) 

Tiberius sikkimensis, Kuwert, 1898, p. 188. 
(Tiberius sikkimensis, Zang, 1905a, p. 163.) 

Macrolinus i sikkimensis + sMhsp. ftavoyanus. Gravely, above, pp. 243 244, pi. xiii, figs. 42-4211. 

' There are specimens in Berlin labelled "Andamans, dc Roepstorflf " : but it is very unlikely, I 
think, that the species really occurs outside the Nicobars. 

IOI4.] F- H. Gravely : An Account of the Oriental Passalidae. 295 

Localities : — 

I. M. SIKKIMENSIS, Stoliczka, s. str. :— 
E. Himalayas : Darjeeling District— crr. 1500 ft., 2 miles. E. of Punkabari. 
Dafia Hills— Dikrang Valley ; Harmutti (base of hills). 
Abor Country— Kobo, 400 ft. ; Janakraukh, 600 ft.; Rotung, 
1400 ft. 
Assam : Khasi Hills. 
Naga Hills. 

2. Subsp. TAVOYANUS, Gravely : — 
Upper Burma : Carin Cheba, 3000-3500 and 4000-5000 ft. 
Lower Burma : Tenasserim— between Misty Hollow and Thingannyinaung , 

Dawna Hills, Amherst District, at various altitudes between 

900 and 2500 ft. ; Tavoy. 

Macrolinus crenatipennis, Kuwert. 

Macrolinus ^crenatipennis, Kuwert, 1898, p. 185. 
Tiberius crenatipennis , Zang, iQOSfl, p. 163. 
Macrolinus "<! crenatipennis. Gravely, above, p. 244. 

Locality : — 

• Macroiinu? rotundifrons, Kaup. 

Macrolinus rotundifrons, [? part], Kaup, 1871, pp. 44-5- 
(Macrolinus rotundifrons, Wystman, 1884, p. 334.) 
(Macrolinus rotundifrons, Kuwert, i8gi, p. 165.) 

Macrolinus rotundifrons + singhalensis, Kuwert, 1898, p. 1S5 

Tiberius ^rotundifrons + singhalensis, Zang, igosrt, p. 163. 

Macrolinus '\ rotundifrons. Gravely, above, pp. 244-245, pi- xiii, fig. 43. 

Localities : — 

Ceylon : Central Province— Kandy ; Peradeniya ; Nalanda ; Patipolla ; Gala- 
Kaup's Chinese record has never been confirmed, though Kuwert accepts it with 
a query in his 1891 catalogue, and definitely in his later work. Possibly Kaup failed to 
distinguish between this species and T. sikkimensis ; or possibly the mistake has 
arisen through some error in the labelling of Cantor's collection, to which the Chinese 
specimens belonged. 

Macrolinus waterhousei, Kaup. 

Macrolinus Waterhousei, Kaup, 1871, pp. 43-4. 
(Macrolinus Waterhousei, Wytsman, 1884, p. 334.) 
(Macrolinus Waterhousei, Kuwert, 1891, p. 165.) 

Macrolinus Waterhousei, Kuwert, i8g8, p. 185. 
(Tiberius Waterhousei, Zang, 1905a, p. 163.) 

Tiberius -f waterhousei , Gravely, above, p. 245, pi. xiii, fig. 44. 

296 Memoirs of tJie Indian Museum. [\'oi.. Ill, 

Localities : — 

Ceylon : Ratnajnira Distict — Bulutota. 

[Macrolinus sulciperfectus, Kuweit. J 

Macrolinus sukiperfectus, Kuwert, 1891, p. 165. 
Macrolinus y sukiperfectus. Kuwert, 1898, p. i8^. 

Locality : — 

South Celebes: Bonthain. 

Macrolinus weberi, Kaup. 

Macrolinus Weberi, Kaup, i868rt, p. 19. 
(Macrolinus Weberi, Gemtninger and Harold, 1868, p. 971.) 

Macrolinus Weberi, Kaup, 187 1, p. 44 
[Macrolinus Weberi, Wytsmati, 1884, p. 334.) 
(Macrolinus Weberi, Kuwert, 1891, p. 165.} 

Macrolinus ^weberi, Gravely, above, p. 245. 
Locality : — 

Philippine Islands. 

Macrolinus latipennis (Percheron). 

Fassalus marginepunctatus , Dejean, 1837, P- 194- 

Passalus latipennis, Dup. in Percheron,' 1841, pp. 8-9, pi. Ixxiii, fig. 3. 

Passalus latipennis, Burmeister, 1847, pp. 464-5. 
(Passalus latipennis, Smith, 1852, p. 6.) 

Macrolinus latipennis, Kaup, 1868a, p. 19. 
(Macrolinus latipennis, Gemminger and Harold, 1868, p. 970.) 

Macrolinus latipennis, Kaup, 1871, p. 43. 
(Macrolinus f latipennis + -f weberi, vStoliczka, 1873, p. 155.) 
(Macrolinus latipennis, Wytsman, 1884, p. 334.) 

Macrolinus latipennis, Kuwert, 1891, p. 165. 

Macrolinus ^latipennis + ab. orbatus + javanus + dissimilis + frateriius + paralkHpcnnis + hatesi + 
^weberi, Kuwert, 1898, pp. 186-7. 

Macrnlinus f latipennis, Gravely, above, pp. 245-246, pi. xiii, figs. 45-46. 

Gemminger and Harold regard marginepunctatus, Dejean, as a synonym of 
latipennis, Percheron. As the former name has priority over the latter, and has been 
accepted by nobody, it is presumably not accompanied by a description. In any 
it seems a pity to drop Percheron' s well-known name. 

I have seen specimens supposed (by Zang, I think) to belong to each of the 
species orbatus, javanus, dissimilis, parallelipcnnis and batesi, Kuwert, but I can find 
no satisfactory character by which they can be distinguished one from another A 
little cleaning showed even the maxillary palp of the so-called M. orbatus to be 
perfectly normal. And it is scarcely likely, I think, that Kuwert's types are any 
more distinct one from another than these. 

Localities: — 

See above, p. 270, footnote. 

I9I4-] F. H. Gravely : An Account of the Oriental Passalidae. 297 

Malay Peninsula : Johore. 
Sumatran Islands: Nias— Ombolata. 

Sumatra — Deli. 
Java: Buitenzorg. 
S. E. Borneo. 

Philippine Islands : S. Palawan. 
Molucca (? = Malacca). 

The principal home of this sub-family is in the islands east of Wallace's Line ; 
but certain species are found further west, one or two getting as far as Southern 



Parapelopides symmetricus, Zang. 

Parapelopidcs ■fsyynmetriciis, Zang, 1904(7., pp. 695-7, fi^s. 1-2. 

Parapelopides symmetricus, Zang, 1905a, p. 189. 

Parapelopides -f symmetricus, Gravely, above, pp. 246-247, text-fig. bA-B. 

Locality : — 

Borneo: Mt. Kina-Balu, c 5000 ft. 

Genus TRAPEZOCHILUS, Zang, 1905. 
= Eriocnemis [part], Kaup, 1871 ; = Phraortes, Kuwert, 1898. 
The genus Trapezochilus occurs in Sumatra, the Malay Peninsula, and the 
extreme south of Burma. 

Trapezochilus dorsalis (Kaup). 

Eriocnemis ■\dorsalis, Kaup, 1871, p. 41. 
(Eriocnemis dorsalis, Wytsman, 1884, p. 334.) 
(Eriocnemis Dorsalis, Kuwert, 1891, p. 168.) 

Eriocnemis dorsalis + Phraortes generosus + respectabilis, + nobilis, Kuwert, 1898, pp. 323 

& 326. 
Trapezochilus f nobilis + f respectabilis. Gravely, above, pp. 247-248, text-fig. 5C-E, pi. xiu, 

fig. 48, 
The two specimens of this species in the Indian Museum collection, and also those 
examined later in Europe, seem to indicate the identity of generosus and nobilis, 
Kuwert. One of the two Indian Museum specimens has six distinct antennal lamellae, 
though one of them is very small ; but in the other (the one figured) the anterior face 
of the fifth joint bears only a swelling whose proximal face is steeper than its distal 
one. This fact, and doubt as to the locality of the type of T. generosus,' has led me 
to use the name nobilis in part 4 of this paper in preference to generosus, although 
the original description of the latter precedes that of the former by a few lines. Since 
that part of the paper went to press I have examined a fine series of specimens from 
four localities in the Malay Peninsula, submitted to me through the kindness of P rof. 
I Kuwert states that he had only one specimen of T. generosus, but gives both Perak and Sumatra 
as its localities. 

2g8 Memoirs of the Indian Museum. |\'0i,. Ill, 

Poulton under whose care they are preserved in the Oxford Museum. These prove 
conckisively that T. nobilis and T. respectahilis are also indistinguishable from one 
another; and the type of Eriocnemis dorsa/is, Kaup, has proved to belong to same 
species. Consequently none of the names given by Kuwert in this genus can be 

Localities : — 

Lower Burma: Tenasserim — Tavoy. 

Malay Peninsula : Perak — Taiping, 40(K)-5<)()() ft. ; Telom,40()0 ft. ; Larut Hills, 
4000-4500 ft. 
E. Siamese Malay States — Nawngchik(Bukit Resar, 2500 ft. ; 
and Ban Sai Kau, base of Bukit Besar). 
Sumatra: Deli. 

(ienus GNAPHALOCNEMIS,' Heller, iqou. 
= Ocythoe [part], Castelnau, 1850' 4- Eriocnemis [partj, Kaup, 1868. 
This genus occurs throughout that part of the Oriental Region which lies south- 
east of Burma ; it is perhaps also found in the Philippines and the Moluccas. 

Eriocnemis gelon, Schaufuss (1885, pp. 187-8) has been identified by Kuwert 
with a species of Plesthenus from Macassar in Celebes, so has been omitted from tlie 
following list. 

Gnaphalocnemis simplex, Gravely. 

Gnaphalocnemis "f simplex. Gravely, above, p. 248, text-fis;. o. 
Locality : — 

Malay Peninsula : Perak. 

Gnaphalocnemis burmeisteri (Kaup). 

Eriocnemis f Burmeisteri, Kaup, 18681/, p. 22. 
(Eriocnemis Burmeisteri , Gemm'mget and Harold, r868, p. 971.) 
Eriocnemis Burmeisteri, Kaup, 1871, pp 41-2. 
t Eriocnemis Burmeisteri, Wytsman, 1884, p. 334.) 
(Eriocnemis Burmeisteri, Kuwert, 1891, p. 168.) 
Eriocnemis burmeisteri + ^ faheri , Kuwert, 1898, p. 323 
Gnaphalocnemis f burmeisteri, Gravely, above, p. 249, pi. xiii, fisa;. 4(). 
Localities : — 

vSumatra : .Somgei Lalah (Indragiri); vSoerian Platation (near Solok); Deli; 
Pedong ; Peinan; Battak Mountain ; Kepahiang ; Redjang-Lebong. 

Gnaphalocnemis monticulosus (Smith). 

Piissaliis f monticulosus. Smith, 1852, p. 6. pi. i, fii^ i. 
Eriocnemis tridens, Kaup, 1868(7, p. 2>. 

(Aceraius monticulosus, Gemminger and Harold, 1868, p. 972.) 
Eriocnemis monticulosus, Kaup, i86g, pp. 38-9. 

' In addition to the species here meationed " Pelopides " gravidus, Kuwert (1891, p. 168; and 
1898, p. 332) from .Mindanao perhaps belongs to this genus (see Zang, igoSa, p. 316, & 1905c, p 227). 

* This name has priority. But it has been so long fotgotten in this coimection that it seems a pity 
to revive it after it has been adopted for a genus of Cephalopoda. 

1914-] F- H. Gravely : An Account of the Oriental Passalidae. 299 

Eriocnemis inonticidosus + tridcns, Kaup, 1871, pp. 41-42. 
(Eriocnemis monHculosus , Stoliczka, 1873, p. i55) 
(Eriocnemis monticulosus, Wytsman, 1884, p 333.) 
(Eriocncniii, dispur + monticulosus, Kuwert, 1891, p. 16S.) 

Enocnciiiis ^ dispar + ignotus + ^ monticulosns, Kuwert, 1898, p. 324. 
{Eriocnemis iiio)ii!ciilosus, Zang, 19035, p. 419-) 

Gnuplialocnemis f monlicnlosns, Gravely, above, pp. 249-20, pi. xiii, tigs. 49-49i(. 
Localities : — 

Malay Peninsula : Penang, Tengah Mountain. 

Sumatra: Tandjong Morawa .(Serdang) ; Deli; Solok ; Lampung ; Medau. 
Borneo : Sarawak — Kuching. 

British N. Borneo — Mt. Kina-Balu. 

Gnaphalocnemis tridens (Wiedemann). 

Passaliis -f tridens, Wiedeuiaii, 1823. pp. 109-110. 
Passdlus laterisctdptiis , Perty, 1831, p. 37, tig. 2. 
Passalus tridens, Percheron, 1835, pp. 24-6, pi. ii, fig. 3. 
*Passalus ^orientalis, Dejean, 1837, p. 194. 
Passalus tridens, Percheron, 1841, p. 5. 
Passalus tridens, Burraeister, 1847, pp. 461-3. 
Ocythoe tridens, Castelnau, 1850, II, p. 170. 
(Passalus tridens, Smith, 1852, p. 6.) 
(Passalus IriJois Redtenbacher, 1867, p. 49.) 
Eriocn.ini'i Min\:cchi, Kaup, i868«, p. 22. 

(Eriocnemis M 11 1 s:.cchi + tridens, Gerriminger and Harold, i8h8, pp. 971-972.) 
Eriocnemis Mniszechi, Kaup, 1871, p 41, pi. iv, figs. 3-3«. 
(Eriocnemis ^tridens, Stoliczka, 1873, p. 155) 
(Eriocnemis tridens ■¥ Mniszechi, Wytsmau, 1884, p. 334.) 
Eriocnemis Mniszechi -^Tridens, + \raLX. Angustior Kuwert, 1891, p. 168. 
Eriocnemis mniszechi + ^tridens, + ab. angustior, Kuwert, 1898, pp. 322 and 324. 
Gnaphdlocnemis "f tridens, Zang, 19046, p. 185. 
Gnaphalocnemis tridens. Gravely, above, p. 250, pi. xiii, figs. 5o-50«. 

lyocalities : — 

Sumatra : Palembang. 

Javanese Islands : Java— Ardjoeno ; Tengger Mountain; Tjibodas, c. 5000 ft. ; 

Soekaboemi; Preanger ; East Java. 
? Moluccas: Amboina.' 

Genus GONATAS, Kaup, 1871. 
= Acei'uius [partj, Kaup, 1868; =Goiiatas [part], Kaup, 1871 ; =Gonatas, 

Kuwert, 1897. 
This genus is found in Australia, and the islands between there and Java ; it seems 
to be centred in the region east of Wallace's Line. I have simply accepted the 
opinions of previous authors in the following synonymy. 

I This record rests on Kuwert's authority. It is the only one yet made outside the Oriental 

300 Memoirs of the Indian Museum. | \'ol. Ill, 

[* Gonatas tridentatus, Kuwert.J 
Gonatus Tridentatus, Kuwert, 1891, p. 169. 
Gonatus tridentatus, Kuwert, 1898, p. 313. 

Locality : — 
New Guinea. 

[* Gonatas schellongi, Kuwert.] 

Gonatas Schellongi , Kuwert, 1891, p. 169. 
Gonatas schellongi, Kuwert, 1898, p. 314. 

Locality : — 
New Guinea. 

[* Gonatas differens, Kuwert.J 
Gonatas Schellongi var. f differens, Kuwert, 1891, p. ibS. 
Gonatas differens, Kuwert, 1898, p. 314. 

Gonatas differens [inch G. tibialis Zang, MS.], Zaufj, iqo^a, p. no. 
Localities: — 

Moluccas: Amboina. 

Aru Island. 

New Guinea : Kaiser Wilhelms Land — Finschhafeu. 

Bismark Archi])lego : New Britain. 

[* Gonatas albertisi, Kuwert.J 

Gonatas Albertisi [Yorkensis Fairm. ?J,' Kuwert, iXgi, j). 169. 
Gonatas albertisi, Kuwert, 1898, p. 314. 
Localities : — 
New Guinea. 
? Bismark Archipelago: New Britain (omitted in Kuwert' s later work). 
Australia : Queen.sland — Cape York. 

[* Gonatas major, Kuwert.J 

Gonatas Major, Kuwert, 1891, p. 169. 
Gonatus major, Kuwert, 1898, p. 314. 
Localities: — 
Aru Islands. 
New Guinea. 

[* Gonatas novacbritanniae, Kuwert. | 

Gonatas novaebrittinniac, Kuwert, 1898, {>. 314. 
Gonatas novaebritanniae , Zang, i895((, pp. iio-i. 

Locality : — 
Bismark Archipelago : New Britain — Kinigunang. 

' Kuwert gives uo reference to the place where this name was published, and I have been unable 
to trace it. 

I9I4-] F- H. Gravely : An Account of the Oriental Passalidae. 301 

Gonatas germari, Kaup. 

Aceraius Germari, Kaup, i86Srf, p. 30. 
Aceraius Germari, Kaup, i868i, p. 4. 
{Aceraius Germari, Gemminger and Harold, 1868, p. 972.) 
Gonatas Germari, Kaup, 1871, p. 51. 
Gonatas ■\ Germari, Stoliczka, 1873, p. 158. 
[Gonatas Germari, Wytsman, 1884, p. 335.) 
Gonatas Germari, Kuwert, 1891, p. 169. 
Gonatas germari + ab. brevis, Kuwert, 1898, p. 315. 
Gonatas ^germari. Gravely, above, pp. 250-251, pi. xiii, figs. 47-47^. 

Localities : — 

Moluccas : Batchian ; Ceram. 
Tern ate. 
Australia : Queensland — I^izard Islands. 

[* Gonatas cetioides, Zang.] 

Gonatas cdiuuics, Zang, 1905.-7, p. 316. 
Locality: — 

[* Gonatas peregrinus, Kuwert.] 

Gonatas peree,rinus, Kuwert, 189S, p" 315. 
Perhaps only a variety of the following species, according to Kuwert. 
Locality : — 
New Guinea. 

* G. naviculator (Percheron). 

Passalus naviculator , Percheron, 1844, pp. 1-2, pi. cxxxiv, fig. i. 
? Passalus naviculator [part], Burmeister, 1847, pp. 467-8. 

Aceraius naviculator, Kaup, 1868a, p. 31. 

Aceraius naviculator, Kaup, 18686, pp. 3 & 7. 

Aceraius naviculator, Gemminger and Harold, 1868, p. 972. 

Gonatas naviculator , Kaup, 1871, pp. 50-1. 

Gonatas naviculator, Stoliczka, 1873, pp. 157-8. 

Gonatas naviculator, Kirsch, 18776, p. 141. 

Gonatas naviculator, Kuwert, 1891, p. 169. 

Gonatas naviculator, Kuwert, 1898, p. 315. 
(Gonatus naviculator , Arrow, 1907, p. 445.) 

Localities : — 


Moluccas: Batchian; Ceram; x\mboina. 
Papuan Islands : New Guinea— Dore. 

302 Memoirs of the Indian Museum. [\'0L. HI, 


|*Gonatas altidens, Heller.] 

Gonatas nUidens, Heller, 1910, pp. 15-1O, pi. I, fiu. ij. 
Locality : — 
New Guinea, c. 3000. 


This subfamily occurs throuj^diout the tro])ical parts of the Indo- Australian region. 

Genus LEPTAULAX, Kaup, 1868. 

incl. Lcptaidacides , Zang, 1905. 

The characters used by Kuwert in his tables, especially those by which he 

divided the genus up into groups, have unfortunately proved to have, for the most 

part, no taxonomic value whatever. After eliminating the characters which are either 

worthless or of varietal value only, very little is left. In the following list I have 

grouped as best I can, with the help of the little that does remain, the names of such 

species as I am miable to recognize, under the names of such as appear to be distinct ; 

but in places the process has been little better than guess-work. Fortunately the 

principal zoogeographical results of this process are free from the doubts to which the 

synonymmy must remain open, as most of the synonyms fall under the names of 

L. dentatus or L. bicolor ,\ioth. of them species of whose wide distribution and variability 

there is abundant proof of quite a definite nature. 

The subfamily Leptaulacinae is found in all parts of the Indo-Australian region 
inhabited by Passalidae. 

Leptaulax beccarii, Kuwert. 

Leptaiilax beccarii, Kuwert, 1891, p. 189. 

Leptaulax ^beccarii, Kuwert, 1898, pp. 294-5. 

Leptaulax f hiimerosiis [part], Zang, igo$a, p. 112. 

Leptaulax f beccarii, Gravely, above, p. 251. 

LocaUty: — 

Sumatra. t i. 1 1 t- .u 

Leptaulax humerosus, Kuwert. 

Leptaulax humerosus, KiiWL-rt, i8()i, p. 189. 
Leptaulax humerosus, Kuwert, 1898, pp. 289 and 294. 
Leptaulax humerosus, Zang, 19046, p. 185. 
Leptaulax ^humerosus [part], Zang, 1905a, p. 112, 
Leptaulax humerosus, Gravely, above, pp. 251-252, pi. xiii, tig. 51 
Localities : — 

Sumatran Islands : vSumatra — Battak and vSolok Mountains. 

