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THOMAS LINCOLN 
CASEY 
LIBRARY 
1925 


Oa ‘Ap Account of the ee 
AL -PASSALIDAE. (COLEOPTERA \) 


_ based primarily on 


oe : : a8 ~ ae By os 
Re “GRAVELY, M.Sc. Sect 


Assistant Supe dg: Indian Museum. 


PUBLISHED BY ORDER OF THE TRUSTEES OF THE INDIAN MUSEUM. 


Calcutta : 
PRINTED AT THE BAPTIST MISSION PRESS. 


SEPTEMBER, IQT4. | ste 


‘Price, Five Rupees. 


Miscellaneous Zoological Publications. 


Rs. As. Rs. As. 
Account of the Deep-sea Brachyura collected by Echinoderma of the Indian Museunn: Littoral 
the R.I.M. S. ‘‘ Investigator.’’ By A. Alcock, Holothurioidea collected by the R.I.MS. 
M.B., C.M.Z.S * 6 0 ‘* Investigator.’’ By R. Koehler and\C. Vaney 2 0 
Account oe the. Despre Madreporaria collected Echinoderma of the Indian Museum: Deep-sea 
by the R.I.M.S, ‘‘ Investigator.’’ By A. Alcock, Ophiuroidea collected by the R.ILM.S. ‘* Investi- 
M.B., C.M.Z.S. ig ae Wea Bay gator.’’ By R. Koehler 3 .. 10¥'0 
Account of the Triaxon (Hexactinellid) sponges Echinoderma of. the Indian Museum: Shallow- 
collected by the R.I.M.S. ‘‘ Investigator.’’ By water Ophiuroidea collected by the R.I.M.S. 
F. E. Schulze, Ph.D., M.D. .. os auto" 0 ‘ Investigator.”’ By R. Koehler at 40 


Account of the Alcyonarians collected by the Echinoderma of the Indian Museum, Part v: 
R.IMS. ‘‘Investigator.’’ PartI. By J. Arthur An account of the Deep-sea Asteroidea col- 
Thomson, M.A., and W. D. Henderson, M.A., lected by the R.I.M.S. ‘‘ Investigator.’’ By R. 

B.Se. es +: Ap 20 : +. 16 0 Koehler .. + ee eee) 

Account of the Alcyonarians collected by the Echinoderma of the Indian Museunt, Part VI: An 
R.LMS. ‘‘Investigator.’’ Part. By J. Arthur account of the Shallow-water Asteroidea. By R, 


Thomson, M.A., and J. J. Simpson, M.A., B.Sc. 20 0 oehler “Shee 
Aids to the identification of Rats connected with ) ah 
Plague in India. By W. C. Hossack, M.D. B'S Echinoderma of the Thdian Mubenin, ‘Part VI: 


Annotated List of the paietie Beetles in the Indian The Crimoids of the Indian Ocean. By A.H,Clark 20, 0 