Java: Tengger Mt. ; Ardjoeno ; and an unrecorded locality at an altitude of 

5000 ft. 
Borneo: Sarawak — Kuching; 10 miles south of Kuching. 

iqi4-] F- H. Gravely : An Account of the Oriental Passalidae. 303 

Leptaulax anna, Zang. 

Lcptanlax ■\anna. Zang, :qo5(/. p. 316. 
Locality : — 

Leptaulax timoriensis (Percheron). 

Passaliis timoriensis, Percheron, 1841, pp 19-21, pi. Ixxviii, fig. i. 
{Leptaulax timoriensis [part], Wystiuan, 1884, p. 332.) 
?} Leptaulax dentatus var. Timorensis, Schaufuss, 1885, pp. iih-7. 
Leptaulax \glahricollis, Kuwert, 1898, pp. 293-4. 
Leptaulax timoriensis, Zang, 19050, p. 223. 

Locality : — 

Leptaulax dentatus (Fabricius). 

Passat us dentatus, Fabricius.. 1792, p 241. 

Passalus dentatus, Fabricius, i8or, p. 256. 

Passalus dentatus, Weber, 1801, p. 82-3. 

Passalus quadridentatus , 1826, Sturm, p. 182. 

Passalus dentatus, Percheron, 1835, pp. 66-7, pi. v, fig. i. 

Passalus dentatus, Percheron, 1841, p. 21. 

Passalus timoriensis + dentatus, Burmeister, 1847. pp. 473 and 477-8. 

Passalus dentatus, Castelnau, 1850, p. 178. 

Passalus Timoriensis ^- dentatus. .Smith, 1852, pp. 17 and 20. 

Passalus dentatus, Redtenbacher, 1867, p. 49. 

Passalus dentatus, Kaup, 1868a, p. 114. 
(Leptaulax dentatus, Gemminger and Harold, 1868, p. 969.) 

Leptaulax timoriensis + dentatus, Kaup, 1871, pp. 33-34- 

Leptaulax -f dentatus + -fbicolor [part] Stoliczka, 1873, p. 155. 

(Leptaulax timoriensis {gaiX\i\ + dentatus, Wystman, 1884, p. 332.) 

Leptaulax dentatus + ? var. Timorensis, Schaufuss, 1885, pp. 186-7. 

Leptaulax ? Ribhei + Saigonicus + ? Dentatus + ? var. Bornensis + ? var. Indicus + Timoriensis + 
Darjeilingi, Kuwert, 1891, pp. 188-190. 

Leptaulax ? abdominibarbatus + malitiosus + submeiius ¥ saigonicus + ^niicus + ?ab. insignis + 
timoriensis + exterris + tdarjeelingi + f celebensis + t dentatus + mixtus + inter ponendus + 't bor- 
nensis + ^ribbei + ?geminatus, Kuwert, 1898, pp. 293-300. 

Leptaulax -fmalitiosus, Zang, 19046, p 185 

Leptaulax ^ tontiinensis ^^humerosus [part], Zang 1905". PP- 102-4 and 112. 

Leptaulax ^dentatus, Gravely, 1914a, p. 31. 

Leptaulax -f dentatus + var. ^ glahrivcntris. Gravely, above, pp. 252-255, pi. xiii, fig. 52-520'- 
This species is so common, so widely distributed, and so variable that it is not sur- 
prising to find that isolated colonies or individuals have received a number of different 
specific names 

It is impossible to be sure from Kuwert' s description alone where L. abdomini- 
barbatus should be placed. The three Bornean specimens that I have identified as his 
L. anibarbis from Borneo, agree, however, with his description of that species per- 
fectly as far as it goes; and the essential characters in which L. anibarbis dm^ts 
from L. abdominibarbatus are just those in which the specimens I have identified as 

304 Memoirs of the Indian Museum. [\'ol. Ill, 

L. anibarbis differ from L. dentatus.^ Kuwert's descriptions of /,. nbhei and L. gemmi- 
natus leave it uncertain whether the specimens from which thej- were drawn up 
belong to L. dentatus or to L. novaeguincae , but as they are both larger than the 
latter species is known to become they probably belong to the former. A specimen 
determined by Kuwert'^ as the former species, which I saw in Berlin, confirms this 
suggestion. It is possible that L. dentatus var. glabrivcntns maj' be identical with 
one or more of the forms described by Kuwert , but as there is no indication of this 
in any of the descriptions I have been compelled to adopt a new name for it. 
Locahties : — 

Madras Presidency : Nr. Vizagai)atam. 
?W. Himala3^as: Mussoorie. 
R. Himalayas: Darjeeling District. 
Dafla Hills — Dikrang Valley ; Burroi (base of hills); Harmutti 

(base of hills). 
Abor Country — Kobo, 400 ft. ; Janakmukh, 600 ft. ; Rotung, 
1300-1400 ft. ; Upper Rotung, 2000 ft. ; Reuging to Rotung, 
2600 ft. ; Kalek, 3800 ft. 
Mishmi country— Beside the La-ai, a tributary of the Kalem 
Assam: Kochugarh, Goalpara District; Chaduar ; Sibsagar ; Dunsiri Valley ; 

Silonbari, base of N. L,akhimpur Hills ; E. Cachar. 
Chittagong Hill Tracts : Kapti. 
Formosa : Kosempo ; Chikutoge ; Taihorin ; Taihorinsho ; Hoozan ; Fuhosho ; 

Poli.sha ; Chi])-Chip ; vSokutsu, Banshoryo Dist. ; Suishfryo. 
Lower Burma : Pegu. 

Tenasserim— Kawkareik, Amherst District, c. 300 ft.; Misty 
Hollow to Thingannyinaung, Dawna Hills, Amherst District, 
various altitudes between goo and 2500 ft. ; Tavoy ; Mergui. 
Andaman Islands. 
French Indo-China : Tonkin— Mt. Mausson, 2000-3000 ft. 

Cochin China — Saigon. 
Malay Peninsula : Perak ; Johore; vSingapore 
Sumatran Islands : vSumatra ; Nias. 
Java : Ardjoeno. 
Borneo : Matang ; Kuching. 
Philipi)ines : ? Mindanao. 
Sumbawa '\ 

' It appears from the collection of the Deutsches Entoniologisches Museum, that Zang api)lied the 
name abdominibarbalus to a form of L. bicolor which I am unable to recognize as distinct. 
' There are specimens from this island in the Hamburg Museum. 

1914] F. H. Gravely : An Accomit of the Oriental PassaUdae. 305 

New Guinea. 

Many of the recent records of this species east of Wallace's line are appended 
to descriptions that may refer either to it or to L. novaeguineae ; so it may 
ultimately prove to be rarer and less widely distributed there than is indicated by the 
evidence at present available. The occurrence of any species of the genus in Guada- 
loup Island or Brazil seems so improbable that these two records (made by Fabricius 
and Kuwert respectively) have been omitted from the above Hst. Concerning the 
Mussoorie record see above, p. 254. 

Lcptaulax cyclotaenius, Kuwert. 

Leptaulax cyclotaenius, Kuwert, 1891, p. 188. 

Leptaulax himalayae + cyclofaenius + fimgustifroHs, Kuwert 1898, pp. 285-6. 
Leptaulax f cyclotaenius + suhsp. f himalayae, Gravely, above, pp. 255-256, pi. xiii, fig. 5^,. 
This species is very near the last, and may prove to be nothing more than a 
variety of it. 

Kuwert's description of L. angustifrons contains not even a hint of the position 
of the form to which he applied this name. I have, however, seen a specimen identi- 
fied by Kuwert as L. angustifrons, and found it to be £. cyclotaenitis , s. str. 
Localities : — 

I. ly. CYCLOTAENIUS, Kuwert, s. str. 

Malay Peninsula : Johore ; Perak ; Singapore. 

Sumatran Islands : Sinkep. 

Borneo : Sarawak— Kuching ; Braang ; Matang Road (near Kuching) ; Mt. 


2. Sub-sp. HIMALAYAE, Kuwert. 
E. Himalayas : Dafia Hills — Dikrang Valley. 

Abor Country : Renging, 2150 ft. ; Upper Renging, 2150 ft. 
Tonkin: Mt. Manson 2000-3000 ft.' 

Leptaulax macassariensis, Schaufuss. 

Leptaulax Macassariensis. Schaufuss, 1885, p. 186. 

Leptaulax Macassariensis. Kuwert, 1891, p. 189. 

Leptaulax macassariensis -^ papiia + anibarbis, Kuwert, 1898, pp. 291 and 293. 

Leptaulax macassariensis subsp. t anibarbis. Gravely, above, p. 256, pi. xiii, fig. 5 
The Bornean species which I have identified as L. anibarbis, Kuwert, agrees 
with Kuwert's description of that species as far as it goes, and does not appear to 
differ from L. macassariensis or L. papua in any constant character. The inner 
margin of the lateral area of the metasternum is slightly straighter in one of the 
specimens before me than in the other two, making the area appear slightly narrower 

There is a specimen from this locality in Berlin. 

3o6 Memoirs of the Indian Museum. [\'Oh. Ill, 

behind ; but the difference is quite a trivial one. A difference in the widths of these 
areas appears to be the only character in the least degree likely to be of importance 
between L. macassariensis and L. papua as described by Kuwert. 
Localities : — 

I. L. MACASSARIENSIS, Kuwert, s. str. 
South Celebes : Macassar. 
New Guinea. 

2. subsp. ANiBARBis, Kuwert. 
Borneo : .Sarawak, 

Leptaulax anipunctus (Zang). 

Ltptaiilacidfs ^ aiii/^iimliis, Zaiig, iqotk. pp. 234-5. 

Lepi.aiiLjx ^anipunctus. Gravely, above, pp. 256-257, pi. xiii, fig. 55. 
Localities : — 

Burmo-Chinese Frontier : Sansi Gorge, 6000 — 8000 ft. 
Upper Burma: Catchin Cauri.' 

[* Leptaulax novaeguineac, Kuwert.] 

Leptaulax Novaeguineac -^^ ? conseqmns, Kuwert, iSgr, pp. 188-9. 

Leptaulax novaeguineae + ah. morator ■^- hansermnni ^ ^. consequent, Kuwert, 1898, pp. 287-300. 

The principal differences between L. novaeguineae and L. hansemanni, as des- 
cribed by Kuwert, seem to be that the pronotum is somewhat less and the inter- 
mediate areas of the metasternum are somewhat more extensively punctured in the 
former than in the latter ; I do not think that these differences are very likely to 
prove constant. It is impossible to tell with certainty from the structural characters 
mentioned in Kuwert's descriptions whether L. batchianae , cicatrosus and consequens 
differ in any constant character either from this species or from L. bicolor ; or whether 
ribbei and geminatiis differ in any constant character either from this species or from 
L. dentatus. But the two last are much larger than any known specimen of 
L. novaeguifieae or of any of the forms I have identified with it ; so for the present 
their names are best associated with that of L. dentatus. I have moreover, seen 
specimens identified by Kuwert as L. batchianae and L. cicatrosus, all of which 
appeared to me to be L. ftjco/or; while a specimen identified by him as L. ribbei was 
L. dentatus. It is possible, I think, that all the forms here grouped together as 
L. novaeguineae, may ultimately prove to be identical with one or other of the 
dominant forms dentat^ts and bicolor. 
Localities: — 

? Batchian (L. consequens). 

New Guinea. 

There is a specimen from this locality ia the Kgl. Zool. Museum in Berli 

I0I4.] F. H. Gravely : An Account of the Oriental Passalidae. 307 

[* Leptaulax obtusidens, Kuwert.] 

Leptaulax obtusidens, Kuwert, 1891, p. 188, 
Leptaulax obtusidens, Kuwert, 1898, p. 292. 
This species is perhaps only a variety of the last. 

Locality: — 
New Guinea. 

[Leptaulax glaber (Kirsch).] 

Trichostigmits glaber,K\rsch, 1877&, pp. 139-140. 
[Trichostigmus glaber, Wytsman, 1884, p. 331.) 
Leptaulax glaber, Kuwert, 1891, p. 188. 
Leptaulax glaber, Kuwert, 1898, p. 292. 
Leptaulacides 1; glaber, Zang, 1905^, pp. 227-8. 
Leptaulacides ^ glaber ^^ piikltelhis, Arrow, 1907, pp. 466-7. 

L. piilchellits from New Guinea is apparently a distinct variety, for Arrow was 
able to compare his type with several specimens of the typical glaher. As all he actu- 
ally saw of the latter were collected by Wallace in Batchian, however, their unifor- 
mity may have been due to their being all from a single colony, and a direct com- 
parison of these and of the type of L. pulchellus with the type of L. glaber from Jobi 
is much to be desired. 
Localities: — 

MoUuccas: Batchian. 

Papuan Islands : New Guinea ; Jobi. 

Leptaulax bicolor (Fabricius). 

Passalus bicolor, Fabricius, iSoi, p. 256. 

* Passalus bicolor, Schonherr, 1806-17. 

Passalus bicolor, Percheron, 1835 pp. 69-70 ; p). v, fig. 3. 

* Passalus bicolor + innocuus, Dejean, 1837, P- I95- 
Passahis bicolor, Percheron, 1841, p. 21. 
Passalus vicinus, Hope in Percheron, 1844, p. 11. 
Passalus bicolor + vicinus, Burmeister, 1847, pp. 478-9. 

(Passalus bicolor + vicinus , Smith, 1852, pp. 17 and 20.) 
Leptaulax EschschoUzi + bicolor, Kaup, i868«, pp. 14 and 16. 
Leptaulax bicolor [part] + EschschoUzi, Gemminger and Harold, 1868, p. 969. 
Leptaulax EschschoUzi + bicolor, Kaup, 1871, pp. 323. 
Leptaulax ^bicolor [part], Stoliczka, 1873, p. 155. 
Leptaulax bicolor, Kirsch, 18776, p. 140. 

(Leptaulax EschschoUzi + bicolor [part], Wystmau, 1884, p. 332.) 
Leptaulax bicolor, Schaufuss, 1885, p. 186. 
■ Leptaulax Manillae + Malaccae+?Consequens + tBatchianae + Bicolor + ^Cicalrosus + Abdoinim- 
sculptus + Differ entispina + Incipiens + Eschscholtzii + Aurivillii -(- Separandus,' + vars. 
Medius and Maxillonotus, Kuwert, 1891, pp. 188-190. 

' Since drawing up this synonymy I have seen a specimen determined by Kuwert as I. separandus 
and found it to be I. dentatus, not L. bicolor. As this is the only instance in which I have not found 

3o8 Memoirs of the Indian Museum. [Voi,. Ill, 

Leptaulax t malaccae + manillae + dindigalensis + f batchianae + f rocpstorfi [part] + f cicalrosus + 
? co«sr^/((;«s + t subsequens + t bicolor [part] + insipiens + sumalrac + t calcuttae + abdomini- 
sculptus + t geininiis + sequens + t «''«'■ + t eschschoUzi + t aurivillii + tenasserimensis + evidens f 
■\ differenlispina + separandus' + muxiUonoiiis + (/it-rfnts + ab. divaricatus, Kuweit, 1898, 
pp 286-300. 

Leplaulacides ifrtigiilosiis + t Friihslorferi + ^ EschschoUzi ¥ ■\ barbicaiida + j Andaiiiananiin + ^pala- 
waniciis + -\anaiilax-\-'fNielnen, Zang, 1905./, pp. 100-2, 106-9, 164-7, 2324, 235-6. 246-7. 

Leplaulacides vicinus, Arrow, 1907, p. 445. 

Leplaulacides. ] papaitanus ^ ■\ analis, Zang, 1906^). 

Leplaulax ^bicolor, Gravely, 1914(1, p. 31. 

Leptaulax f iiVo/o/- + var. ti'»c««»s, Gravely, above, pp. 257-259, pi. xiii, figs. 56-56((. 

Kuwert's descriptions of L. dindigalensis and L. consequens agree as well with 
the characters of L. novaeguineae as with those of the present species ; zoogeographi- 
cal considerations, however, seem to preclude the possibility of the identity of the 
former with that species. 

The best reason for placing many of Kuwert's species here rather than under 
L. dentatiis is a purely negative one — the absence of any reference to the great width 
of the lateral punctured grooves of the elytra that is such a conspicuous feature of 
L. dentatus and its allies. This, however, is a character which Kuwert is not likely to 
have omitted when it was present ' ; and I do not think that the long list of synonyms 
that has resulted from the procedure is bigger than was to be expected in view of 
the extraordinary variability of the specimens which I have been compelled to regard, 
for the present at least, as belonging to the single species, L. btcolor, with one variety, 
vicinus. It is of course possible that some of the names given above as synonyms may 
in reality apply to distinct species, sub-species or varieties ; but I do not think this 
will ever be settled without reference to Kuwert's types, and until some one is al)le to 
approach the question with a far bigger and more representative collection of the 
genus than I have had to deal with, when it is possible that some of the above 
names may have to be revived. To attempt to distinguish such now would, I am 
convinced, only throw into yet greater confusion the nomenclature of this already 
complicated genus. 

It is not at all surprising that Zang, using to a large extent the same characters 
that Kuwert had regarded as of importance, should have described a number of new 
species. from specimens, many of which are unique or from a single collection from a 
single locality and so very likely from a single colony. 

the synonymy adopted here confirmed by such specimens determined by Kuwert as I have since seen, 
and as Kuwert's inability to recognize his own species a second time is shown by his having, applied the 
name L. roepslorfi, to specimeus of what I take to be this species and to specimens of L. bicolor in differ, 
ent boxes belonging to a single collection, I prefer in. the absence of the type, to leave the name 
separandus provisionally in the place in which his description renders it most proljable that it ought 
to go. 

' See footnote on previous page. 

^ It should, however, be pointed out here that this character is not equally strongly developed 
m all specimens of L. dentatus ; and I find that Zang has identified specimens of this species in which 
the punctures in these grooves are so little elongated as to remain almost round, with Kuwert's 
L. inaxillonolu-i and inedius, names which appear in the above list as synonyms of L. bicolor. 

I9I4-J P- H. Gravely : An Account of the Oriental Passalidae. 309 

Localities : — 
.Ceylon : Central Province — Ganiduwa ; Dimbula ; Pundaloya. 
Madras Presidency : Nilgiri Hills — Karkur Ghat, 1500 ft. ; Nadgam, 2500 ft.; 

Madura ; Dindigal. 
? Bengal : Calcutta. 
E. Himalayas: Darjeeling District. 

Daffla Hills— Dikrang Valley. 
Abor Country — Jariahmukh, 600 ft. 
Assam : Dunsiri walley; Silonbari, base of N. Lakhimpur Hills; Naga Hills. 
Lower Burma : Hanthawadi District of Pegu— Rangoon. 

Amherst District of Tenasserim — Between Sukli and Misty 
Hollow, Dawna Hills, 2100-2500 ft. ; Moulmein. 

Formosa: Kosempo ; Polisha. 
Siam . 

Tonkin : Than-Moi ; Mt. Maason, 2000-3000 ft. 
Malay Peninsula. 

Sumatran Islands: vSumatra ; vSinkep ; Nias. 
Java, 4,000 ft. 

Borneo : Sarawak— Kuching ; Matang Road ; 4th mile. Rock Road ; Lingga 
(on the Batang Lupar River). 
British N. Borneo. — ]\it. Kina-Balu. 
Philippines : Manilla; S. Palawan. 
S. Celebes. 
Moluccas : Batchian. 
Papuan Islands : New Guinea ; J obi. 
I have omitted from the above list of localities Kuwert's " ? Old Calabar," as it 
i.s almost certainly incorrect. His Calcutta specimens probably came from some 
collection here, or from imported timber, as no Passalids seem to live anywhere in the 
Indo-Gangetic Plain. 

Leptaulax roepstorfi, Kuwert. 

Leptaiilax \ planus [part], Stoliczka, 1873, p. 153. 

Leptaulax f roepstorfi [part] + planicollis, Kuwert, i8g8, pp. 28S-290. 

Leptaulax ^roepstorfi, Gravely, above^ p 260, pi. xiii, fig 57. 

Localities : — 

? Bengal : Calcutta. 

E. Himalayas : Abor County— Yambung, iioo ft. ; Rotung, 1400 ft. 

Lower Burma : Tenasserim — Misty Hollow, W. side of Dawna Hills, Amherst 

District, c. 2200 ft. ; Tavoy. . 

310 Memoirs, oi the Indian Miisc'iini. [\'OL. Ill, 

Leptaulax planus (lUiger). 

Passalus planus, Illiger, iSoo, p. 104. 
[Leptaulax hicolor [part], Gemminger and Harold, 186S, p. 969.) 

Leptaulax ] planus [part], Stoliczka, 1S73, p. 155. 
{Leptaulax hicolor [pint], Wytsman, 1884, p. ^2.) 
(Leptaulax planus [part], Kuwert, 1891, p. 188.) 

Leptaulax ^planus, Kuwert, 1898, p. 285. 

Leptaulacidcs f planus, Zang, 1904&, p. 185. 

Leptaulax if planus. Gravely, above, pp. 260-261, pi. xiii, fig. 58. 

Localities : — 

Lower Burma : Tenasserim — Tavo}-. 

Malay Peninsula ; Johore. 

Suniatran Islands : N. E. Sumatra — Serdang. 

T Sinkep Island. 

Java. ^ 

Borneo : Sarawak — Kuching ; Mujang ; Sadong ; Matang Road ; Paku. 

This genus appears to be restricted to Celebes and the Philippines. 
Trichostigmus thoreyi, Kauj). 

Leptaulax Thoreyi, Kaup, i868rt, pp. 1.5-14. 
{Leptaulax Thoreyi, Gemminger and Harold, 1868, p. 970.) 

Trichostigmus Thoreyi, Kaup, 1871, p. 31. 
(Trichostigmus Thoreyi, Wystman, 1884, p. 331.) 
(Trichostigmus Thoreyi, Kuwert, 1891, p. 187.) 

Trichostigmus Thoreyi, Kuwert, 1898, p. 283. 

Trichostigmus ■\ thoreyi, Gravely, above, p. 261. 
Locality : — 

Philippine Islands : Middle Luzon.' 

[Trichostigmus ursulus (Schaufuss).] 

Leptaulax "<[ ursulus, Schaufuss, 1885, p. 1S7. 
Trichostigmus Ursulus, Kuwert, i8gi, p. 187. 
Trichostigmus ursulus, Kuwert, 1898, p. 283. 
Locality : — 

South Celebes (according to the labels on the type specimens ; vSchaufuss gives 
no definite record). 

The above summary of our present knowledge of the geographical distribution 
of the Oriental genera of Passalidae, shows that the family occurs in the Oriental 
region chiefly in the neighbourhood of hills. There are one or two records of speci- 
mens coming from Calcutta, but as these are the only records from any part of the 
Gangetic Plain, they may safely be regarded as records of the place from which speci- 
mens were dispatched after having been caught elsewhere. Geographically, there- 
fore, the species found in the Indian Peninsula are at least as widely separated from 
' There is a specimen from this locality in the Hamburg Museum collection. 

I9I4-] F- H Gravely : Ati Account 0/ the Oriental Passalidae. 311 

those found in the Himalayas, Assam, Burma and the Malay Peninsula as from those 
found in Ceylon. Further, no Passalids are known from the dry hills of the Deccan 
or from the northern parts of the Western Ghats ; and in the Himalayas only one species 
appears to extend further west than the Darjeeling District, this being Ophrygoniits 
cantor i which has been recorded from the Dehra Dun District. 

There does not appear to be much to be learnt from the distribution of the 
subfamilies in which asymmetry does not occur. 

Of these the Aulacocyclinae are at present centred towards the east of the Indo- 
Australian area; but the occurrence of a species of Aulacocyclus in the Indian Penin- 
sula suggests that the subfamily may once have been commoner towards the west, 
and some of the genera more widely distributed, than is now the case. The genus 
Ceracupes is intermediate between the aberrant Chinese and Japanese genera Cylin- 
drocaulits and Auritulus on the one hand, and the rest of the subfamily on the other, 
not only in certain structural peculiarities, but also in locality. 

The Pleurariinae occur only in the Indian Peninsula and Sumatra — ^a di.s- 
continuous distribution which also suggests that the subfamily formerly occupied a 
more extensive area. 

The most curious features in the distribution of the Macrolininae are their 
absence, so far as is known, from the Indian Peninsula, which contrasts strangely 
with their abundance in all other parts of the Oriental Region, including Ceylon ; 
and their occurrence outside the Oriental Region only in Celebes. 

The lycptaulacinae appear to be much rarer in the Indian Peninsula and Ceylon 
(from which countries only the two- commonest and most widely distributed species 
are known) than in any other part of the Oriental Region.' 

The distribution of the Aceraiinae and Gnaphalocneminae, in which asymmetrical 
forms occur, is of much greater interest. 

The former subfamily occurs, usually in abundance, in all parts of the Oriental 
Region where any Passalids at all are to be found, except the Andamans and Nicobars. 
The distribution of the different genera belonging to it is peculiar. In Ceylon we 
find the somewhat rare sj^mmetrical species Episphenus moorei, which does not appear 
to live gregariously as only isolated examples ever seem to have been found; 
and the slightly asymmetrical, abundant, gregarious, and highly variable species 
Episphentis comptoni. In the Indian Peninsula we find two almost equally common 
but more markedly asymmetrical species of Episphenus , E. indicus, and E. neel- 
gherriensis. Tiie former has the anterior angles of the head prominent, and resembles 
Epis ^henus camp oni in its gregarious habits and its great variability in size : while 
the latter has the anterior angles of the head obtuse and not at all prominent, and 
resembles what little we know of the species of E. moorei in its less markedly 
gregarious habits and in that all specimens are of an approximately uniform size. 

On the other side of the Gangetic Plain the symmetrical genus, Tiber ioides, is 
found, as a rarity, in the north only. Throughout the whole of the E. Himalayas, 
Assam, Burma, Siam, Indo-China, the Malay Peninsula, and the Sunda and Philippine 
' See also below, p. 330. 

312 Memoirs of the Indian Museum. [Vol. Ill, 

Islands we find the genus Aceraius, of which one species, A. grandis, resembles Epis- 
phenus comptoni and EpispJieniis indicus in its gregarious habits and its wide range of 
variability, resembling or rather exceeding the latter in the prominence of the anterior 
angles of its head, and resembling the former (if not the latter also) in being the most 
markedly asymmetrical species found in the region which it inhabits. I-'inally, in the 
greater part, if not the whole of this area, we find the genus Ophrygonius, a])parently 
less common, and nearly related to the genus Aceraius. 

It will be noticed that whenever one species is markedly more abundant than 
any of its allies inhabiting the same region, it is always found to be much more vari- 
able than them in size, and to be more markedly asymmetrical, in addition to which 
it may have the anterior angles of the head more prominent. Now asymmetry, 
and the prominence of the anterior angles of the head, are both of them peculiar 
characters which probably imply a high degree of specialization in their possessor ; 
from which it will be seen that the dominant species of the sub-family Aceraiinae, in 
Ceylon and in the continental area east of the mouths of the Ganges, is in each case 
apparently the most highly specialized species of the sub-family found there, as well 
as being the most variable in size, and always gregarious in its habits. And it m^y 
be inferred that Episphenus indicus, which differs from E. neelgherriensis in having 
the anterior angles of the head prominent, in its very variable size, and in its 
somewhat more markedh^ gregarious habits, either is, or is likely to become, the 
dominant species of the subfamily in the Indian Peninsula, although at present it 
does not appear to be markedly more common than E. neelgherriensis. 

The curious processes of the canthus in Aceraius occulidens suggest that this 
species, which is very closely allied to A . grandis, is perhaps even more highl}^ special- 
ized. If this is the case the former will presumably oust the latter, in due course, 
from its position as the dominant species of the genus. But it is also possible that 
the higher specialization indicated by these processes is not of an advantageous char- 
acter, in which case A . grandis may be expected to hold its own until the appearance 
of some form better fitted to succeed it. 

To explain the distribution of the Aceraiinae one must suppose that a migration 
of more and more highly specialized forms has taken place from the east towards 
the head of the Bay of Bengal and then south-westwards towards Ceylon.' With the 
exception of the one symmetrical genus Tiberioides, which has been able to hold its 
own in the Eastern Himalayas and the far east of Assam , the symmetrical or very 
slightly asymmetrical forms have been driven into Ceylon. There they have been cut 
off from further aggression, and so have been able to perpetuate their race, and even to 
produce one species, Episphenus comptoni, having all the biological and morphological 
characteristics of a dominant form except perhaps as regards the anterior angles of 
the bead. Similarly the Indian Peninsula forms of the genus Episphenus, both of 
them more strongly asymmetrical than either of the Ceylon forms, but with the 

' The possibility of the migration having taken place in a reverse direction, the degree of speci.iliza- 
tion increasing in response to a fresh environment, is firecluded by the occurrence of species allied to 
Episphenus moorci in Australia (see below, p. 315). 


F. H. Gravely : .4?? Account of the Oriental Passalidae. 313 

lowest terminal tooth still well developed on both mandibles and the anterior 
angles of the head even when prominent not asymmetrical, have been driven 
beyond theGangetic Plain; but one of them has developed all the characteristics of a 
dominant form , except perhaps a higher degree of asymmetry than its allied competitor, 
although this competitor is still quite common. In the genera Ophry genius and 
Aceraius, which inhabit the Oriental Region west of the Bay of Bengal, Aceraius 
grandis has alone developed all the characteristics of a dominant form. It is more 
abundant than any other species of either genus; its distribution is wider; and, 
with the exception of A. occulidens, it is structurally the most highly specialized 
member of the whole subfamily. It seems likely, moreover, to be the progenitor 
of two species, now apparently in course of evolution, one in the south and the other 
in the north, to which the names A. grandis, s. str. (with rectidens as a variety) 
and A. grandis sub-sp. hirsutus, have respectively been applied above. 

The genus Aceraius is so completely united by transitional forms to the genus 
Ophrygonvus , as to render its origin therefrom almost certain. In Ophrygonius, 
although the anterior margin of the head is asymmetrical the mandibles are sym- 
metrical; whereas in all except a few (transitional) species of the ^enus Aceraius, the 
mandibles are also markedly asymmetrical ; so it is quite in keeping with what has been 
said above of the great abundance of the most highly speciahzed species found in 
different parts of the Oriental Region, to find that the genus Aceraius is much more 
abundant than the genus Ophrygonius both as regards individuals and number of 
species; and that the somewhat isolated and symmetrical genus Tiberioides, which 
occurs only in the northern portion' of the area inhabited by these genera, is less 
abundant than either. Many of these facts are illustrated and compared with similar 
ones relating to the Gnapholocneminae in text-fig. 7 (p. 314)- 

Although Palk Strait and the Gangetic Plain appear to have influenced the dis- 
tribution of the Aceraiinae in a much more striking manner than has the line separat- 
ing continental Asia from the East Indian Archipelago (including the Malay Penin- 
sula), the influence of this line can also be seen. Aceraius grandis is the only species of 
the subfamily found on both sides of this line. The northern and southern races of 
this species occur one on each side of this line towards the west; but further east the 
northern race has established itself not only in Hainan and Formosa, but also in 
the Philippine Islands, whose fauna should presumably be aUied rather to that of 
the Archipelago, and from which two representatives of this ianna— Aceraius laevicollis 
and Aceraius borneanus— have already been recorded. 

The Gnaphalocneminae, most of which are asymmetrical, appear to hold much the 
same position in the fauna oftheislandseastof the Straits of Macassar as the Aceraiinae 
do in the countries west of it. But this zoogeographical boundary has not offered 
the same difficulties to their migration, as it has to that of the Aceraiinae; for one 
whole group of Gnaphalocneminae is found in, and perhaps confined to, the Oriental 
Region; and at least one species of the genus Gonatas occurs in the Sunda Islands.' 

I G. g«mwn and Ma7;«c«/«/or have been recorded; but all the specimens I have seen labelled with 
either of these names appear to me to belong to a single species. 


Memoirs of the Indian Museum. 

[Vol. Ill, 

Text-figure 7. 

Diagram illustrating tbe evolution of the different types of asymmetry found in the Aceraiinae and 
Gnaphalocncminae,. as far as this can be done by reference to known genera, all of which are recent : and 
illustrating the geographical distribution of these genera. The only known exceptions to the distribution 
as shown here are: — (i) a species of lipiaphcnoides which lives in New (Guinea instead of Australia; (2) 
one or two species of Gonatas which occur in the Sunda Islands; (3) a species of Gnaphalociicmis 
which has been recorded from Amboina; and (4) the genus Plcsthcnus, whose anomalous distribution (in 
Australia and Celebes) calls for further study. The genus Kaupioloidcs is in some ways intermediate 
between the Australian genera and Cdejus-Anahichcs from New Guinea, in which position it is placed 
here. The head is, however, less perfectly symmetrical than in some species of the latter pair of genera ; 
and its metasternum suggests the possibility of a closer relationship with Protomococlns than is here indi- 
cated. The ])articular structures whose various modifications the diagram is designed to illustrate are 
shown by thickened lines. The figure of P/£'s//)t;«)/s is from Kaup; those of P(;/o/)!V/<'s and Tati'iis are 
from Kuwert; that of Atircliiis is partly from Kuwert, partly from memory; and that of the sym- 
metrical species of Cet ejus-. Anal aches entirely from memory. 

1914-] F. H. Gravely : An Account of the Oriental Passalidae. 315 

The general nature of the evolution and distribution of this subfamily, together 
with that of the Aceraiinae, is shown in the accompanying figure (text-fig. 7). From 
this it will be seen that the forms of Gnaphalocneminae which come nearest to the 
most primitive of the Aceraiinae' — Epispheniis nioorei from Ceylon — are Australian; 
so it seems that in the Gnaphalocneminae also, the most primitive forms have been 
replaced by more highly specialized ones in the central parts of the Indo-Australian 
area, but have survived where they are cut off from aggression by some geographical 
barrier which the latter have as yet been unable to cross. 

The only simple and symmetrical forms of Gnaphalocneminae found outside 
Australia are : (i) a single species— pectinigem , Heller — from New Guinea, of the other- 
wise purely Australian genus Episphenoides; and, (2) a few species of the closely allied 
genera Cetejus and Analaches. All the remaining extra- Australian symmetrical species 
are specialized in other ways. For instance, in the whole of the Hyperplcsthenus group 
of genera, no member of which is very highly asymmetrical, and some of which are 
perfectly symmetrical, the lateral and intermediate areas of the metasternum are fused ; 
and in addition, the central tubercle of Aiircliiis, and the antennae of Labienus, are of 
somewhat unusual form. 

A comparsion of the geographical distribution of the Aceraiinae and Gnaphalo- 
cneminae with that of the Arachnid family Thelyphonidae, is not without interest. 
The Passalidae and Thelyphonidae (of the Oriental Region at least) inhabit the damp 
jungles of more or less hilly country ; both families are absent from the Gangetic Plain ; 
and in India the northwestern boundary of the geographical range of both is approxi- 
mately the same, being dependent, in both cases probably on climatic conditions. 
Finally, in the Thelyphonidae, as in the Aceraiinae and Gnaphalocneminae, we have a 
structural index to the degree of specialization found in different species, and find that 
species are most numerous and most highly specialized in the neighbourhood of Borneo 
and Celebes, and less numerous and less highly specialized the further one goes from 
these islands in any direction.' This seems to indicate a radial pressure of highly 
specialized on less specialized forms, resulting in a radial migration, one line of which 
especially — that in a westerly direction— follows very much the same course as 
appears to have been followed by the advance of more and more highly specialized 
forms of the Passalidae now under consideration. 

The fact that this type of distribution is found in two such widely separated 
groups of Arthropods suggests that it may occur in a number of other groups also. 
And the peculiar richness of the fauna of the Malay Archipelago, which has long been 
known, and has contributed largely to the advancement of the hypothesis that insular 
conditions favour the rapid evolution of species, is evidence in favour of this sug- 
gestion. But when, as appears to be the case in the beetles now under discussion, 

' Episphenus moorei appears to be more primitive than aay species of the geaus Tiberioides. For the 
latter, although equally symmetrical, differ from all other species of the subfamily in having the frontal 
tubercles less widely separated, and so probably form a divergent line of descent (see diagram). 

* See J.A.S.B. (N.S.) VII (1911), Proceedings, pp. c.xxiii-cx.KV 

3i6 Memoirs of the Indian Museum. \\o\.. Ill, 

the more higlily specialized species have l^een able to establish themselves on all the 
islands on one side or other of " WalUace's Line ' ' and even in a few instances {Gonatas 
spp. and perhaps Gnuphalocnemis tridens) to establish themselves on both sides of this 
important boundary-, it is difficult to see why more of them have not been able to 
cross it, and why Palk Strait and Torres Strait should have formed such serious 
obstacles to migration. 


In the key to the genera of Aceraiinae given on pp. 196-7 of the present 
paper the genera Basilianus, Ophrygonms, and Aceraius were defined in accor- 
dance with the views that had previously found general acceptance. Since that part 
of the paper went to the press, however, certain new species (all of them described 
above) have been submitted to me, which completely bridge the gaps between these 
three genera as there defined. I have therefore been compelled to reconsider the limits 
of the genera, and to adopt definitions more like those used in redefining the genera 
of Gnaphalocneminae (above, pp. 199-203). Indeed, the general course of evolution 
followed by the subfamilies Aceraiinae and Gnaphalocneminae respectively has proved 
to be very much the same (see diagram, p. 314); and the characters by which the 
Aceraiinae are distinguished from the Gnaphalocneminae as a whole, are probably of 
no greater importance than those by which certain groups of the latter subfamily are 
distinguished from one another. The majority of the genera of Gnaphalocneminae 
were known to me only through Kuwert's descriptions until after the outlines of the 
classification adopted above had been irrevocably fixed so far as this paper is 
concerned. Otherwise I would have reversed the positions of the Aceraiinae 
and Macrolininae, merging the former in the Gnaphalocneminae as the Aceraius 

Not only do the genera Basilianus (old sense), Ophrygonius and Aceraius grade 
oiie into another, but the genera Episphenus, Chilomazus, and Basilianus do so also; 
for Chilomazus comptoni is even more variable than I at first supposed, and besides 
including specimens hardly more distinctly asymmetrical than Episphenus moorei, 
contains forms whose asymmetry is almost as great as that of the genus Basilianus, 
which they somewhat resemble in the structure of the anterior margin of the 

Turning now to the structure of the mandibles in these three genera, the extent to 
which the dentition of the right mandible is reduced in any species is found to be 
correlated with the extent of the asymmetry of the head, in all cases except that of 
Basilianus cantori, a species in which the head is highly asymmetrical, but the 
tnandibles scarcely more so than in . the symmetrical species Episphenus moorei. In 
this it resembles Ophrygonius inaequalis, with which it is further connected, both 
stucturally and zoogeographically, by certain of the new species already referred to. 
The gaj) hitherto supposed to exist between Basilianus cantori and the genus Aceraius 
has likewise been filled ; and the greatest gap that now remains in the Aceraiinae is 
that between Basilianus cantori from the Himalayas and Assam, and the species from 

I9I4-] P- K- Gravely : An Account of the Oriental Passalidae. 317 

the Indian Peninsula with which it has hitherto been associated. In view of which 
I have found it necessary to transfer B. cantori to the genus Ophrygonins. 

The monospecific genus Chilomazus is ahnost as closely related to the monospecific 
genus Episphenus on the one hand, and to the genus Basilianus on the other, as are 
the species Basilianus neelgherriensis and B. indicus to one another. Consequently, 
as a genus consisting of only four species cannot be regarded as inconveniently large, 
it seems best to unite all the species of Aceraiinae found in Ceylon and the Indian 
Peninsula in a single genus, for which the name Episphenus must be retained. 
The range of degrees of asymmetry found in this genus is very little greater than 
that found in each of the genera Cetejus fed Analachcs. 

The species of Aceraiinae found on ;the other side of the Gangetic Plain are, 
however, much more numerous; and, although transitional forms exist, the majority 
either have almost symmetrical mandibles, or have both the lowest terminal and 
anterior lower teeth on the right side quite rudimentary, the former tooth being normal 
and the latter enlarged on the left side. The genus Ophrygonius must therefore be 
defined so as to include all species of the former class, and the genus Aceraius so as to 
include all species of the latter. 

The latter genus appears to have been derived from the former in at least two 
different ways ; for the transitional species described above under the names Ophry- 
gonius singapurae and Aceraius wallacei seem to be allied to quite a different section 
of the genus Aceraius from that to which the transitional forms Aceraius acquidens 
and A. minor are allied. 

Definitions, based on the structvire of the mandibles, could be framed so as to 
include any of these four species in either genus ; so I have thought it best to follow 
accepted definitions as far as possible, and to use the presence or absence of hair on 
the sides of the elytra as the crucial test, although Aceraius wallacei is transitional in 
this respect also, being much less hairy than most of the species belonging to the 
genus in which I have had to place it. 

Four genera of Aceraiinae may then be recognized. They can be distinguished 
from one another as follows : — 

j Inner tubercles separated by a space J— | as long as that 
j separating outer tubercles . . 

\ Inner tubercles separated by a space j— i times as long as that 
I separating outer tubercles . . . . 

Left outer tubercle acute, and little or no larger than right ; 
or much larger and curved inwards, with its extremity rounded 
rather than truncate, and never angular on the outer side in 
front. Dentition complete in symmetrical species ; lowest ter- 
minal tooth always present on both sides; right anterior lower 
2' tooth smaller than left in the more highly asymmetrical species. 
Left outer tubercle always larger than right, directed more or 
less inwards, truncate distally, outer angle of truncation dis- 
tinct, forming a more or less forwardly directed apex to the 
tubercle; dentition complete, or both lowest terminal and 
^ anterior lower teeth reduced 

Tiberioides, Gravely, 

pp. 215 & 280. 

Episphenus, Kaup, 

pp. 217 & 281. 


Mi')uoiis oj the Iiidimi Mhscidu. 

[Vol. Ill, 

f A little hair present on elytra at shoulders oulj' ; dentition 
always complete, left anterior lower tooth rarely enlarged. 
I Elytra hairy at sides also; right lowest terminal and anterior 
I lower teeth almost always rudimentary; left anterior lower 
I tooth almost always enlarged 


Zang, pp. 224 & 284. 