Figures and Descriptions of nine Species of Squil- 
Pounielioe! sy et "Anandale, De, aud W. ; lide from the Collection of the Indian Museum. 
ia R Pid rn By J]. Wood-Mason, F.Z.S., ete., edited es 
Biological tollec tions: "OE the ope T.M. Ss. *« Investi- A. Alcock, M.B., C. M. ZS... 240 
Guide to the Zoological Collections exhibited in 
gator.’’ List of Stations, 1884-1913 HeakRirdiGallesce ak ii 
Catalogue of Indian Crustacea. Part T.—Introdue- bagel’ ns + Se Sah ag Indian ‘Museum. ee 
tion and Brachyura Primigenia. By A. Alcock, inn, BA EZS + Pits 
M.B., LL.D., F-R.S. 7! 0 Guide to the oolouia al Collections eaniited in 
Catalogue of Indian Decapod Crustacea. Part I.— . the Fish Gallery of the Indian Museum, By 
Brachyura. Fasciculus II.—Indian Freshwater A. Alcock, M.B., C.MLZ.S. .. o 8 
Crabs—Potamonide. By A. Alcock, C.1.E., Guide to the Zoological Collections exhibited in the 
M.B., LL.D., F.R.S td OO: Invertebrate Gallery of the Indian Museum. 
Catalogue of the Indian Decapod Crustacea, Part | By A. Alcock, M.B., C.M.Z,S. (Out of print.) 
JI.—Anomura. Fasciculus I.—Pagurides. By Guide to the Zoological Collections exhibited in the 
A. Alcock; M.B., LL.D., F.R.S., C.LE. 1470.14 Reptile and Amphibia Gallery of the Indian Mu- 
Catalogue of the Indian Decapod Crustacea. Part | seum. By <A. Alcock, M.B., C.M.Z.S. (Out of 
I1.—Macrura. Fasciculus I—The Prawns of print.) 
the Peneus Group. By A. Alcock, M.B., LL.D., ' Hand List of Mollusca in the Indian Museum 
ERS. CLE, 4 7 0 Parts I and II, and Fasciculus E. By G. Nevill, 
Catalogue of Indian Deep-sea Crustacea : : Decapoda C.M.Z.S., etc. Index Parts I and II. By W. 
Macrura and Anomala in the Indian Museum. Theobold . w 4c en 
By A. Alcock, M.B., LL.D., C.M.ZS.. 10 0 Illustrated Catalogue of Asiatic Horns and 
Catalogue of Indian Deep-sea Fishes in the Indian | Antlers in the Indian Museum. By 'T. Bentham 2 o 
Museum. By A. Alcock, M.B., C.M.Z.S. .. § © |. List of Batrachia in the Indian Museum, By W. L. 
Catalogue of Mammalia in the "Indian Museum, Sclater, M.A., F.Z.S. Y eMe ay th 
Part I. By J. Anderson, M.D., LL.D., F.R. s. List of Birds in the Indian Museum. "Part I— 
Part II. By W. L: Sclater, M. A. FZS. SONG, os} Corvide, Paradiseide, Ptilonorhynchide and 
Catalogue of Mantodea in the Indian Museum, Crateropodide. By FP. Finn, B.A., F.Z.S. <0 the 
Parts I and II. By J. Wood-Mason, F.Z.S.,etc. 2 0 | List of Snakes in the Indian Museum. By W. L. 
Catalogue of Moths of India, Parts I to VII. By ) Sclater, M.A., F.Z.S. 10 
E. C Cotes and C. Swinhoe, E.L:S. F.Z,S., ete. 5 12 Monograph of the Asiatic Chiroptera and Catalogue 
Echinoderma of the Indian Museum : Account of of the Species of Bats in the Indian Museum. 
the Deep-sea Holothurioidea collected by the By G. E. Dobson, M.A., M.B., F.R.S . pe 
R.I.M.S. ‘‘ Investigator.’ wi R. Koehler and | Monograph of the Oriental Cicadide, “Parts I to 
C. Vaney .. i elo. [10 Vil. By W. L. Distant, F.E.S Se 22 Gly 14 


The above can be obtained from the Superintendent of the Indian Museum, Calcutta, and from Messrs. Friedlander 
& Sohn, 11, Carlstrasse, Berlin. 


Other Publications edited and sold by the Superintendent of the Indian Museum (also 
obtainable from Messrs. Friedlander & Sohn) issued by the Director 
of the Royal Indian Marine. 


Illustrations of the Zoology of the R.I.M.S. ‘‘ Investigator,’’ 1892. Fishes, Plates I to VII. Crustacea, Plates 
I to V, 1894. Fishes, Plates VII to XIII. Crustacea, Plates VI to VII. Echinoderma, Plates I to III, 1895. 
Echinoderma, Plates IV and V. Fishes, Plates XIV to XVI. Crustacea, Plates IX to XV, 1896. Crustacea, 
Plates XVI to XXVII, 1897. Fishes, Plate XVII. Crustacea, Plates, XXVIII to XXXII. Mollusca, Plates I to 
VI, 1898. Fishes, Plates XVIII to XXIV. Crustacea, Plates XXXIII to XXXV. Mollusca, Plates VII and VIII, 
1899. Fishes, Plates XXV and XXVI._ Crustacea, Plates XXKVI -to (XLY,, 1900. Fishes, Plates XXVIT to 
XXXV. Crustacea, Plates XLVI to XLVIIJ. Index, Part I, 1901. Crustacea, Plates XLIX to LX. Mollusea, 
Plates IX to XIII, 1902. Crustacea, Plates LVI to LXV Ii. Crustacea, Plates LXV Ill to LXXVI._ Fishes, Plates 
XXXVI to XXXVIII, 1905. Crustacea (Malacostraca), Plates LXXV Il to LXXIX. Crustacea (Entomostraca), 
Plates J and IJ. Mollusca, Plates XIV to XVIII, 1907. Fishes Plates XXXIX to XLIJI. Crustacea (Entomostra- 
ca), Plates IJ] to V. Mollusca, Plates XIX and XX, 1908.—Re. 1 per plate. Mollusca, Plates XXI to XXIII, 1909.— 
As. 6 per plate. 