Aocniiiis, Kaup, 

pp. 228 & 286- 



C. joveicollis, p. 267. 

In most of the following keys Oriental species only are dealt with. Of the 
seven genera occurring in both the Oriental and Australian Regions five (Contaciipes, 
Macrolinus, Gnaphalocnemis [?], LeptaiUax and Trichostigmus) are chiefly Oriental, 
and I have included all their species, basing my definitions of such as I have not 
seen' on the descriptions of previous writers. The genera Aulacocyclus and Gonatas, 
on the other hand, are found mostly in the Australian Region, and as I am unable 
to give any revised definitions of their species, only the Oriental forms are included 
in the keys. 

Genus Comacupes, Kaup. 

I Mesosternuni strongly punctured all over, abdominal sterna 
^ with at least a few hair-bearing punctures in fresh specimens.. .. .. 2. 

(Mesosternum unpunctured except at sides, abdomen unpunc- 
tured and hairless 
'' Lower margin of overhanging portion of central tubercle of 
head usually rather long, always horizontal, anterior part of 
upper margin descending obliquely to meet it in a more or less 
acute angle ; upper surface of same tubercle usually more or 
less distinctly grooved longitudinally, or excavate 
Lower margin of overhanging portion of central tubercle of 
head short, or ascending obliquely to meet upper margin which 
is always horizontal; tubercle usually truncate or concave 
anteriorly, not sharply pointed, narrower, keeled or rounded 

Central tubercle pedunculate (almost as in Aulacocyclus) 
Central tubercle not pedunculate 
Central tubercle broad and strongly excavate above 

Central tubercle narrow, at most longitudinally grooved above C. stoliczkae, pp 
Whole anterior end of central tubercle raised well above supra- 
orbital ridges, not truncate or concave in front . . . . C. masoni, pp. 207 & 
Central tubercle less elevated, truncate or concave in front, 

very variable . . . . C. cyclindraccus, pp. 


C. basalts, p. 267. 

C cavicornis, pp. 204 

& 268. 

207 & 

' Several species not seen in time for inclusion in the descriptive part of this paper (part 4) are 
redefined here from personal observation. The only species not so defined are those marked with an 
asterisk ( * ) in part 6. 

1914-] F. H. Gravely : An Account of the Oriental Passalidac. 319 

Genus Taenioers, Kaup. 

j' Upper surface of central tubercle about twice as long as broad, 
j flat, punctured, bordered by very distinct liorse-shoe-shaped 
^ ! ridge which is open in front .. .. .. T. bicantha/iis, pp. 208 & 270. 

I Upper surface of central tubercle relatively broader as a rule, 

less flattened, unpunctured, marginal ridge often indistinct or 
1^ absent across middle-line behind as well as in front .. .. .. 2. 

^ Anterior tibiae very broad . . .. .. .. T. platypus, p. 2yo. 

( Anterior tibiae slenderer . . . . . . . . . . , 3. 

Anterior margin of canthus meeting side of head a considerable 

distance behind anterior angle; external angle of canthus 

obtuse . . . . . . . . . . T. pygmaeus, pp. 209 & 271. 

Anterior margin of canthus meeting side of head a very short 

distance behind anterior angle; external angle of canthus 

sharper . . . . . . . . . . T. bicuspis, pp. 210 & 272 

Genus Aulacocyclus, Kaup. 

Apical portion of central tubercle not very strongly bent over 
forwards, directed a little upwards rather than downwards, the 
upper margin straight . . . . . . . . . . _ 2. 

Apical portion of central tubercle more strongly bent over for- 
wards, never upwardly directed, at least the extreme distal 
portion of the upper margin bent a little downwards . . . . . . 3. 

/Dorsal grooves of elytra unpunctured . . . . . . A. rosenhergii, p. 274. 

2' Dorsal grooves of elytra very distinctly punctured, as in the 
(^ the two following species . . . . . . ..A. dilatns, p. 276. 

j Central tubercle as seen from above not or scarcely broader in 
o j front than behind . . . . . . . . a 

\ Central tubercle seen from above much broader in front than 
[^ behin 

4) war 

d .. .. .. .. A. andrewsi, p. 2:1 & 2y^. 

Apical portion of central tubercle very short, not bent down- 
wards as a whole .. .. .. .. A. amensis,p. 2J7. 

Apical portion of central tubercle much longer, very slightly 

but distinctly bent downwards as a whole .. .. A. parryi, p. 2'^6. 

Genus Ceracupes, Kaup. 

f Apex of horn formed by fusion of central tubercle with ante- 

I rior margin of head bifid . . . . . . . . . _ 2. 

(^ Apex of this horn acute . . . . . . . . C. austeni, pp. 212 ct 278. 

( Upper surface of this horn roughly parallel-sided throughout . . C. arrowi, p. 278. 

( Uper surface of this horn much broader in front than behind C. frDnticornis pp. 212 i^ 277. 

Genus Cylindrocaulus, Fairmaire. 
Onh' one species, C. bucerns, p. 279. 

Genus Auritulus, Zang. 
Only one species, A. patalis, p. 279. 


Memoirs of the Indian Museum. 

fVoi.. Ill, 

Genus Pleurarius, Kaup. 
I have not seen a specimen of P. pilipes, and can find nothing in Kaup's descrip- 
tion and figure by which to separate it from the Indian species. P. pilipes, from 
Sumatra (above, p. 279), and P. hrachyphyllus , from India (above, pp. 213 and 280), are 
the onh' species of the genus that have yet been described. 

Genus Tiberioides, Gravely. 

Lateral grooves of elytra narrow, normally punctured 
I Lateralgiovesof elytra broad, their punctures transversely linear T. kuwerii, pp. 215 & 
protuberance on anterior margin, somewha" as 
om/)/o;n', and a stron;.^ t ransverse ridge a little 
further back .. .. .. .. T. borealis, ■p. 2d,i. 

Mentum without any such ridge or protubeiance . . . . T. aiisteni, pp. 216 «& 

( Mentum with a ] 
j in Epispheniis co 

Genus Episphenus, Kaup. 

( Anterior margin of head symmetrical ; anterior margin of men- 

, j turn not depressed or grooved 
j Anterior margin of head more or less asymmetrical ; anterior 
Vraargin of mentum more or less depressed or grooved 
( Anterior margin of head not very strongly asymmetrical as a 
I rule; anterior maigin of mentum strongly grooved on either 

I side of a strong median tubercle (occasionally paired) 
I Anterior margin of head strongly asymmetrical; mentum 
V^without any strongly marked tubercle 
( Anterior angles of head not prominent 

( Anterior angles of head more or less prominent . . 

Genus Ophrygonius, Zang. 

( At least five well developed and pubescent antennal lamellae 
' i Antennal lamellae very short, only four of them pubescent . . 

r Anterior lower tooth of both mandibles normal, conical and 
^ J acute 

j Anterior lower tooth of both mandibles very broad, more 

[^ obtuse, flattened dorso-ventrally 

I Left outer tubercle moderately stout, directed more or less 

E. moorci, pp. 217 and 281. 

E. coiiiptoni, pp. 21S & 281. 

222 & 

3' forwards 
I Left outer tubercle 

.•ery slender, directed strongly inwards 

E. neclgherriensis, pp. 

E. indicus, pp. 220 & 2S2 

O. inaeqnalis, pp. 227 & 283 

O. singapurae, pp. 226 & 285. 

0. cantori, pp. 224 & 284. 
O. birmanicus, pp. 226 & 285. 

Genus Aceraius, Kaup. 
Many of the characters by which the species of this genus are distinguished one 
from another are .somewhat variable even in fresh specimens, and are verj- often 
modified by friction. When a series of each species is available for reference their 
determination is not very difficult ; but when .single specimens are to be determined 
the difficulty is often great, and size and locality will sometimes be found to afford a 
simpler clue to their identity than this key. The full range of the variation that I 


F. H. Gravely : An Account of the Oriental Passalidae. 





have seen in each species has been carefuUy noted in the descriptions given in the 
more extended account of the genus (above pp. 228-240). 

Upper tooth of both mandibles distinct; lowest terminal 
tooth of right mandible rudimentary or absent; anterior lower 
tooth of right mandible minute or absent, that of left mandible 

always very large .. .. ■ .-. •■ •• •• 2- 

Upper tooth of both mandibles obsolete ; lowest terminal tooth 
of both mandibles normal ; anterior lower tooth of right man- 
dible well developed, that of left si le not always abnormally 


No convexity of upper margin of left mandible, behind base of 
hinder margin of simple or bifid upper tooth ; this margin con- 
cave, straight, or convex the whole way from tip of tooth 
backwards (see figs. 26a, 271?, 286, 29) 

Upper tooth of left mandible always simple, set in a hollow in 
front of a convexity of the upper margin from which it is dis- 
tinctly separated at base (see fig. 25a, pi. xii; & 4C, p. 234) 
Upper tooth of left mandible very variable ; a small denticle 
sometimes present below the apex (figs. 28a, 32), but the apex 
itself not bifid (figs. 26*, 27a, 28, 29, 30, 32, 33) except some- 
times in species in which the anterior angles of the head are 
very distinctly prolonged forwards (fig. 286) 
Anterior angles of the head obtuse, never prolonged forwards ; 
apex of upper tooth of left mandible always more or less dis- 

.tinctly bifid in unworn specimens (see especially fig. 31) 
Right outer tubercle simple, elongate, acute (figs. 26, 32, 

\ 33); anterior angles of head truncate or at most moderately 

I prolonged 
4 ; Anterior angles of head at least moderately prolonged; right 
outer tubercle usually shorter, truncate or rounded, some- 
times divided into two separate tubercles of which the outer 

^ one or both may be more or less elongate and acute 

I All six lamellae of antennae very long and slender; inner 
margin of right outer tubercle parallel to axis of body at base ; 
5 anterior angles of head not prominent 

(Proximal lamellae of antennae shorter ; inner margin of right 
outer tubercle oblique throughout 

-• Apex of right outer tubercle truncate or imperfectly bifid in 
profile, the suture distinctly bent downwards; anterior angles 
I of head not prominent ; mentum punctured sparsely or not at 
, ; all in middle (left upper tooth very variable) 

Apex of right outer tubercle simply pointed in profile, suture 
not bent downwards; anterior angles of head more or less dis- 
tinctly prominent; mentum more or less strongly punctured 

' Anterior angles of head not very prominent ; tip of right outer 
tubercle as in A. psrakensis. not b^nt outwards (fig. 33) 
Anterior angles of head more prominent; tip of right outer 
I tubercle bent somewhat abruptly outwards 

A. wallacei, pp. 228 & 286. 

A. perakensis, pp. 229 & 287. 

A. imsMeri, pp 229 & 287. 
A . illegalis, pp. 230 & 287. 


Memoirs of the Indian Miisciiiti. 

[Vol. Ill, 

Caiithus without any upwardly directed tubercle ., 
A stout erect tubercle arising from dorsal surface of canthus 
immediately in front of eye 

At most 34 mm. long ; anterior angles of head scarcely more 
prominent than in A. illegalis, somewhat variable; apparently 
confined to the Malay Peninsula and Archipelago, and the 

At least 33 nun. long, specimens from the Malay Peninsula 
and Archipelago always much bigger than this; anterior angles 
of head often strongly produced forwards, especially in small 
'^ specimen? 

(Specimens at least 43 mm. long ; right outer tubercle rounded 
or obsolete except in one species always over 47 mm. long . . 
Specimens at most 40 mm. long; right outer tubercle always 
more or less distinct and pointed 

Right outer tubercle distinct; ridge between lateral and inter- 
mediate areas of metasternum obtuse, rough; tenth rib of 
elytra punctured and hairy anteriorly . . 

Right outer tubercle more or less obsolete ; ridge between 
lateral and intermediate areas of metasternum acute, polished; 
tenth rib of elj'tra usually hairless and unpuactured 

'^Supra-orbital ridges and inner tubercles normal; right outer 
tubercle not completelj' absent 
Supra-orbital ridges with the apical angle very strongly deve- 

\ loped, and situated on inner side of, rather than behind, the 
anterior truncation; left inner tubercle situated on inner side 

l^of base of left outer tubercle ; right outer tubercle missing . . 

A. occiilidens, pp. 234 Si 290. 

A. laevicoUis, pp. 230 & 2S7. 

A. grandis'^, pp. 231 & 288. 

A. lani'^cr, pp 234 & 290. 

.1. kv.werti, pp. 235 & 290. 

A. Iriconiis, pp. 235 & 290. 

' These distinctions are much more definite in reality than might be supposed. Out of the several 
hundred specimens I have examined I have only been in doubt as to the identity of one; and as this was 
from Tonkin, where A. grandis is known to occur not infrequently, and .4. laevicoUis has never been 
found, it must I think have belonged to the former species. 

■^ Occurring under three (? or four) imperfectly differentiated forms: — 

A. Northern race, confined to continental Asia north of the Malay 

Peninsula, the Philippines, Formosa, etc. Seventh rib of 
elytra usually strongly punctured, rarely without punc- 
tures; hair on elytra always quite short in specimens from 
Assam and the Himalayas, inclined to be longer in speci- 
mens from Burma and Tonkin ; shape of head very variable sub-sp. Iiirsiiliis, Kuwert. 

B. Southern race, confined to the Malay Peninsula and Sunda 

Islands. Seventh rib of elytra never punctured : hair very 
variable in length, usually longer than in the northern 
race. Occurring in two forms :— 
(/. Anterior angle of left side of head at least moderately 

prolonged and curved inwards 
I). .'Vnterior angle of left side of head at most moderately 
prolonged, directed outwards or forwards 
?C. Chinese race with elytra like those of the southern race 
[Mr. Arrow tells me that Burmeister's type resembles recHdcns; iti 
throughout this paper called e/andis, s. str., becomes var. addendus, Kuwert J 

A. grandis, Burmeister. s. str. 

var. reclideiis, Kuwert. 
sub-sp chinensis, Kuwert 
this case the form here and 


F\ H. Gravely : An Accoitnt of the Oricntnl Passalidae. 



At least 34 mm. long; very variable in structure; right outer 
tubercle simply pointed in profile in unworn specimens ; scars 
on mesosternum almost always distinct 

IAt most 32 mm. long; less variable; right outer tubercle 
slightly truncate in profile in perfectly unworn specimens; 
scars on mesosternum obsolete 
I Posterior part of tenth and whole of eighth ribs of elytra uu- 
4 ' punctured 
I Seventh to tenth ribs of elytra (inclusive)punctured throughout 
/-Convexity of upper margin of left mandible very high and 
I strongly curved — usually much higher than in any other 
I species; left outer tubercle rather slender; right outer tuber- 
5 ; cle more or less truncate in profile 

Convexity of upper margin of left mandible moderately high 
and strongly curved ; left outer tubercle somewhat stouter ; 
right outer tubercle variable 

(Free portion of left outer tubercle distinctly longer than broad ; 
right outer tubercle often very broadly bifid in profile 
\ Free portion of left outer tubercle more or less square ; right 
I outer tubercle not very broadly bifid in profile . . 
( Right outer tubercle distinctly bifid in profile 
I Right outer tubercle at most imperfectly bifid in pro'ile 
Anterior lower tooth of left mandible very large, as in all 
preceding species of the genus, much larger than that of left 
mandible; posterior part of tenth and whole of eighth ribs of 
i8( elytra unpunctured . . ... 

Anterior lower tooth of left mandible scarcely larger than that 
of right; s.>venth to tenth ribs of elytra (inclusive) somewhat 
sparsely punctured throughout 


A. laevimargo ,^ pp. 238 & 291. 

A. pilifer, pp. 235 & 291. 


A.borneanm, pp. 23S& 292. 

A .(iliUaccoslerHiis , pp. 236 & 291. 


A. heifer i, pp. 238 & 292. 

A. assamnisis, pp. 237 & 292. 
A. hiimlayensis, pp. 236 & 292. 

A. minor, pp. 240 & 293. 
.4. aeqiddens, pp. 240 & 29J. 

M. urns, p. 294. 

M. duivenbodei, p. 294. 

Genus Macrolinus, Kaup. 

( luner tubercles, and anterior part or whole of frontal ridges, 

j obsolete 

j Frontal ridges complete and well developed, inner tubercles 

( more or less distinct 

^Apex of central tubercle approximately rectangular in profile, 

I directed upwards, not overhanging . . 

j Apex of central tubercle acute, directed forwards, somewhat 

\^ overhanging . . 

(•Ridge joining inner tubercles separated from anterior margin 

of head throughout its whole length by a more or less concave 
3 surface . . . . . . . . . . . . . . 4. 

Ridge joining inner tubercles closely approximated to anterior 
l_ margin of head either in middle or throughout .. .. .. .. 6. 

<! I,ateral grooves of elytra narrow, punctures normal . . . . . . 5. 

1, I<ateral grooves of elytra broal, punctures transversely linear M. sikkimemis, pp. 243 & 294. 

* Or the slightly larger .4. perakensis (pp. 229 & 287) in which the upper margin of the left 
mandible is very variable and may be indistinguishable from that typical of the section of the genus to 
which A. laevimargo belongs. 

324 Memoii's 0/ the Indian Miisenm. [Vol. Ill, 

/'Outer tubercles sleuder in profile, truncate; ridge joiuiiig 

! iuuer tubercles concave . . . . . . .. M. nicobaricus, pp. 241 & 294. 

j Outer tubercles stouter and distinctly bifid in profile, ridge 

I joining inner tubercles straight . . . . .. M. ^42&2g4. 

I Inuer tubercles situated distinctly l)eliind anterior margin of 

, ' bead . . . . . . . . . . , 

6 ••../. 

j Inner tubercles situated on (or vertically above) anterior margin 

Vof head . . . . . . . . . . . . _ q 

/'Grooves of elytra coarsely punctured, lateral grooves almost 
I as broad as intervening ridges, their punctures very coarse 

7 > indeed .. .. .. .. .. M.crena(ipennis,pp.2^4 & 2^)$. 

j Grooves of elytra less coarsely punctured, lateral grooves 

Vniuch narrower than intervening ridges . . . . . . . . 8. 

( Ridge between inner tubercles convex, evenly curved through- 

) out; anterior angles of pronotum strongly punctured . . .1/. rjlmi lifron-;, pp. 2 [i &2g5. 

J Ridge between inner tubercles straight throughout almost its 

(^ whole length ; anterior angles of pronotum unpmctured .. M. w i(efkoitsin, i)p. 2^^Si2g5. 
I Third lamella of antennae distinctly shorter than fourth ; 
' median groove of pronotum distinct, complete . . . . M. sHkipjrfeclus, p. 296. 

' j Third lamella of antennae not distinctly shorter than fourth ; 
1 median groove of pronotum obsolete . . . . . . . . 10. 

!Tip of second lamella not reaching line joining tips of first and 
third when antenna is furled . . . . . . M. webtri, pp. 245 & 296. 

\ Tips of all six lamellae arranged in a straight line when antenna 

I is furled .. .. .. .. .. ,1/. /((///>t')(»/s, pp. 245 & 20b. 

Genus Parapelopides, Zang. 
Only one species, P. symmelricus, pp. 246 and 297. 

Genus Trapezochilus, Zang. 
Only one species, T. dorsalis, pp. 247 and 297. 

Genus Gnaphalocnemis, Heller. 

(Left outer tubercle consisting of a single, somewhat slender, 
obliquely truncate process; right outer tubercle consisting of 
) a similar but broader and slightly bifid inner process, together 

I' with smaller pointed outer and middle processes . . G. simplex, pp. 24S & 2^8. 

Both tubercles consisting of three denticles more or less fused 
.together, the middle one sometimes obsolete . . . . . . . . 2. 

i Lateral grooves of elytra all narrow, simply punctured . . G. bimneislcn, pp 249 & 258. 

Grooves 5-7 of elytra more or less broad; each containing a 
polished flattened band which is marked by a single row of 
I punctures, and defined on either side by a more less distinct 
roughened line, with which the punctures may be to some 
I extent confluent 


F. H. GravEIvY : An Account of the Oriental Passalidae. 


Grooves 5-7 of elytra somewhat variable in width ; the poste- 
rior part of groove 8 rarely' wider than the anterior part, 
3 ' never as wide as groove 7 . . 

(^.-.— — - •— c / •• ■■ • •• G. ;«o«<tc«/osHS, pp. 249 &29S. 

Grooves 5-7 of elytra always very broad; the posterior part 
of groove 8 like them . . . . . . .. G. tridens, pp. 250 & 299. 

It is possible that " Pelopides'.' gyavidus, Kuwert, whose position is uncertain, 
may belong to this genus. It diiTers from all the species included in the key in having 
the left outer tubercle more strongly developed than the right. 

Genus Gonatas, Kaup. 
Only two species of this genus are recorded from the Oriental Region. I have 
only seen one of them. 

( heit mandible less elongated and more curved ; lamellae of 
antennae longer and slenderer . . . . .. G. gennari, pp. 250 & joi. 

] Left mandible longer and less curved, lamellae of antennae 

(^shorter and stouter .. .. .. .. G. naviciilatoy, p. ^oi. 

Genus Leptaulax, Kaup. 
j Depressed surface of two or three outermost grooves of elytra 

dull, the sculpturing somewhat worn-looking 
I Elytra polished throughout • 
A more or less distinct polished tubercle formed out of each 
of the transverse ridges in lateral grooves of elytra 
No such tubercles present 

Pronotum flatenned, densely punctured laterally, sides practi- 
cally straight . . 

Pronotum convex; punctured only in marginal groove, in and 
close round scars, and (usually) in anterior angles; sides dis- 
tinctly curved . . 

Inner tubercles at most twice as far from one another as from 
outer tubercles ; puncturing of lateral grooves of elytra some- 
what variable, but normally transverse to a marked degree in all 
species ; intermediate areas of metasternum closely punctured, 
j uupunctured band along outer margin absent or quite narrow 
) Inner tubercles at least twice as far from one another as from 
outer tubercles; puncturing of lateral grooves of elytra not 
markedly transverse; intermediate areas of metasternum often 
less closely punctured, and always with an unpunctured band 
\ along the outer margin 

/Inner and outer tubercles approximately equal, the former at 
I most twice as far from one another as from the latter 
( Inner tubercles much longer than outer, the later almost 
j obsolete ; the former at least twice as far from one another as 
(^ from the latter 

L. beccarii, pp. 251 & 302. 

L. humerosus, pp. 251 & 302. 
L. anna, p. 303. 

L. inacassariensis, pp.256 & 305. 

' I have no specimens of L. novaeguineae , L. obtusidens, or L. glabey before me, and cannot be cer- 
tain that these characters are correctly noted here in their case. 

3-6 Memoirs of tlw Inditiu Museum. \\o\.. Ill, 

/- Pronotum unpuiictured, except in scars and marginal grooves L. timoricnsis, p. 303. 

Sides of pronotum more or less thickly covered with strong 

I, punctures . . . . . . . . . . 7. 

I f Central area of metasternum uupunctured .. .. I. (/f«/«/»s, pp. 252 & 303. 

\ At least a few strong punctures on central area of metasternuni L. cydntaenius , pp. 255 & 305. 
, Puncturing of abdominal sterna very variable; punctures, 
g I when present, of the usual type .. .. .. .. 9. 

1 Abdominal sterua covered all over with somewhat obscure, 

I broad, shallow punctures . . . . . . . . L. pluiius, pp. 260 & 310. 

/ Posterior intermediate areas of metasternutn punctured . . . . . . 10. 

t Posterior intermediate areas of metasternuni unpunctured . . L. roefyslorfi, pp 260 & 309. 

I Central part of mesosternum sharply delined on each side by a 
pair of ridges which, starting from the anterior angles, extend 
in a straight line backwards and a little outwards till close to 
iqI ' ^^^ posterior margin, where they curve strongly outwards; the 

I. whole of the mesosternum outside these lines depressed and 
granular ; only the central part smooth 
Mesosternal scars smaller, not occupying the whole of the si les 
of the mesosternum 
I I^ateral grooves of elytra moderately broad, punctures slightly 
ii\ transverse 

( Lateral grooves of elytra narrower, punctures rounder 
J 2 < Frontal ridges not reaching anterior margin of head 
( Frontal ridges reaching anterior margin of head . . 
( Posterior intermediate areas of metasternum finely punctured 

1 in inner angle ; elytra uniformly coloured . . . . L. oblnsidens, p. 307. 
I Posterior intermediate areas of metasternum strongly and 
V confusedly punctured ; elytra reddish in front and black behind L. glaber, p. 307. 

ipnncliifi, pp. 256 & 306. 

L. novaeguincac, p. 306. 

L. hkolor, pp 257 & 307. 

Genus Trichostigmus, Kaup. 

'About 24 mm. long; a few punctures in anterior angles of 
pronotum ; marginal groove of pronotum broad and deep and 
I coarsely punctured 

About 17 mm. long; surface of pronotum unpunctured except 
I close to scars ; marginal groove very fine or almost obsolete . . 

T. ilioreyi,Kz\x^; pp. 26i&3ro. 
T. iirsiiliis, Schaufuss; p. 310. 

Since writing the earlier part of this paper, I have received for examination a 
very remarkable Passalid from New Guinea. Although this insect is not from the 
Oriental Region, it may conveniently be described in the present paper ; for it will 
have to be made the type of a new genus, whose position in the classification of Indo- 
Australian genera outlined above is of peculiar interest. It appears, moreover, to 

' It is quite possible that this character is of less value than in this case it at present seems to 
be, and that these two species are not really distinct. 

This distinction is by no means a satisfactory one; but the position of L. novaegiiineae, as ex- 
plained above (p. 306), has not yet been properly determined. 

1914-] F. H. Gravely : An Account of the Oriental Passalidac. 327 

throw light 011 the position of that most aberrant of all Indo-Australian Passalids, 
Tarquinius paradoxus, Knwert, concerning which I have not hitherto dared to hazard 
an opinion. 

Pseudepisphenus, n. gen. 

Lamellae of ant<innac six in number, all moderately long. Mandibles as in Tar- 
quinius: symmetrical, the groove between the upper and middle terminal teeth very 
sharply defined on the lower side, branched just above the external angle of the 
mandible, the upper and more conspicuous branch extending obliquely across the 
externo-dorsal surface to end in the upper tooth which is obtuse and set very far back ; 
remaining teeth normal. Mentiim as in Episphenus: primary scars absent ; secondary 
scars small and confined to anterior margin. Left outer tubercle of head composed of 
two widely separated parts, of which only the outermost is represented on the right 
side. Lateral and intermediate areas of metasternum distinct. 

Pseudepisphenus perplexus, n. sp. 
Text-fig. 8, A & B. 
Described from a single specimen collected by Mr. A. F. R. WoUaston during the 
Utakwa River Expedition in Dutch New Guinea, and preserved in the British 

Text-figure 8. 

A. Psettdepisptienus perplexus, head x 4. 

B. Do. front of head from side x 8. 

C. Tarquinius paradoxus, head x 4. 

D. Do. front of head from side x 8. 

Length 24 5 mm. The lamellae of the antennae are moderately long and slender, 
the last three much longer than the first three. The labrum is punctured and hairy, 
its anterior margin is practically straight , its angles are rounded and the left one is 
distinctly more prominent than the right. The general form of the mandibles has 
been described in defining the genus ; it is unlike that found in any other Passalid 
known to me except Tarquinius paradoxus, Kuwert ; the external angle is, how- 
ever less pronounced than in that species, as is also the lateral keel behind it, 
which is only indicated by a fine groove just above the outer margin ; the anterior 
lower teeth, too, are somewhat shorter and stouter, and that on the left 
mandible is no larger than that on the right. The mentmn is extraordinarily like that 
of Episphenus indicus, and in the single specimen before me I can find no characters 
by which it can be distinguished. The head is highly polished, and entirely smooth 
except for a few punctures in front of the parietal ridges. Its general form is shown in 

3-2^ Memoirs of the Indian Museum. [Vol. Ill, 

text-figs. 8A& B. The right outer tubercle is very deep and broadly truncate as seen 
from the side. The outer portion of the left outer tubercle is exactly like it ; but be- 
tween this and the middle-line is a large rounded process which does not occur on the 
right side. The surface of the head between the outer tubercles and the supraorbital 
ridges is more or less level, not excavate. The apical angles of the supraorbital 
ridges are very distinct, of about 120°, and slightly peaked. The pronotum is un- 
punctured except in the marginal groove and in the small, round, hairless .scars. The 
marginal groove is broadly incomplete before and behind; but the median groove, 
which is deeply impressed, is practically complete. The plates of the loioer side of the 
prothorax bear punctures and hair distributed as usual in the family, but not so 
thickly or extensively as sometimes. The scutellum is very indistinctly punctured all 
over ; the mesothoracic epistcrna are coarsely punctured above and along the anterior 
margin ; the mesosterntim is highly polished throughout, except in the deeply im- 
pressed semi-lunar scars, and even these are scarcely dull. The lateral areas of the 
metasternum are narrow, parallel-sided and slightly roughened ; they are very sharply 
-separated from the anterior and posterior intermediate areas, which are broadly 
continuous with one another, and are coarsely punctured except along their outer 
margins. The posterior coxae are smooth. The scars of the abdominal sterna are 
more or less roughened and punctured. The elytra are hairless, and are unpunctured 
except in the grooves, of which the lateral are much more coarsely punctured than 
the dorsal, their punctures being, however, scarcely transverse. 

The systematic position of this form is somewhat difficult to determine. The 
structure of the anterior margin of the head at once suggests relationship with the sub- 
family Gnaphalocneminae ; and the Aceraiine form of mentum seems to place it near 
the genus Hyperplesthcnus. In Hyperplesthenus and all other genera of the group to 
which it belongs, however, the lateral and intermediate areas of the metasternum are 
fused, whereas in Pseudepisphenus no trace of any such fusion is found; and all 
known species of the Hyperplesthenus group are much larger insects than Pseudepis- 
phenus perplex us. 

In size and general appearance, Pseudepisphenus resembles rather the Protomo- 
coelus group, the simpler members of which have, like it, a metasternum of the ordinary 
type. The possibiUty of the absence of primary scars from the mentum in this group 
has been pointed out above (p. 194, footnote) ; and it is quite likely, I think, that in 
Pseudepisphenus we have a case in point. 

Although the precise systematic position of Pseudepisphenus is open to this much 
doubt, its asymmetry and consequent obvious connection with some group of the 
Gnaphalocneminae are of great interest on account of its apparent affinity, on the other 
hand, with the aberrant genus Tarquinius. The peculiar structure of its mandibles 
is essentially the same as that found, so far as I know, in Tarquinius alone of all 
Indo-Australian Passalidae. The chief difference between the mandibles of the two 
genera lies in the fact that the peculiar external keel is more strongly emphasized in 

igi4.] F. H. Gravely: An Account of the Oriental Passalidae. 329 

Tarquinius than in Pseudepisphenus, i.e. that the pecnharities which have appeared 
in Pseudepisphenus tend to be accentuated in Tarquinius. Quite in keeping with 
this is the fact that the scars on the mentum, of which the primary pair has 
disappeared and the secondary is small in Pseudepisphenus, have completely dis- 
appeared in Tarquinius. And the outer tubercles of Tarquinius very closely re- 
semble the right outer tubercle of Pseudepisphenus and that part of the left which 
is symmetrical with it — all that is required to produce a head shaped like that of 
Tarquinius from that of Pseudepisphenus, is a shift forwards of the inner tubercles 
to the anterior margin, where that of the left side would replace the asym- 
metrical inner portion of the left outer tubercle. The outer tubercles in the Leptaula- 
cinae, on the other hand, are simple and more or less acute; and the lyeptaulacinae 
are the only known Indo-Australian forms to which Tarquinius bears even a 
superficial resemblance. It is further separated from them by the presence of six 
well-developed lamellae on each antenna instead of only three, by the structure of its 
mandibles, "and by the absence of scars from the mentum. 

The differences between the Aceraiinae and Gnaphalocneminae seemed very great, 
so long as my knowledge of the latter subfamily was practically confined to the 
information I could glean from previous authors. The Aceraiinae are, undoubtedly, 
somewhat isolated from a geographical point of view ; and the separation has been a 
convenient one for the purposes of the present paper. That their recognition as a 
distinct subfamily on anatomical grounds is less easy to justify than it at first 
appeared to be, and that it may have to be abandoned, has already been suggested 
in Appendix I. If Tarquinius is also to be included in the Gnaphalocneminae, it is 
difficult to see why the Macrolininae and perhaps also the Pleurariinae and even the should not be included as well. Whether these several series are to rank 
as subfamihesor as groups, is purely a question of convenience; and it cannot be 
satisfactorily settled by one who has no special knowledge of African and American 

Pseudepisphenus and Tarquinius appear to have originated from an asym- 
metrical ancestor within some group of the Gnaphalocneminae as defined at the begin- 
ning of this paper. There is no reason to suppose that they have had a separate 
ancestry from symmetrical forms, as is the case with the Aceraiinae. It is, however, 
impossible to include them in the Gnaphalocneminae without redefining that sub- 
family. This I could not do without either splitting it up into groups, or else enlarg- 
ing it to some extent and thus raising the larger issues referred to in the preceding 
paragraph ; and these I am not in a position to deal with. For the present, therefore, 
it seems best to regard the genera Pseudepisphenus and Tarquinius as constituting a 
distinct subfamily Tarquiniinae, distinguished from all others by the strong groove 
which extends from the upper tooth to the outer angle of both mandibles. 

The genera Pseudepisphenus and Tarquinius may be separated thus: — 
Inner tubercles situated behind anterior margin of head ; outer 

tubercles asymmetrical .. .. .. .. Pseudepisphenus, Gravely. 

Inner tubercles situated on anterior margin of head as in Lep- 

iaulax; outer tubercles symmetrical .. .. •• Tarquinius, K.\i\\ext. 

330 Memoirs of the Indian Museum. [Vol. Ill, 

The evolution of a symmetrical Lcptaulax-like form from an asymmetrical ancestor, 
suggests that the union of the inner tubercles with the anterior margin of the head 
represents an even higher degree of specialization than does the asymmetrical condition . 
The extraordinarily wide distribution and exceptional variability of the two dominant 
species of Leptaitlax tend to support this suggestion. From which it may be inferred 
that the Leptaulacinae are of comparatively recent origin, and that their scarcity in the 
Indian Peninsula and Ceylon is due to the fact that they \i'\vi not yet had time fully 
to establish themselves there. 

Whether the high degree of specialization thus indicated in Tarqitiimis will prove 
sufficiently advantageous to its po.ssessor to enable it to become a dominant form 
remains to be seen. At present the only known species of the genus appears to be 
extremely rare. 

Before finally leaving the question of the many different forms assumed by 
different species of Gnaphalocneminae and their allies, a further aspect may be 
emphasized of the fact that in Pscmiepisphenus perplexus we have a specie§, obviously 
of Australian and not Oriental extraction, whose mentum is indistinguishable from 
that of an Oriental species. Its Australian ancestrj^ is indicated, apart from zoogeo- 
graphical considerations, only by the form of the anterior margin of the head, 
and this is the only structural indication of such ancestry that we could expect to 
find in a species with a mentum of this form . 

If one species of Australian ancestry has such a mentum , there is no reason why 
another should not have it also ; and the alteration needed in the form of the anterior 
margin of the head of many Gnaphalocneminae to make them resemble the Aceraiinae 
in this respect also, is no greater, and would be no more remarkable, than the 
alteration that appears to have taken place in the evolution of Tarqiiiiiiiis. Two so- 
called species, "Laches" infantilis and piurilis, Kaup, which I have been unable 
to distinguish from Episphenus neelgherriensis, are recorded from Vanicoro (Santa 
Cruz Islands) and the Aru Islands respectively. In the absence of any confirmation 
of these records since the " species " were first described in 1871, the probability is, as 
pointed out above (p. 284), that they are incorrect. Should either or both of them be 
confirmed, however, convergence would, I consider, offer a much more plausible 
explanation of such anomalous distribution, than migration in a manner utterly at 
variance with that otherwise adopted throughout the Aceraiinae. It is, moreover, by 
no means improbable that species may yet be found in Australian islands which, 
though really allied to some group of Gnaphalocneminae, and distinct from any 
known species of Aceraiinae, have, nevertheless, the characters of the latter rather 
than of the former subfamily. 

Largely as a result of work done in European museums while the earlier parts of 
this paper were going through the press the following additions have recently been 
made to the Indian Museum collection. 

iqi4.j F. H. Gravely : An Account of the Oriental Passalidae. 331 


Genus COMACUPES, Kaup. 

Comacupes cylindraceus (Perty). 

Regd. No. ^^*-f," Telom, S. Perak, 4000 ft. Oxford Museum. 

Comacupes foveicollis, Kuwert. 

Regd. No. '-^ Baram R., Sarawak British Museum. 

Genus TRISTORTHUS. Kuwert 
Tristorthus tricuspis (Kaup). 

Regd. No. 

VahouL", New Caledonia K. Zool. Mus., Berlin. 

Taeniocerus platypus, Kaup. 

Regd. No. *^i;-* Deli, N.E. Sumatra K. Zool. Mus., Berlin. 

Aulacocyclus deyrollei, Kaup. 

Regd. No. *^ Ballarat Djst., Victoria Deutsches Entom. Mus. 

Aulacocyclus sp. 

Regd. No **-f Australia ? 

Genus CERACUPES, Kaup. 
Ceracupes arrowi, Heller. 

Regd. No. ^-If Kosempo, Formosa Deutsches Entom. Mus, 

Genus PURITULUS, Zang. 
Auritulus patalis, Lewis. 

Regd. No. -^ (cotypes) Yuyama, Japan British Museum. 

Subfamily ACERAIINAE. 


Ophrygonius cantori subsp. convexifrons, Zang. 

Regd. No. '~^ Sin Lum, Bhamo, hooo ft. British Museum. 

7293-4 „, 

,, -^TTT — Bhanu) ,, 

332 Mi'iiioii's 0/ the Iiiduui Mtisiitni. [Vol. Ill, 

Ophrygonius inaequalis (Burmeister). 

Regd. No. ^ Singapore British Museum. 

*ir I'enang 

Aceraius perakensis, Kuwert. 

Regd. No. -5jf Taiping, 4000-4500 It. K. Zool. Mus.", Berlin. 

"1^ Telom, S. Ferak, 4000 ft. Oxford Museum. 

Regd. No. 

Aceraius laevicollis (lUiger) 

I W. Sumatra (" Bandar Buat b. I 


\ PadaugSchoedeS. G.") 

! IS.. />ooi. .MUS., joeriHi. 


Delhi, Sumatra 




Deutsches Eutoui. Mus. 



K. Zool. Mus., Berlin. 


S. Palawan 

Deutsihes Entom. Mus 



K. Zool Mus., Berlin. 

Aceraius grandis (Burmeister), s. str. 

Regd. No. ~ Sumatra K. Zool. Mus., Berlin. 

,^ ^-i*i Tengger Mountain, Java Deutsches Entom. Mus. 

«»« m I i K. Zool. Mus., Berlin. 

•• . 20 20 ' - ' ' 


,, !^' ) ( Deutsches Entom. Mus. 

1?J Bandjermasin K. Zool. Mus., Berlin. 

A. grandis var. rectidens, Kuwert. 

Regd. No. 'i!,'' Deli, Sumatra- K. Zool. Mus., Berlin. 

"20 J a a I , 

^^ {A. Magnus, ? Deutsches Entom. Mus. 
Kuwert det.) 

A. grandis subsp. hirsutus, Kuwert. 

Regd. No. "^" Catchin Canri Deutsches Entom. .Mus. 

.Mt. Mauson, 2-3000 ft. Tonkin K. Zool. Mus., Berlin. 

Tonkin Deutsches Entom. Mus. 

Hainan K. Zool. Mus., Berlin. 
Fuhosho, S. Formosa 

Kosempo, Formosa Deutsches Entom. Mus. 

Polisha, Formosa K. Zool. Mus., Berlin. 

Sokutsu, Formosa Deutsches Entom. Mus. 

IOI4.] F- H. Gravely : An Acconiit of the Oriental Passalidae. 333 

Aceraius kuwerti, Zang. 

Regd. No. ^' Mt. Kiua-Balu, c. 5000 ft. Deutsches Entom. Mus. 


Regd. No. 

Mt. Kiiia-Balu, 45oo ft. Sarawak Museum. 

Aceraius laevimargo, Zang. 

Mt. Kina-Balu, c. 5000 ft. Deutsches Entom. Mus. 

Aceraius pilifer (Percheron). 

Regd. No. '.tr Preanger, Java, 4000-6000 ft. Hamburg Museum. 

Miy Tiibodas, c. 5000 ft. K. Zool. Mus., Berlin. 

«7ii3 Tjibodas • Hamburg Museum. 

5598 Borneo H. E. Andrewes. 

" 20 

Sarawak Museum. 

Regd. No. 

Mt. Kina-Balu, 4500 ft. 

Aceraius helferi, Kuwert. 

Sin Lum. Bhamo, 6000 ft. British Museum. 


Aceraius borneanus, Kaup. 

Regd No. i^' ^^^- Kina-Balu, c. 6000 ft. Deutsches Entom. Mus. 

«L7 S. Palawan 

Aceraius minor, Gravely. 

Regd. No. '-^(cotype) Taiping, 4000-5000 ft. K. Zool. Mus., Berlin. 

'Ml±:8 Telom S. Perak, 4000 ft. Oxford Museum 

>> 20 ' 


Genus MACROLINUS, Kaup. 

Macrolinus crenatipennis, Kuwert. 

• , ,, is-io , J J, ■l^ r^,.i,>n K. Zool. Mus., Berlin. 

Regd. No. *^^ (cotype') Ce>lon 

Macrolinus sulciperfectus, Kuwert. 

Regd. No. 'f (cotypc) Bonthain, Celebes K. Zool. Mus., Berlin. 

Episphenoides quaestionis (Kuwert). 

Regd. No. if Sidney Deutsches Entom. Mus 

I Larva, pupa and adult in spirit. 

■^ Kuwert says that he only saw one specimen; but he gives the lengths of two. 

334 Mi-»i('irs of the Iiidimi MiiSiiim. [Vol. Ill, 

Mastochilus polyphyllus (MacLeay). 