RECORDS 
of the 


INDIAN MUSEUM 


(A JOURNAL OF INDIAN ZOOLOGY) 


Vol. I, 1907. 


Part I.—Contributions to the Fauna of the Arabian Sea. Records of Hemiptera 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 Stratiomyide. Description of an Oligochete Worm allied to Chetogaster. The 
Fauna of Brackish Ponds at Port Canning, Lower Bengal, IV. Further Note on a Polyzoon from the Himalayas. 
Reports on a collection 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 IL. Miscellanea. 


Part III.—Report on the Marine Polyzoa in the collection of the Indian Museum. The Fauna of Brackish Ponds at 
Port Canning, Lower Bengal, VI. A third note on Earwigs (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- 
Jection-of marketable fish from Akyab, with a description of a new species of Lactarius. Description of two fresh- 
water Oligochete 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 Nycteribiide from India. Annotated Catalogue of Oriental Culicide. 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 list 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 Siam. Descriptions of new species of Marine and 
Freshwater Shells in the collection of the Indian Museum. Notes on Oriental Syrphide, I. Description of a new 
variety of Spongilla lovicata. Notes on Oriental Diptera, V. Miscellanea. 


Part II.—Gordiens du Musée Indien. The Fauna of Brackish Ponds at Port Canning, Lower Bengal, IX. Description 
of a new species of Danio from Lower Burma. Rhynchota Malayana, I. Cimex votundatus, Signoret. Notes on 
Freshwater Sponges, IX. Fruit Bats of the genus Pferopus inhabiting the Andaman and Nicobar Archipelagos. 
A new species of Sun-Bird obtained near Darjiling. Three Indian Phylactokemata. On two new species of Bagle- 
Rays (Myliobatide). Description of a new species of the genus Sesayma, Say., from the Andaman Islands. Des- 
criptions of new species of Land, Marine, and Freshwater Shells from the Andaman Islands. 


Part I[J.—The Fauna of Brackish Ponds at Port Canning, Lower Bengal, X, XI. On some Oriental Solifugee with 
descriptions of new forms. The difference between the Takin (Budorcas) 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 Charaves from the Bhutan Frontier. First Report on the Collection of Culicidee and Corethride 
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 Psychodide. Description of a new species of mouse from 
the Madura District, Madras. Some Cleride of the Indian Museum. 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 Tyygon from the Bay of Bengal. New Micro-lepidoptera from India and Burma. Notes on some 
Chrysomelid Beetles in the collection of the Indian Museum. Six new Cicindeline from the Oriental Region. Des- 
cription of a new slug from Tibet. 


Part V.—Revision of the Oriental Leptide. Revised and annotated Catalogue of Oriental Bombylide, with descriptions 
of new species. 


Vol. III, 1909. 
Part I.—The Races of Indian Rats. 


Part II.—Notes on Freshwater Sponges, X. Report on a collection of aquatic animals made in Tibet by Capt. F. H. 
Stewart in 1907, IJ. Note on some amphibious Cockroaches. Description de quelques nouvelles Cécidomyies 
des Indes. Description of new land and marine shells from Ceylon and S. India. Description of two new species 
of Caranx 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 III.—Notes on the Neuroptera in the collection of the Indian Museum. New Indian Leptide and Bombylide, 
with a note on Comastes, Os. Sac., v. Heterostylum, Macq. Notes on the Trichoptera in the collection of the Indian 
Museum. Diagnoses of new species and varieties of Freshwater Crabs. 1—3. Report on a small collection of | 
Lizards from Travancore. Descriptions of three new Cicindeline from Borneo. The relation between fertility and 
normality in Rats. Descriptions of a Barnacle of the genus Scalpellum from Malaysia. The Hemipterous family 
Polyctenidz. Notes on Freshwater Sponges, XI. Descriptions of two new shells from S. India. Preliminary note 
on a new genus of Phylactolematous Polyzoa. Miscellanea. 


Part IV.—Description of a minute Hymenopterous insect from Calcutta. The Insect Fauna of Tirhut, No. 1. Descrip- 
tions of new species of Botia and Nemachilus. New Oriental Sepsine. A new species of Fredericella from Indian 
lakes. Diagnoses of new species and varieties of freshwater crabs, No. 4. On some new or little-known Mygalo- 
morph spiders from the Oriental region and Australasia. 


Vol. IV, 1910. 


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

Nos. II and III.—The Thdian species of Papataci Fly (Phlebotomus). Taxonomic values in Culicide. 

No. IV.—Revision of the Oriental blood-sucking Muscide (Stomoxine, Philematomyia, Aust., and Pristirhynchdmyia, 
(gen. nov.). 

No. V.—A new arrangement of the Indian Anopheline. 

No. VI.—A revision of the species of Tabanus from the Oriental Region, including notes on species from surrounding 
countries, 

No. VII.—New Oriental Nemocera. Miscellanea :—Synonymy in Corethrinae, Indian Phlebotomi. 


Nos. VIII and 1X.—A revision of the Oriental species of the genera of the family Tabanidae other than Tabanus. Con- 
tributions to the fauna of Yunnan, Part VII. 


No. X.—Annotated Catalogue of Oriental Culicidae—Supplement. 


Vol. V, 1910. 


Part I.—The Hydroids of the Indian Museum, I. Notes on Freshwater Sponges, XII. Descriptions of new Shells in 
the collection of the Indian Museum from Burma, Siam and the Bay of Bengal. Materials for a revision of the 
Phylactolematous Polyzoa of India. Studies on the aquatic Oligochzta of the Punjab. An undescribed Burme 

= Frog allied to Rana tigrina. - Miscellanea. ‘ 

Part II,—Description d’Ophiures nouvelles provenant des derniéres campagnes de ‘‘ I’Investigator’’ dans 1’Océan 
Indien. Description d’ Holothuries nouvelles appartenant au Musée Indien. The races of Indiax rats, Il. 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 fish from India and Persia, with descriptions of new species. 


Part I1I.—A new genus of Psychodid Diptera from the Himalayas and Travancore. The Indian barnacles of the sub- 
genus Smilium, with remarks on the classification of the genus Scalpellum. On a subspecies of Scutigerella unguicu- 
Jata, Hansen, found in Calcutta. The distribution of the Oriental Scolopendride. Notes on Decapoda in the Indian 
Museum, I, Description of a new species of Nemachilus from Northern India. Notes on the larve of Toxorhynchites 
immisericors, Wik. Description of a South India frog allied to Rana corrugata of Ceylon. Contributions to the 
fauna of Yunnan, Introduction and Part I. Miscellanea. 


Part [V.—Notes and descriptions of Indian Microlepidoptera. On some. aquatic oligochete worms commensal in 
Spongilla carteri. On Bothrioneurum iris, Beddard. Notes on nudibranchs from the Indian Museum. On the 
classification of the Potamonide (Telphusidae). Catalogue of the pheasants, pea fowl, jungle fowl and spur fowl 
in the Indian Museum. On certain species of Palaemon trom South India. A//uaudella himalayensis, a new species 
of degenerate (.) cockroach, with an account of the venation found in the genera Caydax and Alluaudella. 


Rhynchota 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. Contributions to the fauna of Yunnan, Parts II to V. Pedipalpi in the Indian 
Museum, I and II. Six new species of shells from Bengal and Madras. Mziscellanea :—Fleas from India and 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 Vermetidae. Aquatic animals from Tibet, 
Ill. Cyprinidae from Tibet and the Chumbi Valley, with a description of a new species. of Gymnocypris. New 
species and varieties of Crustacea Stomatopoda in the Indian Museum. The development of some Indian Ascala- 
phidae and Myrmeleonidae. Miscellanea :—The occurrence of Dactylopius citri, Risso, in the Himalayas. Note on 
Aquatic Rhynchota. 