Regd. No. ',]f; Australia K. Zool. Mus., Berlin. 

Pharochilus sp. 

Regd. No. f Australia ? 

(ienus KAUPIOLOIDES, (.ravely. 
Kaupioloides trigonophorus (Zang). 

Regd. No. y-]' Toricelli -Mts., Kaiser Wilheloislaiid Deutsches Kntom. .Mus. 

Genus CETEJUS, Kaup. 
Cetejus grabowskyi, Kuwert. 

Regd. No. ''^''•' Stephaasort, Kaiser Wilhelmslaiid K. Zool. Mus., IJjrliu. 

Protomocoelus heynei (Kuwert). 

Regd. No. ^^^{' Kaiser Wilhelmsland, New (iuiiiea K. Zool. Mus., Berlin 

Genus KAUPIOLUS. Zang. 
Kaupiolus moluccanus (Percheron). 
Regd. No. *^ Moluccas Deutsches Hutoin. .Mus. 

Kaupiolus compergus (Boisduval). 

Regd. No. S51 Kapaur, Dutch New Guinea K. Zool. Mus., Berlin. 

Genus LABIENUS, Kaup. 
Labienus, sp. 

Regd. No. ^■' ? K. Zool Mus., Berlin. 


Bukana, Huon Gulf, German New 

Hyperplesthenus gracilis (Heller). 

Regd. No. "^ (tolype) Toricelli Mts. Kaiser Wilhelmsland Deutsches Entom. Mus. 

Parapelopides symmetricus, Zang. 

Regd. No. ■y^" Mt. Kina-Balu, c. 5000 ft. Deutsches Entom. Mus. 

iqi4-] P- H. Gravely: Au Account of the Oriental Passalidae. 335 

Trapezochilus dorsalis (Kaup). 

Regd No. ^^ Taiping, 4500-5000 ft. K. Zool. Mus., Berlin. 

"^ Bukit Besar, 2500 ft. ^ 

6218-24 T, , J., > Oxford Museum. 

—2;; — Telom, 4000 ft. J 

*~ Deli, Sumatra Deutsches Entom. Mus. 

Gnaphalocnemis burmeisteri (Kaup). 

Regd. No. ^ Padang, \\'. Sumatra Deutsches Entom. Mus. 

~ Deli, Sumatra 

~ Teinau, Sumatra K. Zool. Mus., Berlin. 

^' Kepahiang 

Gnaphalocnemis monticwlosus (Smith). 

Regd. Nq. —' Deli, Sumatra K. Zool. Mus., Berlin. 

^ Sumatra 

*|f Sarawak 

Gnaphalocnemis tridens (Wiedemann). 

Regd. No. ~ Java Deutsches Entom. Mus. 

^-^ Asia 

,, 21, Asia • ,, 

,, **.^"' Tjibodas, Java K. Zool. Mus., Berlin. 

Genus OMEGARIUS, Kuwert. 
Omegarius minimus, Kuwert. 

Regd. No. ^ Samberi, Dutch New Guinea K. Zool. Mus., Berlin. 

,, ^ Kaiser Wilhelmsland, New Guinea ,, 

,, ^—^ N. Bucht Squally I., New Britain Hamburg Museum. 

Gentis TARQUINIUS Kuwert. 
Tarquinius parodoxus, Kuwert. 

Regd. No. i^ Wandesi, New Guinea Deutsches Entom. Mus. 


Genus LEPTAULAX, Kaup. 

Leptaulax humerosus, Kuwert. 

Regd. No. ~ W. Sumatra Deutsches Entom. Mus. 

^ Ardjoeno, Java .. 

>) ^ Tengger Mountain, Java ,. 


Mcmoira of the huiiau Museum. 
Leptaulax anna, Zang. 

[Vol. Ill, 

Regd. No. *^^^ {co/ype) Suinhawa 

Hamburg Museum. 

Regd. No, 


No. "^' 


Regd. No. 

Regd. No. 

4460 m\i 44*!4 

Leptaulax timoriensis (Perchcron). 

'I'iinor K. Zool. Mus., Berlin. 

L. dentatus (Fabricius) s. str. 

Mt. Mauson, 20oo-,',0oo ft. K. Zool. Mus , Berlin. 

Fuhosho, Formosa Deutsclies Kntom. Mus. 

Poiisha, Formosa 
Sokutsu, Formosa 

L. dentatus, \^ar. glabriventris, Gravety. 

'^y Deli, .Sumatra K. Zool. Mus., Berlin. 

Leptaulax cyclotaenius subsp. himalayac, Kuwert. 

"i" Mt. Mauson, 2000-3000 ft. K. Zool. Mus., Berlin. 

Leptaulax bicolor Fabricius) s. .str. 

VtlS^i 4.(20 

Regd. No. -^ ,„ 


*t«7v S 



+419 4421 
20 20 

Mt. Mauson, 2000-3000 ft. 
Kosempo, Formosa 
Poiisha, Formosa 
Taiping, 4000-5000 ft 
St. Rambe, Sumatra 
Preanger, Java, 4000-6000 ft. 
Tjibodas, c. 5000 ft. 
Tjibodas, Java 
Banguej' Island 

K. Zool. Mus., Berlin. 
Deutsches Entom. Mus. 

K. Zool. Mus., Berlin. 
Deutsches Entom. Mus. 
Hamburg Museum. 
K. Zool. Mus., Berlin. 
Hamburg Museum. 

Regd. No. 

Leptaulax bicolor var. vicinus (Percheron). 

.Mt. Kiiia-Balu, 4500 ft. Sarawak \[useum. 

I . Classification . 
The Indo-Australian Passalidae have been divided into two main sections (pp. 
191-192). The first section contains the Aulacocychnae of Kuwert, together with 
Cylindrocaulus and Auritulus It has been regarded as a single subfamily, but the 
two last-named genera and Ccracupcs maj^ ultimately have to be regarded as belonging 
to a different subfamily from the rest (p. 192). It contains eight genera (pp. 192-193). 
The second section has been subdivided into the Pleurarius, Aceraius, Macrolinus, 


F. H. Gravely : An Account of the Oriental Passalidae. 337 

Kaupioloides , Protomocoelus, Hyperplesthenus, Gnaplialocnemis, Plesthcnus, Gonatas, 
Tarquinius and Leptaulax groups. Of these groups the first three and the last two appear 
tobeof most importance, and have been provisionally ranked as subfamilies, the remain- 
der 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 (pp. 193-4, 199 and 329). The genera and Oriental species of all 
of them have been redefined (pp. 197, 199-204, 317-8, 329 and 318-326). 

2. External Morphology and Taxonomy. 

The pair of tubercles situated in the Pleurariinae, Aceraiinae, Macrolininae and 
Gnaphalocneminae, between the central tubercle of the head and the processes of the 
anterior margin, have been shown to be homologous with the inner pair of marginal pro- 
cesses found in the L,eptaulacinae ; and the terms inner and outer tubercles have been 
consequently applied to the two pairs of processes found in all members of the second 
section of the family (pp. 184-185). The use of these terms does not commit one to 
any definite system of interpretation of the homologies of the head ; which is advan- 
tageous, inasmuch as there is reason to believe 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 ; 
in which case these tubercles are not really processes of the clypeus as they have hitherto 
been called (p. 185). 

Apart from this no criticism of the accepted homologies of different parts of 
the body has been found necessary"; but considerable changes have been made in 
the taxonomic values assigned to different parts (pp. 179-191), and it has been found 
necessary to use aU characters with much greater caution than has often been the case 
in the past. As a result of this it has been found possible to define species with 
a considerable degree of precision. Five species, however, stand out from all the rest 
by reason of their remarkable variability, in size especially ; and certain structural 
variations are corellated with variation in size, much as in the I^ucanidae (pp. 262- 
265). These species are Episphenus comptoni, Episphenus indicus, and Aceraius 
grandis which appear respectively to be the dominant species of Aceraiinae in each 
of the three divisions of the Oriental Region in which they occur ; and Leptaulax 
dentatus and Leptaulax bicolor which are the dominant species of I,eptaulacinae 
throughout the whole of this region (pp. 311-313). 

In the Aceraiinae and Gnaphalocneminae, the two subfamilies in which markedly 
asymmetrical forms are found, the degree of asymmetry attained by a species appears 
to be an index of the degree of specialization attained : and the dominant species of 
a region is always the most markedly asymmetrical species found there (p. 312 '). 

The nature of the asymmetry is somewhat different in different groups, and the 
asymmetrical condition appears to have been evolved more or less independently in 
each of them (fig. 7, p. 314). The evolution of a symmetrical Leptaulax-like form, 

As regards the Gnaphalocneminae see Gravely, 1914 (b). 

338 Memoirs of the Indian Mtiscitm. [Vol. Ill, 

Tarqiiinius, from an asymmetrical ancestor allied to the (jnaplialocneminae, suggests 
that the union of the inner tubercles with the anterior margin of the head indicates 
an even higher degree of specialization than does the asymmetrical condition (p. 330). 

3. Geographical distribution. 

The material examined during the preparation of this paper does not enable me 
to discuss the relation borne by Indo-Australian Passalidae to members of the family 
found in Africa or America; but it seems by no means improbable, from Kuwert's 
account of the family, that all the seven subfamilies described, with the possible ex- 
ception of the Leptaulacinae, will prove to be confined to the Indo-Australian Region, 
China and Japan (p. 194). 

With the exception of the Pleurariinae, Aceraiinae and Tarquiniinae all these 
subfamilies occur on both sides of the Straits of Macassar, but the AulacocycHnae and 
Gnaphalocneminae are much more strongly represented east of these straits, and in the 
eastern parts of the Oriental Region, than they are further west; while with the 
Macrohninae the reverse is the case (pp. 311 & 313). 

The discontinuous distribution of the genus Aulacocyclits suggests that the 
Aulacocyclinae were once more abundant towards the west than they are now. 
Although the Macrohninae are well represented in Ceylon, they appear to be absent 
from the Indian Peninsula. The Pleurarariinae occur only in the Indian Peninsula 
and Sumatra, which discontinuous distribution suggests that they are probably the 
remnants of a once more extensive group. The Leptaulacinae are centred in the East 
Indian Archipelago. Only the dominant species seem to have established themselves 
in India and Ceylon, and these are less common there than elsewhere (p. 311). 

The distribution of the Aceraiinae and Gnaphalocneminae, the two subfamilies in 
which asymmetrical species are found, is of much greater general interest. It appears 
to have been greatly influenced by the following zoogeographical boundaries: — 
Palk Strait, the Gangetic Plain (in which Passalids seem unable to live — pp. 310-1) 
the Straits of Macassar, and Torres Strait. Of these the Straits of Macassar are by 
far the most important, for no group of either subfamily is found in any abundance 
on more than one side of them. It is therefore somewhat surprising to find that this 
is the only one of the above-mentioned boundaries, which any species of these sub- 
families has succeeded in crossing (pp. 313 & 315-6). Another boundary of some 
importance is that separating the East Indian Archipelago — including the Malay 
Peninsula and presumably also the Philippines — from continental Asia (p. 313). 

The occurrence of different degrees of asymmetry in different species of Aceraiinae 
probably affords an index to the degrees of speciahzation to which they have severally 
attained, and its correlation with locality is very evident. It is concluded that 
a migration has taken place westwards towards the head of the Bay of Bengal and 
then south-westwards towards Ceylon, the less specialized forms being constantly 
displaced by their more specialized allies (pp. 311-314). It may further be pointed 
out that although neither of the two most highly asymmetrical species of the genus 
Episphenus have been able to cross over from India to Ceylon, and the genus Aceraius 

1914.] F. H. Gravely : An Account of the Oriental Passalidae. 339 

has been unable to cross the Gangetic Plain , the latter genus has had no difficulty in 
occupying the Sunda and Philippine Islands, as well as a large part of the Asiatic 

The evolution and migration of different forms of Gnaphalocneminae seems to 
have followed lines similar to those followed by the Aceraiinae, the most primitive 
forms again being driven out from the Archipelago — in this case eastwards to Australia 
instead of westwards towards Ceylon. As a result of this, the most closely related 
(symmetrical) forms of the two subfamilies are found only in two widely separated 
countries on the periphery of Indo- Australian area, the intervening countries being 
inhabited by more highly specialized forms most of which are asymmetrical (pp. 

4. The Dominant Species. 

Five Oriental species stand out from all others on account of their extraordinary 
variability in size. Three of these belong to the Aceraiinae, and two to the Leptau- 
lacinae (p. 262). 

Of the first three, one {Episphenus coniptoni) is confined to Ceylon, one {Epis- 
phemis indicus) to the Indian Peninsula, and one {Aceraius grandis) to the rest of the 
Oriental Region. Each of them differs from the species of Aceraiinae with which they 
are geographically associated in that they are more abundant, more highly asymmetri- 
cal, and always markedly gregarious ; but these distinctions are much less pronounced 
in Episphenus indicus, than in the other two. Aceraius grandis, in addition to these 
distinctions, is by far the most "vvridely distributed member of its genus. It has 
further been pointed out, that of the three genera of Aceraiinae inhabiting the country 
east of the Ganges, Aceraius, which is the most markedly asymmetrical, is also the 
most numerous both in species and in individuals; Ophrygonius, which differs from 
Aceraius principally in having symmetrical mandibles, comes next, and is perhaps 
equally widely distributed ; Tiber io ides, in which the head is symmetrical as well as 
the mandibles, being the smallest and scarcest genus of the three (pp. 311-313). 

No asymmetrical species of Leptaulacinae are known ; but there is reason to 
suppose that the structure of the head found throughout this subfamily indicates 
an even higher degree of specialization than does asymmetry (p. 330). It is 
therefore not surprising to find that the two variable species, both of which are 
gregarious, and much more abundant than any other species of the subfamily, are 
even more widely distributed than any of the dominant species of Aceraiinae, both of 
them being found in all subregions of the Indo-Australian area in which Passalids of 
any kind occur. 

5. Habits. 
Such information as I have been able to gather together concerning the habits of 
Oriental Passalidae is recorded above under the descriptions of the several species 
(Taeniocerus bicuspis, p. 211 ; Pleurarius brachyphyllus, p. 215; Tiberioides austeni, 
pp. 216-7 ; Episphenus comptoni, pp. 219-220 ; Episphenus indicus, p. 222 ; Episphenus 
neelgherriensis , p. 223; Aceraius grandis, sub-sp. hirsutus, p. 233; Aceraius helferi, 

340 Mi'Hiuiys of the Indian Museum. [Yoi.. Ill, 

p. 238; Macroliniis sikkimcnsis, p. 243; Macro/inns rotnndi/rons, p. 245; Leptaulax 
dentatus, p. 255; Leptaulax bicolor, pp. 258-9; and Leptaulax rocpstorfi, p. 260). This 
information, as far as it goes, confirms Arrow's belief that the habits of Oriental 
forms would be found to resemble those of the American forms described by Ohaus, 
on which Arrow found himself compelled to base the account of the habits of the 
family in the introduction to his account of the Indian lyamellicornia in the " Fauna 
of British India" series. 

It has been found that different species differ somewhat in their habits one from 
another in various ways. Probably adults of nearly all species live in decaying wood 
in pairs with their young ; but whereas in certain species large numbers of such 
families are commonly found in a single log, so closely associated with one another 
that it is often impossible to separate out the individuals belonging to any single 
pair, in others each family forms an isolated group. All the five dominant species 
are gregarious. 

Leptaulax roepstorfi, one of the smallest and most markedly flattened of the 
Oriental representatives of the family, inhabits natural cracks in hard logs, into the 
depths of which it penetrates. Most species burrow nearer the surface, and the 
Leptaulacinae, most of which are much flattened, appear to burrow as a rule even 
closer to the bark than do the Aceraiinae, which are less flat. One at least (Leptaul.ix 
dentatus) of the two dominant species of Leptaulacinae is, moreover, only found in very 
rotten wood ; whereas the dominant representative of the Aceraiinae in Assam and 
the adjoining country, Accraius grandis sub-.sp. hirsutus, as well as the only other 
member of the genus whose habits have been studied {A . helferi) are only found in wood 
of a very much tougher consistency. On the other hand, the Aceraiinae inhabiting 
the Indian Peninsula [Episphcnus indicus and neelgherriensis) appear to live in quite 
rotten wood, which may perhaps help to account in some measure for the apparent 
scarcity of Leptaulacinae in that area, where probably at most not more than half 
a dozen separate colonies have yet been noticed. Episphenus neelgherriensis has been 
found under stones as well as in rotten wood ; and Macrolinus rotundifrons has been 
found among decaying vegetable refuse. 

Reproduction appears to be retarded or arrested during dry cold weather 
(p. 22Z). 


Papers marked with ati asterisk (*) are not available in Calcutta. 

1792. l-abricius, J. C. " Entomologia Systematica emenda et aucta." Yo\. I 
(Hafniae, 1892), pp. 240-1. 

* 1800. Illiger, K. ' Vierzig neue Insecten aus der Hellwigischen Sammlvmg in 

Braimschweig " Wiedemann Archiv fiir Zool. u. Zoot. (Berlin and 
Brunswick 1800) I (2), pp. 103-150, and II, pp. 229-230. 

* 1800. Erichson in Wiedemann Archiv fiir Zool u. Zoot. I (1800). 

1914-] F- H. Gravely: An Account of the Oriental Passalidae. 341 

1801. Fabricius, J.C. " Systema Eleutheratorum II " (Kiliae,i8oi), pp. 255-6. 
1801. Weber, F. " Observationes Entomologica " (Kiliae, 1801). 

* 1806-17. Schonherr,C. J "Synonymia Insectoruni,oder : Versuch einer Syno- 

nymie aller bisher bekanten Insecten; nach Frabricii Systema 
Eleutheratorum geordnet " (Stockholm, 1806-17). 
1823. Wiedemann, C. R. " Zweihundert neue Kafer von Java, Bengalen, und 

dem Vorgebirge der Gutten-Hoffnting." Zool. Mag. II (i), p. log. 
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Intertropical and Western Coasts of Australia performed between the 
years 1818 and 1822." Vol. II (London, 1826). 

* 1826. Stunn, J. "Catalog meiner Insecten-Sammlung," I (Niirnberg, 1826). 

* 1831. Perty. "Obs. Nonnulae Coleopt. Indiae Orientalis." 

1835. Boisduval, J. B. A. D. de. "Faune Entomologique de I'Ocean Paci- 
fique. II. Coleoptere et autres Ordres" in "Voyage de 1' Astrolobe 
... .pendent. .. .1826-9 sous le commandent de M. J. Duraont 
d'Urville etc." 

1835. Percheron, A. "Monographic des Passales (Paris, 1835). 

* 1837. Dejean, P. F. M. A. "Catalogue de la Collection de Coleopteres de M. 

le Baron Dejean," 3 ed. (Paris, 1837), pp. xiv, 503. 

1 841. Percheron, A. " Revision critique et Supplement a la Monographic du 

Genre Passale, premiere partie." Mag. Zool., Insectes, 1841. PI. 

1842. Westwood, J. O. " Insectorum no varum Centuria." Ann. Mag. Nat. 

Hist. VIII, 1842, pp. 123-125. 

1843. Guerin Meneville, F. E. " Animaux Articules " in "Souvenirs d'un 

Voyage dans I'lnde execute de 1834 a 1839 par M. Adolphe Deles- 
sert," Pt. II (Paris 1843), pp. 33-98. 

1844. Percheron, A. " Monographic des Passales, Second Supplement." Mag. 

Zool., 1844, pi. cxxxiv-cxxxv. 

* 1845. Hope, F. W. A. (Westwood) " A Catalogue of the Lucanoid Coleop- 

tera in the collection of. . . .F. W. Hope, etc." (London, 1845), p. 31. 
1847. Burmeister, H. " Handbuch der Entomologie," V (Berlin, 1847), pp. 

1850. Castelnau, F. L. de E. de. " Histoire Naturelle des Animaux Arti- 
cules." Insectes Coleopteres par M. le Compte de Castelnau, II 

(Paris 1850), pp. 178-9. 
1852. Smith, F. "Nomenclature of Coleopterous Insects in the Collection of 

the British Museum. Pt. VI Passalidae" (London, 1852). 
1855. Montrouzier, le Pere. " Essai sur la Faune de I'lle de Woodlark ou 

Moiou. " Ann. Sci. Phys. et Nat. d'Agric. et d'Industrie (Soc. Imp. 

d'Agric. etc. de Lyon, 1855) VII (i), pp. 1-114. 


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i860. Montrouzier, le Pere, " Essai sur la Faune Entoniologique de la Nouvelle- 
Caledonie (Balade) et des lies des Pins, Art, Lifu etc." Ann. Soc. 
Ent. France (3) viii, pp. 229-308. 

1862. Fauvel, A. " Coleopteres de la Nouvelle-Caledonie recueilles par M. E. 
Deplanche 1858-60." Normandie, Soc. Linn. Bull. VII, pp. 120-185. 

1867. Redtenbacher, L. " Coleoptera " in " Reise der Osterreichcn Fregatte 

' Novara '; Zool. II " (Vienna, 1S67), 249 pp., 5 pi. 

1868. Gemniinger and Harold. " Catalogus Coleopterorum " II (Munich, 

1868), pp. 968-972. 