Part I[I.—Nouveaux Chironomides de 1’Indian Museum de Caleutta. 


Part IV.—Indian Isopods. Systematic notes on the Ctenostomatous Polyzoa 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 J6/a. 


Part V.—Revision of the Oriental Tipulidae. Five new Indian species of Clavicorn Coleoptera. Tes Chironomides 
(Tendipedidae) de 1’Himalaya et d’Assam, The occurrence of dpus 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 Museum, 
Ill. Fauna of Paresnath Hill. Observations on the shallow-water Fauna of the Bay of Bengal made on the 
‘Golden Crown,’’ 1908-09. Coccidae in the Indian Museum, I. On the nervous system of Ampuillaria globosa, 
Description of a species of Phoridae that causes Myasis in man. Miscellanea :—Mimicry of a Mutiliid by a spider. 
Capture of Limulus on the surface. Development of the larva of Lingula, Distribution of some Indian and 
Burmese Lizards. 


Part II.—The Mosquitoes of Calcutta. Freshwater sponges, XIV. Pedipalpi in the Indian Museum collection, III and 
IV. New Freshwater Crab from S. India. Decapoda in the Indian Museum, 1V, Fauna Symbiotica Indica, [ 
and II. Inverteb:ate Fauna oi the Kumaon Lakes. Fauna Symbiotica Indica, IMJ. The Indian Mud-Turtles. 
Anatomy of Afopos sanguinolenta. Cestodes from Indian Fishes. Miscelianea:—Entoprocta in Indian Waters. 
Rearing of Leeches in Bara Banki District. Habits of Tiger-Beetles. Schizodactylus mons:rosus as bait for birds. 
Macrones menoda vat. trachacanthus. Aquatic Tortoises of the Ganges and Brahmaputra. 

Part III,—Gordiens du Musée Indien. A new Tortoise from South India, New species of Branchiodrilus and other 
aquatic Oligochaeta, with remarks on cephalization in the Naididae. Fauna Symbiotica Indica, IV. Freshwater 
Medusa from the Bombay Presidency. A new species of Indian Thrips. Aquatic Chelonia of the Mahanaddi. 
Recent Crinoids from the Indian Ocean. Fauna of Yunnan, VIII, Earthworms. Asiatic Naiades in the Indian 
Museum. Miscellanea:—Malatia Mortality in the Fringe Area of Calcutta. Larval Habits of Toxorhynchites 
immisericors. East Asiatic species of Apus. 

Part IV.—Notes on the surface-living Copepoda of the Bay of Bengal. Freshwater sponges of the Malabar Zone __ Notes 
on the Habits and Distribution of Limnocnida indica. The Anatomy of melo indicus. Two new species of Scolo- 
pendridae. 


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


An Aceount of the 
ORIENTAL PASSALIDAE (COLEOPTERA). 


ERRATA. 
« 99 
P. 319 line 1, for “ Zaenioers” read “ Taentocerus 


. Lh 
FP 331 line 21, for “Puritulus” read “ Auritulus 


By 


F. H. GRAVELY, MSc. 


Assistant Superintendent, Indian Museum 


el eee, oS 

: | evetsers 

ae ~ ne, ; 
me) Cla 

‘(eD. .- 1 he 


Te 
, we eat 


AN ACCOUNT OF THE ORIENTAL PASSALIDAE (COLEOPTERA), 
BASED PRIMARILY ON THE COLLECTION IN THE INDIAN MUSEUM ~ 


By F. H. Gravety, VW.Sc., Assistant Superintendent, Indian Museum. 


(Plates XI—XIII.) 


CONTENTS. 


Page 
1. Introduction .. oy) ‘ : a 2: fal 77, 
2. External anatomy with special Peicreriee to taxonomic values oc a 370) 
3. Classification of Indo-Australian Passalidae ye es So.) 4 OE 
4. Catalogue of specimens in the collection of the Indian Mises with notes on those 
in certain other collections ae or ne S80 eZ On! 
5. The variable species : BS a4 a8 3 en e202 
6. Geographical distribution and synonymy : as 3d ag AOR 
7. Appendix I. A revised classification of the Acenaiaae Be ae se ESO 
8. 35 II. Keys for the determination of species Be j or 318 
9. », Ill. The genus Targuinius and aremarkable new genus from Wee Guinea . 326 
Io. 5 IV. Supplementary catalogue of specimens in the Indian Museum collection 330 
II. Summary and conclusions .. eee ‘fe =e rae a 336 
12. Bibliography .. hs BY ae ss ae S40 
13. Index as an a ef 0 oC sone S45 


1. INTRODUCTION. 


The present Memoir is in effect 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 ‘‘ List’’ 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 Kast would 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- 


178 Memoirs of the Indian Museum. (VoL... 


tinctive than that system at first sight suggests. I have accordingly given a 
revised classification with keys to all the genera known from this area, except Tar- 
quinius, Kuwert. This genus is as yet only known from one imperfectly described 
specimen from New Guinea which I 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 attempted more than I can successfully carry out in the limited state of 
my personal 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 my 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 brackets, 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. Bainbrigge Fletcher and Captain R. B. Seymour Sewell, who have collected 
valuable series of South Indian species for me; to Mr. S. W. Kemp, who, with the 
assistance of Captain the Hon. M. de Courey 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 supposed 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 
the 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 by Zang from New Guinea without any description (1906b, p. 23). ‘These show that Kuwert’s 
figure of this species is less trustworthy than his description. In general appearance they closely 
resemble species of the genus Leptaulax, from which they differ chiefly in the absence of scars from 
the mentum and in the presence of six well-developed lamellae on each antenna. See also Appendix 
III, pp. 326-330 below. 

* The locality of Semicyclus redtenbacheri, Stoliczka (1873) is probably not Ceylon but Brazil (see 
Kuwert, 1898, pp. 203 and 279); while that of Paxilloides schmidtii and philippinensis, Kuwert (Deutsche 
Ent. Zeitschr., 1890) is probably also Brazil, not the Philippine Islands (see Kuwert, 1891, p. 182 and 
1898, p. 181). Consequently the genera Semicyclus and Paxilloides are not regarded here as belonging to 
the Oriental Kegion. Nor have I accepted the record of Mastochilus politus from Madras (see Stoliczka, 
1873, p. 156). 

* Since this paper was sent to press I have had access to the collections in Hamburg, Berlin, and 


1914.] F. H. GRAVELY: An Account of the Oriental Passalidae. 179 


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 te 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. 


2. EXTERNAL ANATOMY WITH SPECIAL REFERENCE TO THE 
TAXONOMIC VALUES OF DIFFERENT PARTS. 


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 (1g05a, 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. Kraepelin, Prof. Von Brunn, Herr Gebieu, Dr. Horn and 
Mr. Arrow for the help they gave me when studying these collections. Ihave also, through the kindness 
of Mr. René 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 named by 
Kuwert; to complete my account of the widely-distributed Oriental genus Aceraius 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. 


180 Memoirs of the Indian Museum. [Vor. III, 


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 
constancy 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 Ceracupes, 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. 

Symmetry. 

Oriental Passalids may be either symmetrical or asymmetrical. The degree of 
asymmetry 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. 


-Intennae. 


The antennae are always ten-jointed. The basal joint is always long and thick. 
Joints 2—4 are thinner and only about aslong as broad ; the fourth occasionally bears 
a rudimentary lamella. Joints 5-7 always 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 isin the least degree curled (see text-figure 1, A and 
B), as it almost invariably is at this point in dried specimens. The three terminal 
joints 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 grandis subsp. hirsutus 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 distinguisting the species of Macrolinus one from another, and in separating 
the genus Kaupiolus from the genus Labienus; and I have used the same character 
in défining the Gonafas 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 absolutely constant and they are difficult to compare in the case of insects which 


19I4.| F. H. Gravety: An Account of the Oriental Passalidae. 181 


‘TEX!-FIGURE I. 


Fic. A. Leptaulax dentatus, from above. 
es LB. PP from below. 
‘3 C. Left mandible of Aceraius grandis, subsp. hirsutus, viewed obliquely from the inner side. 
Pe D. Anterior margin of head of Leptaulax dentatus, from below. 
5 E. 3 = aA Macrolinus nicobaricus,  ., 
,, EF. Front coxae and prothorax of Leptaulax dentatus, viewed obliquely from in front. 