1868. Kaup, J. ((0 " Prodromus zu einer Monographic der Passaliden." 

Coleopterologische Hefte, III, 1868, pp. 4-32. 
{h) Ditto. Coleopterologische Hefte, IV, 1868, pp. 1-7. 

1869. Kaup, J. Ditto. Coleopterologische Hefte, V, pp. 38-40. 

1871. Kaup, J. "Monographic der Passaliden." Berlin Ent. Zeitschr. 15 

Jahrg. Supplement, 125 pp., pi. iii-vii. 
1871 MacLeay, W. " Notes on a Collection of Insectes from Cxayndah (Cole- 
optera) pt. III." Trans. Ent. Soc, N. S. WalesJI, i87i,pp. 159-205- 
1873. Stoliczka, F. "A contribution towards a Monograph of the Indian 

Passalidae." J.A.S.B. XUI (H), 1873, pp. 149-162. 
1877. Kirsch, T. {a) " Neue Kafer aus Malacca." Mitt. K. Zool. Mus. Dres- 
den, I, 1877, pp. 27-58. 
(6) " Beitrage zur Kenntniss der Coleopteren-Fauna von Neu 
Guinea." Mitt. K. Zool .Mus. Dresden, II, 1877, 
pp. 135-161. 
1880. P'airmaire, L-, in Le Naturaliste, i88u, p. 1.64. 

1883. Lewis, G. " Lucanidae of Japan." Trans. Ent. Soc. London, 1883, 

PP- 333-342, pi. xiv. 

1884. Wystmann, P. " Catalogue Sj^stematique des Passalides." Ami. Civ 

Mus. Genova, (2a) I (XXI) 1884, pp. 326-348- 

1885. Schaufuss, L. W. " Beitrage zur Fauna der Niederlandi.schen Besit- 

zungen auf den Sunda-Inseln." Horae Soc. Ent. Ross. XIX 1885, 

pp. i83-'209. 
1887. Fairmaire L. " Coleopteres de I'lnterieure de la Chine." Ann. Soc. 

Ent. Belg. XXXI, 1887, pp. 87-136. 
'" 1890. Kuwert, A. " Die Passaliden und ihr Monographic von Kaup." Soc. 

Ent. V, 1890, pp. I, 9, and 17. 
1891. Kuwert, A. " Systematische Uebersicht der Passaliden Arten und 

Gattungen." Deutsche Ent. Zeitschr. 1891 (i), pp. 161-192. 
1896. Blackburn, T. "New Genera and species of Australian Coleoptera, 

XX." Trans R. vSoc. S. Australia XX, pp. 233-259. 
1896. 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., i pi. 

T9I4-] F. H. Gravely : An Account of the Oriental Passalidac. 343 

1896. Kuweit, A. "Die Passaliden dichotomisch bearbeitet" Novit. Zool. 

Ill, 1896, pp. 209-230, pi. v-vii. 

1897. Kuwcrt, A. Ditto. IV, 1897, pp. 274-306. 

1898. Kuwert, A. Ditto. V, 1898, pp. 137-205 and 259-349. 

i8g8. Heller, K. M. "Neue Kafer von Celebes." Abh. u. Ber. d. K. Zool. u. 

Anthr.-Ethn. Mus. zu Dresden, VII, 1898-9 (3), 42 pp., i pi. 
1900. Heller K. M. Ditto. IX, 1900 (5), 46 pp. 
* 1902. Fruhstorfer. " Tagebuchblattern " Insekten-Borse XIX, 1902, p. 28. 
1903. Fauvel, A. " Faune Analytique des Coleopteres de la Nouvelle Cale- 
donie." Rev. Ent. franc. XXII, 1903, pp. 203-378. 

1903. Zang, R. (a) " Vorlaufige Diagnosen neuer Indo-Australischer Passali- 

den" Insekten-Borse, 20 Jahrg., No. 43, pp. 338-9. 
{b) " Bemerkungen zur Alteren Passaliden-Litteratur." 
Deutsche Ent. Zeitschr., 1903, pp. 417-420. 

1904. 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," Tijdscht. v. Ent. XLVII, pp. 

1905. Zang. R. {(i) Numerous papers in Deutsche Ent. Zeitschr. for 1905. 

(b) " Anderung in der Nomenclatur der Passaliden (Coleop- 
tera)." Zool. Anz. XXIX, 1905, pp. 154-5. 

(r) " Passalidarum Synonymia. Kritische Revision der von 
Kuwert und anderen Autoren aufgestellten Gattungen 
undArten." Notes Leyden Mus. XXV, 1905, pp. 

(d) "Zwei neue Passahden aus den Gattungen Comacupes, 
Kp., und Aceraius, Kp." Notes Lej^den Mus. XXV, 

1905, pp. 233-8. 

igo6. Zang, R. (u) " Uebersicht der Basihanus-Arten." Deutsche Ent. 
Zeitschr., 1906, pp. 177-183. 
*(h) " Passalini " in " Nova Guinea, Resultats de I'Expedition 
scientifique Neerlandaise a la Nouvelle-Guinee en 
1903, sous les auspices de Arthur Wichmann, Chef de 
I'Expedition." V (i) pp. 23-26. 
1907. Arrow, G. J. "A contribution to the classification of the Coleopterous 
Family Passalidae." Trans. Ent. Soc London, 1906, pp. 441-469. 

1910. Heller, K. M. " Fiinfter BeitragezurPapuanischen Kaferfauna. "Abh. 

u. Ber. d. K. Zool. u. Anthr-Fthn. Mus. zu Dresden, XIII, 1910(3), 
42 pp., I pi. 

1911. Heller, K. M. "Fine neue Ceracupes-Art aus Formosa." Ann Soc. 

Ent. Belg. IvV, 1911, pp. 256-7, I text-fig. 

344 Memoirs of the Indian Museum. [Vol. Ill, 1914-] 

1913. Gravely, Iv H. " Preliminary Account of a revised classification of Indo- 

Australian Passalidae." J.A.S.B. VIII, 1912 (1913), pp. 403-7. 

1914. Gravely, F. H. (a) " H. Sauter's Formosa-Ausbeute — Passalidae." 

Supplementa Entomologica III, 1914, pp. 30-32. 
(h) "The Evolution and Distribution of certain Indo- 
Australian Passalidae, J.A.S.B. (In the Press). 

igT4-J F. H. Gravely: An Account of the Oriental Passalidae. 345 

13. INDEX. 

All names treated as synonyms in pt. 6 of this paper are printed in italics. Page 
numbers which refer to a key, to a full description, to synonymy, or to a figure, are 
printed in ordinary type ; other numbers are in bold face. 

The summar)^ of the paper on pp. 336-349 has been designed partly as a guide 
to the whereabouts of the principal facts recorded. References to it are not included 
in this index. 


abdominibarbatiis (Leptaulax) . . . . 303. 

abdominisculptus (L,eptaulax) . . . . 307. 

Aceraiinae .. .. .. .. 194-197, 215 etc., 280 etc., 311-3, 314, 315, 316-8, 

329, 331 etc. 

Aceraius .. .. .. .. 181, 195, 197, 228 etc., 286 etc., 312-4, 314, 316-8, 

320 etc., 332 etc. 

Aceraius .. .. .. 280, 281, 293, 299. 

addendus (Aceraius) . . . . . . 288, 322. 

aequidens (Aceraius) .. .. .. 240,293,817,323. 

albertisi (Gonatas) . . . . . . 300. 

aliicornis (Aulacocyclus) . . . . • . . 273. 

altidens (Gonatas) ., .. .. 302. 

alutaceosternus (Aceraius) .. .. .. 236,291, 323. 

Analaches .. .. .. .. 195,201,314,315. 

Analaches . . . . . . . . 195. 

analis (Leptaulacides) . . . . . . 308. 

anaulax [Leptaulacides) . . . . . . 308. 

andamanarum (Leptaulacides) . . . . 308. 

andamanensis {Basilianus, Macrolinus, Tiberius) .. 242, 294, 324. 

andamanensis [Basilianus, Tiberius) .. . . 294. 

andrewesi (Aulacocj'clus) .. .. .. 211,275,319. 

angusticornis (Comacupes) . . . . 269. 

angustifrons (Leptaulax) . . . . . . 305. 

angustior (Eriocnemis) . . . . . . 299. 

anibarbis (Leptaulax) . . . . . . 256, 303, 305. 

anipunctus (Z.e/)to/yrt«(^es, Leptaulax). . .. 256,306,326. 

anna (Leptaulax) .. .. .. 303,325,336. 

arcuatoclypeatus (Aulacocyclus) . . . • 274. 

arrowi (Ceracupes) .. .. .. 278,319,331. 

aruensis (Aulacocyclus) . . . . . . 277, 319. 

assamensis (Aceraius) . . . . 237, 292, 323. 

Aulacocyclinae . .. .. 185,188, 189, 191-3, 204 etc., 267 etc., 311, 331. 

Anlacocyclus . . . . . . 191, 192, 193, 211 etc., 272 etc., 311, 318, 319, 331. 

Aulacocyclus .. .. .. .. 267,270,277. 

Aurelius ,. .. ,. .. 202,314,315. 

34^ Memoirs of the Indian Museum. [Vol. VIII, 



Auritulus .. .. 191,192,193,279,311,319,331. 

aurivillii (Leptaulax) . . 307-8. 

austeni (Ceracupes) .. .. 212,262,278,319. 

austeni (Tiberioides) .. .. .. 216,281,320. 

barbicauda (Leplaulacides) . . .. . . 308. 

Basilianus .. .. .. .. 195, 197,280,281,284.293,316-7. 

basalis (Comacupes) . . . . 267, 318. 

batchianae (Leptaulax) . . . . 306, 307-8. 

batesi (Macroliims) . . . . . . 296. 

beccarii (Leptaulaulax) .. .. .. 251,302,325. 

bicanthatus (AiUacocycliis; Passaliis, Taeniocerus) 208, 270, 319. 

hkoloi {Le/ytaidacidi-s, t,eptau\aK, Passalus) 257, 262-295,307,326,336. 

bicolor (Leptaulax) .. .. .. 305,310. 

bicuspis (Aulacocyclus, Taeniocerus) . . 210, 271, 319. 

bihastaliis (Aiilacocycliis, Passalits) .. 272,277. 

hinominis (Basilianius) . . . . . . 283. 

birmanicus (Ophrygonius) . . . . . . 226, 285, 320. 

borealis(C/»7owa2;/.';, Tiberioides) .. .. 281,320. 

borneanus (Aceraius) .. .. .. 238,292,313,323,333. 

borneanus (Aceraius) . . 291. 

borneensis (Comacupes) . . . . 205, 268. 

homensis (Leptaulax) . . . . . . 303. 

brachyphyllus (Pleunirius) .. .. 213,262,280,320. 

brevis (Gonatas) . . . . . . 301. 

bucerus (Cylindrocaulus) .. .. .. 191,279, S^Q- 

burmeisteri [Eriocncmis, Gnaphalocneinis) . . 249, 298, 324, 335. 

caffer (Tiberius) . . . . 194 

calcuttae (Leptaulax) . . . . . . 308. 

canctus (Aceraitis, Basilianus, Tiberius) . . 280. 

cantoiiiAceratiis, Basilianus, 0-p'axygon\\xs,Passalus) 224, 262, 284, 311, 320, 331. 

caniori (Aceraius, Basilianus) .. . . 282, 285. 

Caulifer . . . . . . 1^3. 

cavicornis (^«/rtCOcyc/««s, Comacupes) .. .. 181,204,268,318. 

celebensis (Aulacocyclus) . . . . . . 277. 

celebcnsis (Leptaulax) . . . . . . 303. 

Ceracupes .. .. .. .. iso, 188, 192, 212 etc., 277 etc., 311, ,',19, 331. 

Ceratocupes .. . . . 278. 

certiis (Basilianus) .. 282, 283. 

Cetejus . .. . .. 194,201,314,315,334. 

cetioides (Gonatas) .. .. .. 301. 

ceylonicus (Aceraius) . . . . . . 288. 

Chilomazus .. .. 195,196,281,316-7. 

chinensis (Aceraius) . . . . . . 288 ^22. 

ciccalrosus (Leptaulax) . . . . 306, 307-8. 

collaris (Aulacocyclus) .. .. .. 275. 

Comacupes .. .. .. .. 181, 192,193,204,267,318,331. 

Comatus (A ulacocyclus) . . . . . . 267. 

igi4.] F- H. Gravely: An Account of the Oriental Passalidae. 347 


compergus (Kaupiolus) . . • . • • 334. 

comptoni (Accraim, Episphenus, Laches) . 218, 262-265, 281, 311-2, 314. 320. 

coMsegj«eMS (Leptaulax) .. .. • 306,307-8. 

convexifrons {Basilianns, Ophrygonius) • ■ 225, 284, 331. 

cormocems (Comacupes) . . . • • ■ 208, 269. 

comutus (Ceratocupes) .. ■ ■ ■■ 278. 

crenatipennis (Macrolinus, r«7)m«s) .. -. 244,295,324,333 

crinitus {Heterochilm) . . ■ . ■ • 286. 

cyclotaenius (Leptaulax) .. .. •• 255,305,326,336. 
cylindraceus {Aulococycliis, Comacupes, Passalus) . . 207, 269, 318, 331. 

cylindraceus (Comacupes, Passalus) .. ■■ 268, 269, 275. 

Cylindrocaulus . . . . ■ ■ • • 191. 192, I93> 279, 311, 3I9- 

darjeelingi (Leptaulax) . . . . • . 303- 

darjeilingi (L,epaulax) . . . • 303- 

dentatus (Leptaulax, P^ssrtZ/rs) .. .. 181, 185, 252, 262-265, 303, 326, 336. 

deyrollei (Aulacocyclus, Taeniocerus) .. .. 272,331. 

differens (Gonatas) . . . . • • 300. 

differ entispina (Leptaulax) . . . . • • 30?- 

dilatus (Aulacocyclus) . . • . • ■ 276, 319. 

dindigalensis (Leptaulax) . . . . . ■ 308. 

dispar (Eriocnemis) .. ■ ■ ■ ■ 299. 

dissimilis (Macrolinus) . . • • • • 296. 

divaricatus (Leptaulax) . . . . • • 3o8. 

dorsalis (Eriocnemis, Trapezocbilus) . . . . 247, 297, 324, 335. 

duivenbodei (Macrolinus) . . ' • • 294, 323. 

dunsiriensis (OTphiygonms) .. •• 225,284. 

edentulus (Aulacocyclus, Passalus) .. •• 275. 

emarginatus(Acerams, Passalus) . •• 287,288,290,291,292,293. 

Embryulcus . . ■ ■ ■ ■ ■ • 202. 

EUlaches .. ■■ ■■ 19^. 

Episphenoides .. .. •• 200, 314. 315, 333 

Episphenus .. .. •• •• 189, 194. 196, 217 etc., 281 etc., 311-3, 314. 315, 

316, 317, 320, 330. 

Eriocnemis . . . . • • • 297, 290. 

errans (Aulacocyclus) . . . . • • 275. 

eschscholtzi (Leptaulacidcs, Leptaulax) . . . . 307-8. 

evidens (Leptaulax) . . . . • • 308. 

exterris (Leptaulax) . . . . • • 303- 

fdberi (Eriocnemis) . . • • 298. 

felderi (Comacupes, Tristorthus) . . . • 267. 

filius (Analaches) . . . ■ • ■ 283. 

flachi (Chilomazus, Episphenus, Laches) . . 219, 281. 

flachii (Laches) .. .. . • . • 281. 

foveicollis (Comacupes) . . . . • • 267, 318, 331. 

foveipunctatus (Aulacocyclus) . . • . 274. 

fratermis (Macrolinus) . . . . ■ • 296. 

fratricornis (Aulacocyclus) . . . . . . 274. 

frouticornis (Ceracupes, Passa/Ms) .. .. 212,277,319. 


Memoirs of the Indian Museum. 

[Vol. VIII, 

frunlicornis (Ceracupes) 
fnihslorfen (Laches) 
fruhstor]eri [Leptaulacides) . . 
furcicornis (Passaliis) 

gelon {Eriocnemis , Plesthenus) 
geminatiis (Leptaulax) 
geminus (Leptaulax) 
generosus (Phraorles) 
germari (^«rat«s, Gonatas) 
glaber (Leptaulacides, Leptaulax) 
glabricollis (Leptaulax) 
glabriusculus (Aulacocyclus) 
glabriveiUris (Leptaulax) 


Gnaphalocneuiis group 


Gonatas group . . 

grabowskyi (Cetejus; 

gracilis (Hyperplesthenus, Z.rti(V«i/s) 

grandis (Aceraius, Passaliis) 

gravid us (Pelopides) 

hansemanni (Leptaulax) 
helferi (Aceraius) 
heynei (Protomocoelus) 
himalayac (Leptaulax) 
himalayensis (Aceraius) 
hirsutus (Aceraius) 
humerosus (Leptaulax) 
humerosiis (Leptaulax) 
Hyperplesthenus group 

ignolus (Eriocnemis) 

illegalis (Aceraius) 

imitator (Taeniocerus) 

impar (Hyperplesthenus) 

inaequalis (Aceraius, Busilianus, Oprygouius, Pas- 

inaequalis (Bassilianits) 
incidens (Aceraius) 
incipiens (Leptaulax) 
indicus (jB(/si7a«»s, Episheiius) 
indictts (Leptaulax) 

infantilis (Aceraius, Analaches, Epilacbes, Lacfns) . . 
innocitus (Passat us) 




303, 306. 



250, 301, 313, 325. 

307, 326. 



255 -• 303, 338. 

191, 194, 195, 197-203, 246 etc., 297 etc., 313-6, 

191, 202, 24S etc., 298 etc., 314, 318, 324, 329, 335. 
199, 202. 

203, 250 etc., 299 etc., 313, 314, 318, 325. 
199, 201. 

202, 334. 

181, 231, 262-265, 28S, 312-3, 314, 322, 332. 
201, 203, 298. 


238, 292, 323, 333. 

195, 286. 


256, 305, 336. 

236, 292, 323. 

180, 181, 231, 263, 288, 322, 332. 

251, 262, 302, 325, 335. 

302, 303- 

194. 202, 314, 315, 328, 334. 

199, 201. 


230, 287, 321. 



227, 285, 320, 332. 




220, 262-265, 282, 311-2, 314, 320. 


283, 330. 


1914-] F- H. Gravely: An Account of the Oriental Passalidae. 3.49 


insignis (Leptaulax) . . . . . . 303. 

insipiens (Leptaulax) . . 308. 

interponendus (Leptaulax) . . . . . . 303. 

interrogationis (Basilianus) . . . . . . 284. 

javamis (Macrolinus) . . . . . . 296. 

kaupi (Aulacocyclus) . . . . , 275. 

kuupii (Aceraius) . . . . . . 292. 

Kaupioloides . . . . . . . 200, 314, 334. 

Kaupioloides group . . . - ■ ■ 199, 200. 

Kaupiolus . . . . . . 202, 314, 334. 

Kaupiolus group . . . . . . 198. 

kuwerti (Aceraius) .. .. .. 235,290,322. 

kuwerti (Tiberioides, Tiberias) .. . . 215, 226, 280, 320, 332. 

Labienus .. .. .. .. 191,202,314,315,334. 

Laches .. .. .. .. 195, 281, 330. 

laevicolUs (Aceraius, Passa/»s) .. .. 230,287,313,322,332. 

laevicollis (Aceraius) . . . . . . 293. 

laevicornis (Aulacocyclus , Qom^cn^Q?,) .. .. 181,205,268. 

laevimargo (Aceraius) .. .. 235,291,323,333. 

langier (Aceraius) .. .. .. 234,290,322. 

laterisculptm (Passalus) . . . ■ ■ 299. 

latipennis (Macrolinus) . . . . . . 245, 296, 324. 

Leptaulacides .. .. .. -. 203,204,302. 

Leptaulacinae .. .. .. ' •• 191, 194, 199, 203-204, 251 etc., 302 etc., 311, 329, 


Leptaulax .. .. .. •• 181, 185, 189, 191, 203, 204, 251 etc., 302 etc., 

318, 325-6, 330, 335-6. 

Inzonicus (Aceraius) . . . . • • 287. 

macassariensis ^Leptaulax) .. .. .. 256,305,325. 

Macroliuinae .. .. .. .. 194-195, 197, 240 etc., 293 etc., 311, 329, 333. 

Macrolinus . . . . . . . . 181, 197, 240 etc., 293 etc., 318, ^,2^, 333. 

magnus (Aceraius) . . . . • . 28S. 

major (Gonatas).. .. .. .. 300. 

malaccae 1 Leptaulax) . . . . . . 307-8. 

mahfiosus (Leptaulax) . . . . . . 303. 

manillae (Leptaulax) . . . . . . 307. 

marginepunctaius (Passalus) . . . . . . 296. 

masoiii (Comacupes) .. .. .. 207, 2()8, 318. 

niasoni (Comacupes) . . . . . . 269. 

Mastachilus . . . . . . . . 195. 

mastersi (Aulococyclus, 7'rt(;««oct.THi) .. .. 193,273. 

Mastochilus .. .. .. 178, 195, 200, 314, 334. 

maxillonoius (Leptaulax) . . . . . . 307-8. 

medius (Leptaulax) . . . . . , 307-8. 

meyeri (Aceraius) . . . . . . 287. 

minimus (Omegarius) . . . . . . 335. 

minor (Aceraius).. .. .. .. 240,293,317,323,333. 


Memoirs of (he Indian Miiseiivi. 

[Vol. VIII, 

minor (Cotnacupes) 

minutifrons (Aceraius) 

mixtus (Ivcptaulax) 

tiiniszechi (Eriocueniis, Giiai)halocneinis) 

moeschlc'i (Aceraius) 

nwlossus (Aceraius) 

moluccanus (Kaupiolus) 

monticulosus (Aceraius, Kriocncmis, Gnaphalociiemis, 

moorei (Episphenus) 
morator (Leptaulax) 
moschleri (Aceraius) 

nanus (Aceraius) 

naviculator (Gonatas) 

ncelgherriensis (Aceraius, Basilianus, Episphenus, 

Passalus ) . . 
ncelgherriensis (Aceraius, Basilianus) . . 

iiiae (Leptaulax) 
nicobaricus (Aceraius) 
nicobaricus (Macrolinus) 
nictnen (Leptaulacides) 
nikobaricus (Passalus) 
nobilis (Phraories, TiaY)Qzoch'i\us) 
novaebritanniae (Gonatas) . . 
novaeguineae (Letaulax) 

obtusidens (Leptaulax) 

occulidens (Aceraius) - • • 

oculitesselatus (Hekrochihis) 




orbatus (Macrolinus) 
orientalis (Passalus) 
oroleius (Leptaulax, Ophrygonius, Passalus) 

palawanicus (Leptaulacides) 
palawanus (Aceraius) 
papua (Leptaulax) 
papuanus (Letpaulacides) . . • 
paradoxus (Tarquinius) 
parallelipcnnis (Macrolinus) 
parallelogrammilrons (Laches) 
Parapelopides . . 
parreyi (Aulacocyclus) 
parryi (Aulacocyclus) 
parryi (Aulococyclus) 









249, 298, 325, 335. 

217, 281, 311, 312, 315, 320. 


229, 287, 321. 


301, 313, 325. 

222, 270, 2S3, 311. 314. 320, 330. 





241, 294, 324. 



247, 297. 


306, 326. 

307. 326. 

234, 290, 312-3, 322. 



203, 314, 335. 

196, 224 etc., 284 etc., 311, 312-3, 3: 

r4. 316-7, 

318, 320, 331. 





327, 335. 

202, 246, 
276, 277. 
276, 319. 

297. 314. 324. 334. 


F. H. Gravely: An Account of the Oriental Passalidae. 


parvus (Taeniocerus) 

^&ta\\s {Aulacocyclus, Aiiri/iiiliis, Auritulus) 


pearsoni (Episphenus) 

pectinigera (Epispheiioides) 


Pel op ides 


perakensis (Aceraius) 

percheroni (Aulacocyclus) . . 

percheronii (Aceraius) 

peregrinus (Gonatas) 

perlatus (Aulacocyclus) 

perplexus (Pseudepispheaus) 

perturbans (Taeniocerus) . . - 


philippinensis (Paxilloides) . , 


pilifer (Aceraius) 

pilifer (Aceraius) 

pilipes (Pleurarius) 

planicollis (Leptaulax) 

planus (Leptaulacides, I^eptaulax, PtissaJiis 

planus (Leptaulax) 

platypus (Aulacocyclus, Taeniocerus) 


Plestheuus group 



politus (Mastochilus) 

polyphyllus (Mastochilus) . . 

pontifex [Aceraius) 


prosternisulcatus (Aceraius) . . 

Protomocoelus . . 

Protomocoelus group 

Pseudepisphenus . . ' 

puella (Laches) . . 

puerilis (Aceraius, Laches) . . 

pugaax (Aulacocyclus, Comacupes) 

pulchellus (Leptatdacides) 

punclifrons (Comacupes, Passaius) . . 

P3 gmaeus (Aulacocyclus, Taeniocerus) 

quadridentatus (Passaius) . . 
quadrifer (Ophrygouius) 
quaestionis (Episphenoides) 

rectidens (Aceraius) 
redtenbacteri (Semicyclus) :. 



279. 319, 331. 


218, 2S1. 


197, 199, 201, 314. 

201, 298. 

229, 287, 321, 323, 332. 







200, 314, 334. 



235, 291,323, 333. 


279, 320. 


260, 262, 310, 326. 

270, 319. 

202, 314. 

194, 199, 213 etc., 279 etc., 311, 
194, 213 etc., 279 etc., 320. 

191, 199, 201, 314, 334. 
199, 200. 
327, 329, 330. 

283, 330. 
209, 271, 319. 


233, 288, 322, 332. 


Memoirs of the Indian Museum. 

[Voi. \'TII 

respedabilis (Phraorles, Trapezochilus) 

.. 248.297. 

reticnlaticoUis (Aceraius) 

.. 291. 


. . 286. 

rilhei (Leptaulax) 

.. 303. 306. 

roepstorfi (Leptaulax) 

. . 260, 309, 326. 

rocpslorfi (Leptaulax) 

.. 308. 

rosenbergii (Aulacocyclus) . . 

• • 274, 319. 

rotundatoclypeatiis (Aulacocyclus) 


rotuudifrons (Macrolinus, Tiberius) 


rugulosus (Leptaulacides) 

.. 308. 

saigonicHs (Leptaulax) 

• ■ 303. 

schellongi (Gonatas) 

. . 300. 

schenklingi (Analaches) 

. 200. 

schmidtii (Paxilloides) 

. . 178. 

schraderi (Pelopides) 



. . 178. 

separandus (Le])taulax) 

■ ■ 307-S. 

sequens (Leptaulax) 

.. 308. 

sikkimensis (Basilianus, ^L^crolinus Tiberius) 

• • 243, 394. 323- 

simplex (Giiaphalocnemis) . . 

. . 248, 298, 324. 

singapurae (Ophrygonius) . . 

. . 226. 285, 317. 320. 

singhalensis (Macrolinus, T/ienHs) 


sinkepiciis (B.isilianits) 

. . 286. 

socius {Laches) 

. . 281. 

stoliczkae (Basiliauiis) 

. . 282. 

stoliczkae (Comacupes) 

206, 268, 318. 

submedius (Leptaulax) 

•■ 303- 

subsequens (Leptaulax) 

.. 308. 

stilcatipoiis (Aulacocyclus) 

• • 273. 

sulciperfectus (Macrolinus) 

. . 296, 324, 333. 

sumatrae (Leptaulax) 

. . 308. 

symmetricus (Para pelopides) 

.. 246.297,324,334. 


. . 192, 193, 208 etc., 270 etc., 319. 


. . 272. 


. . 329, 335. 


178, 327, 328, 329 330, 335. 


197, 203, 314. 

tavoyamis (Aceraius) 

• • 237, 292. 

tavoyanus (Macrolinus) 

• • 243, 294 

lenasserimensis (Leptaulax) 


tereoides (Aulacocyclus) 

• • 273. 

teres (Aulacocyclus, Passaliis) 

• • 273. 

thoreyi {Leptaulax, Trichostigmus) 

.. 261,310,326. 


. . 189, 196, 215 etc., 280 etc., 311 3, 314. 



.. 194,195,197,280,293. 

tibialis (Gonatas) 

. . 300. 

timoriensis (Leptaulax, Passalus) 

.. 303,326,336. 

timoriensis (Leptaulax , Passalus) 

• • 303. 

igi4.] F. H. Gravely: An Account of the Oriental Passulidac. 353 

tonkinensis (Leptaulax) 



trlcortiis (Aceraius) 

tricuspis (Tristorthus) 

tridens {Eriocnemis, Gnaphalocnemis, Ocytlioe, Pas- 

Iridens (Eriocnemis) 
tridentatus (Gonatas) 
trigonophorus (Kaupioloides, Kaiipiolus) 

ursulus {Leptaulax, Trichostigmus) 
urus (Macrolinus) 

variolosus (Aulacocyclus) 


vicinus (Passalus, Leptaidacides , Leptaulax) 

wallacei (Aceraius, He/e^-oc/jz'/Ms) 
waterhousei (Macrolinus, Tiberius) 
weberi (Macrolinus) 
weberi (Macrolinus) 
westermanni (Comacupes) 

















































. 324 



yorliensia (Gonatas) 


Fig. I. — Comaciipes cavicornisvax.borneensis (type). Head from above, x 2: 
,y I"- Ditto Head from the side, x 2. 

2.—Comacupes cavicornis var. laevicornis (regd. no. "f ). Head from above, 
X 2. 

-«• Ditto Head from the side, x 2. 

26. Ditto Mentum, etc., x 4. 

„ i.—Comacupesstoliczkae {type}. Head from above, x 2. 

3«- Ditto Head from the side, X 2. 

^■—Comaciipes masoni (type). Head from above, x 2. 

4''- Ditto Head from the side, x 2. 

5-—Com(icHpes cylindracciis (regd. no. 'H.'). Head from above, x 2. 

5". Ditto Head from the side, X 2. 

6.— Another form of the sanie species (from a specimen in the British 
Museum). Head from above, x 2. 

f^c- Ditto Head from the side, X 2. 

y.—Taciiioceyus bicanthatiis (regd. no. •=*«-). Head from above, x 2. 

7«- Ditto Head from the side, x 2. 

7*- Ditto Mentum, etc., X 4. 

S.—Taeniocerns pygmaeiis (regd. no. ^). Head from above, x 2. 

8rt. Ditto Head from the side, x 2. 

g.—Taeniocenis bicuspis (regd. no. '^). Head from above, x 2. 

9«- Ditto Head from the side, X 2. 

,, io.~Aulacocychis andrewesi (from the type in Mr. Andrewes' collection). 

Head from above, x 2. 
.. io«. Ditto Head from the side, X 2. 

,, II. — Ceracupes misteni (type). Head from above, x 2. 
,. ii«- Ditto. Head from the side, x 2. 

,, i2.~Ceracupes fronticornis (regd. no. «f' ). Head from above, x 2. 
,, 13.—Pleurarius brachyphyllns (regd. no. «5f). Head from above, x 2. 
,, I3«- Ditto Mentum, etc., x 4. 

,, 14.— Tiberioides kuwerti [xQgA. no. ^'^^). Head from above, x 2. 
„ 15- — Tiberioides misteni (type). Head from above, x 2. 
M I5«- Ditto Mentum, etc., x 4. 

,, 16.— Episphenus moorei (regd. no. ^"). Head from above, x 2. 
,. i6fl. Ditto Mentum, etc., x 4. 

„ 17.— Episphenus peai'soni' (type). Head from above, x 2. 
„ 18.— Episphenus comptoni (regd. no. °-f^"). Head from above, x 2. 
,, i8a. Ditto Mentum, etc., x 4. 

,, ig.—Epispheftus comptoni var. flachi (regd. no. *•,';" and 5lli'). Frontal area, 

etc., X 3, showing the variability of the shape of this area. 
,, iga. Abnormal mentum of another specimen (regd. no. -'^f) x 4. 
,, 20.— Episphenus indtcus, large specimen (regd. no. '•^). Head from above, x 2. 
,. 20a. Ditto Mentum, etc., X 4. 

,, 206. Same species, small specimen (from a specimen in the Bombay Natural 

Hi.story vSociety's collection). Head from above, x 2. 
,, 21.— Episphenus neelghcrricnsis (regd. no. ';'.f). Head from above, x 2. 
,, 2ia. Ditto Mentum, etc., x 4. 

,, 22.— Ophrygonius cantori (regd. no. '^"). Head from above, x 2. 
,, 22fl. Ditto Mentum, etc., X 4. 

,, 2^.— Ophrygonius cantori var. dunsiriensis (type). Mentum, etc., x 4. 

' E. moorei, see above, p. 281. 

Mem. Ind. Mus.,Vol. lll,4«*e-. 

Plate XI. 

B«mros«,Collo., Derby 



Fjg. 24.— Ophtygonius inaeqnalis (regd. no. '^). Head from above, x 2. 
24a. Ditto Mentum, etc., X4. 

25. — Aceraius borneantis (regd. no. --^). Head from above, x 2. 
,, 25«. Ditto Left mandible from the side, x 4. 

,, 256. Ditto Mentum, etc., x 4. 

,, 26. — Aceraius wallacei (regd. no. ~). Head from above, x 2. 

26a. Ditto Left mandible from the side, X 4. 

,, 266. Ditto Mentum, etc., x 4. 

,, 27.— Aceraius laevicollis (regd. no. ?|5). Head from above, x 2. 
,, 27^. Ditto Left mandible from the side, x 4. 

,, 28. — Aceraius grandis subsp. hirsuius, a specimen of moderate size (regd. 
no. "^). Head from above, x 2. 
28a. A large specimen of the same species and subspecies (regd. no. ^^'). 
Head from above, x 2. 
,, 28b. Left mandible of a similar specimen to the last from above, x 4. 
, , 29. — Left mandible of another specimen of the same sub-species (regd. no. ^^) 

from above, x 4. 
,, 30.— Aceraius grandis var. rectidens (from a specimen in the Sarawak Museum 

collection). Head from above, x 2. 
,, 31. — Aceraius kuwerti (from a specimen in Mr. Audrewes' collection). Head 

from above, x 2. 
,, 32. — Aceraius illegalis (regd. no. ^'-f). Head from above, x 2. 
„ 32c7. Ditto Mentum, etc., x 4. 

,, 33- — Aceraius moschleri (regd. no. •^). Head from above, x 2. 
,, 34- — Aceraius alutaceosternus (regd. no. ^'). Head from above, x 2. 
,, 34rt. Ditto Mentum, etc., x 4. 

,, 35- — .4 ceyfl/»s />^7^;■/ey (from a specimen in Mr. Andrewes' collection). Head 

from above, x 2. 
,, 36.— ^cemm,9 /n'wrt/aymsts (type). Head from above, x 2. 
,, 36rt. Ditto Anterior part of head from the right side, x 4. 

37. — Aceraius assamensis {xegA. no. ~). Head from above. 
,, 37^^ Ditto Anterior part of head from the right side, X 4. 

38. — Aceraius iavoyanus' (regd. no. *•?). Head from above, x 2. 
,, 38fl. Ditto Anterior part of head from the right side, x 4. 

,, 3g.— Aceraius helferi (regd. no. -^i"). Head from above, x 2 

y)a. Ditto Anterior part of head from the right side, X 4. 

A. helferi, .see above, p. 292. 


Mem. Ind. Mus., Vol. Ill, iW8? 

Plate Xll. 




Fig. 40. — Macro/ill us nicobaric7is (typQ). Head from above, X2. 

,, 40a. Ditto Anterior part of head from the side, X4. 

,, 41- — Macroliniis andcimdttoisis (regd. no. ■'^'^''). Head from above, X2. 

>, 41^- Ditto Anterior part of head from the side, x 4. 

,, 42. — Macrolinus sikkimensis {tYYte). Head from above, X2. 

,, 42a. Ditto. Mentum, etc., X4. 

,, 43- — Macrolinus rotnndijrons (regd. no. "j'f). Head from above, X2. 

,, 44.—Mactolinus waterhousei (regd. no. -^^). Head from above, X2. 

,, 45. — Macrolinus latipennis {regd. no. ^'^f). Head from above, X2. 

,, 46. — Another specimen of the same species (regd. no. -j-"). Head from 

above, X2. 

,, 47. — Gonatas germari (regd. no. ^|f). Head from above, X2. 

,, 47a. Ditto Mentum, etc., X4. 

,, 48 — Trapezochilns dorsalis (regd. no. %"'). Head from above, X2. 

,, 49. — Gnaphalocnemis bmmeisteri {regd. no. '^^f). Head from above, X2. 

,, 4g«. Gnaphalocnemis monticulostis (Tegd. no. ^^'■■). Head from above, X2. 

,, 50. — Gnaphalocncmis tridens (from a specimen in Mr. Andrewes' collection). 

Head from above, X2. 

,, 50rt. Ditto Mentum, etc., X4. 

,, 51. — Lepimdax humerosits {regd. no. ^-^). Head from above, X2. 

,, 52. — Leptaidax dentains, large specimen (regd. no. ^"). Head from above, 

X 2. 
,, 52rt. Ditto Mentum etc., x 4. 

,, 526. Frontal areas of some specimens of the same species (regd. nos. ^-, ^, 

*{"), all taken from one colony, x 2. 
,, 52c. Frontal areas of specimens of the same species (regd. nos. '*', ^,^, ^) from 

other colonies, x 2. 
,, 52^. Small specimen of the same species (regd. no. yi'). Head from 

above, X2. 
,, 53. — Leptaidax cyclotaenius, s. str. (regd. no. "^). Head from above, x 2. 
,, 54.— Leptaulax macassariensis subsp. anibarlis (regd. no. ";"''). Head from 

above, x 2. 
,, 55. — Leptaidax anipunctus (regd. no. ^^). Head from above, x 2. 
,, 56. — Leptaulax bicolor, s. str., large specimen (regd. no. —''). Head from 

above, x 2. 
,, 56rt. The same species (var. vicinus), small specimen (regd. no. ■^'). Head 

from above, x 2. 
,, 57- — Leptaulax roepstorfi (regd. no. i^^). Head from above, x 2. 
,, 58. — Leptaidax planus (regd. no. 'ff). Head from above, x 2. 


Mem. Ind. Mus.,Vol. Ill, •*#*&■ 

Plate Xlll. 

40 a: 'fp 



%MU ^^ '^'^ 

Bemrose.Collo, Derby. 


Vol. VIII, 1912. 

Zoologiciii Results of the Abor Expedition, 1911-12. 
Part /.— Introduction. BaUachia. Reptilia. Lepidoptera. Porifera. Scolopendridae. Hymenoptera. Authoplula. 
Part II. — Hammals. Coleoptera, I-III. Arachuida, I. Dermaptera. Diptera. 
Part in. — Coleoptera. IV-VI. Thysauoptera. Arachnida, II. MoUusca, I. Temnocephalidae. Hymenoptera II, 

III. Birds. Crustacea Decapoda. Orthoptera, I. Freshwater Planaria. 
Part /F.— Hymenoptera. IV. Orthoptera, II. Odonata. Neuropteroid In.sects. Reptilia (Supplement). Mollusca. 

II. Oligochaeta. 
Pari r.— Coleoptera, VII. Orthoptera, III. Termitidae. Lepidoptera, II. Hymenoptera, V. Crustacea Amphipoda. 

Vol. IX, 1913. 

Part I. — Notes ou the collection of Coccidae in the Indian Museum, II. New Indian Empidae. 

Part II. — A new variety of freshwater crab from Travancore. Ou some Thysanura in the Indian Museum. The 

Tortoises of Chota Nagpur. Notes from the Bengal Fisheries Laboratory, Indian Museum, No. I. Cryptostoraes 

from the Indian Museum, 
Part III. — Nouvelle etude sur les Chironomides de I'lndiau Museum de Calcutta. Description de quelques nouvelles 

Cecidomyies des Indes. Description d'uu nouveau Mymaride des Indes Orientales. The Anthribidae in the Indian 

Museum. Description of a new Sand-boa from the Persian Gulf. 
Part IV. — A note on certain Ophiuroids in the Indian Museum. The Indian Barnacles of the subgenus ScalpeUuin. 

Notes on Freshwater Sponges, XV. Preliminary Note ou a new genus of Onychophora from theN.-E. Frontier of 

India. Miscellanea: — Some Salps taken by the R.I.M.S.S. "Investigator" in the Bay of Bengal and Andaman 

Sea. Adaptation in the Habits of a Tabanid Fly. Indian Blood-sucking Midges. Further Notes on the Habits 

and Distribution of Limnocaida indica. 
Part F.— On a small collection of Birds from the Mishmi Hills, N.-E. Frontier of India. New and interesting Diptera 

from the Eastern Himalayas. On some Indian Cestoda, Pt. I. Some new and interesting Batrachia and Lizards 

from India, Ceylon and Borneo. The Indian Geckos of the genus Gymnodactylus. Butterflies of the genus 

Paf«05Sj«s in the collection of the Indian Museum. Nemestriniden (Dipt.) aus dem Indian Museum in Calcutta. 

Miscellanea :—A subfossil Bat's skull from Rodriguez I A list of the Reptiles obtained by Mr. H. Stevens in Upper 

-issam and the Eastern Himalayas. Notes on Plankton from the Chilka Lake. 

Vol. X, 1914. 

Part I. — On the Pseudoscorpions of the Indian Museum. Critical Review of genera in Culicidae. Further records of 
Indian brackish water Mysidae with descriptions of a new genus and species. 

Part //.—Notes on Crustacea Decapoda in the Indian Museum. V. Notes on Indian Fish, I-II. Further notes on the 
sponges of Lake Baikal. Fauna Symbiotica Indica, V. 

Pari III.— On a new species of Blepharocerid fly from I'ashmir, with a description of some larvae from the same locality. 
Studies in Indian Helminthology, No. I. Studies in Indian Helininthology , No. II. On some new terrestrial Isopods 
from the Andaman Islands and Southern India. Miscellanea .-—'Sote on the genas A naclinia. Metamorphosis of 
Zoanthella. Change of name io an Indian genus of Echinoids. Note on some Amphipods collected on the Pamirs 
at an altitude of 15,000ft. Xenapsylla iiesiotes : a correction. 


of the 


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