,, G. Front coxae and prothorax of Comacupes cavicornis, vat. laevicornis, viewed obliquely 
from in front. 


aa. anterior angle of head; a.z.a. anterior intermediate area of metasternum; a./.é. anterior lower 
tooth: an.z. basal joint of antenna; a. prs. anterior plate of prosternum ; D.e/. base of elytron ; c. coxa; ¢.d. 
central area of metasternum; c.a.;. anterior part of hind coxa; cam. canthus ; c.p.3. posterior part of 
hind coxa; c.f. central tubercle; ¢. elytron; f femur; /r.a. frontal area; fr.v. frontal ridge; g./. fourth 
groove of elytron; i.¢. inner tubercle; /.a. lateral area of metasternum; /by. labrum; /. p. fused lateral 
plates of lower surface of prothorax; 1.é t. lowest terminal tooth; m. mentum ; m.l.¢. middle lower tooth ; 
ms.em mesothoracic -epimeron ; 77/s.es. mesothoracic episternum ; mss. mesosternum ; mt.em. metatho- 
racic epimeron ; mdé.es. metathoracic episternun ; m.t.t. middle terminal tooth: o.f. outer tubercle; p.2.a. 
posterior intermediate area of metasternum ; /././. posterior lower tooth; p.prs. posterior plate of proster- 
num; ~.r. parietal ridge; prn. pronotum; s. scar: sc. scutellum; s.0c. supra-occipital ridge; s.ov. supra- 
orbital ridge; swf. suture; ¢. tibia; fv. trochanter; ¢.f. terminal teeth ; #./. upper tooth ; w././. upper- 
most terminal tooth; v.f. ventral tubercle; J-VZ abdominal sterna. 


182 Memoirs of the Indian Museum. [Vo1,. III, 


differ from one another in size or in the extent to which their antennae happen to be 
unfurled. It is so difficult, 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. 


Manatbles. 


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 asa 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 1) may be described as follows:— 

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

Three terminal 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 question belongs.' It is set further back in 
the latter case than in the former. 

Three Jower teeth. Of these the anterior one varies considerably in form in 
different genera; the middle one is always a chisel-shaped lamella which is hinged 
on to the mandible except in certain 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 sui face of head. 

The principal features of the upper surface of the head are shown in text-figure 
1A. 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 ridges, 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 obsolete 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-family Aulacocyciinae, and it is often convenient to refer to this as the first section of the 
family, as opposed to the remaining subfamilies which together constitute the second section. ‘These 
subfamilies are considered in the present paper to be four in number, and I have termed them Pleur- 
ariinae, Aceraiinae, Gnaphalocneminae and Leptaulacinae. See below, p. 191 and onwards. 


1914. | F. H. GRAvELY: An Account of the Oriental Passalidae. 183 


the angle at which the upper and anterior surfaces of the supra-orbital ridges meet is 
moderately 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 Cylindrocaulus 
and Auritulus 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. 20), and fig. 28 with 
fig 28a). The part of the anterior margin of the head from which this process is 
developed may be called the anterior angle of each side of the head. The size of this 
angle is often of some taxonomic importance. From it a canthus 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 Caulifer. Some- 
times, however, even in the Aulacocyclinae, the inward prolongations of the posterior 
ends of the supra-orbital ridges evice 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-oceipital 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 Auritulus, 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 tubercle. 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 Dara eee “he supra- pecial ridge unites the supra-orbital ridges, and is also con- 
tinued outwards beyond them. 


184 Memoirs of the Indian Museum. fVor. Tae 


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 fusion, 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 examples, fig. 19, and figs 52 and 
52b,c and d). 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 /rons 
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 c/ypeus, or plate 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 plates together before going 
on to enquire into this. 

The pair of tubercles in which the frontal ridges end may be called the inner 
tubercles, 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 appear, 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 different shape from those which they replace. 


* Before this can be done the whole of the labrum must be removed. 


IQT4. | F. H. Gravety: 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. 1, D and EK). 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. 1 D and E, and figs. 36a, 37a, 38a, 39a, 40a 
and 41a}. 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 reality to what is here considered to be the frons instead 
of to theclypeus. 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 Leftaulax 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—7.e. 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 Gnaphalocnemis affords the most important exception I know. 


186 Memoirs of the Indian Museum. [Vor THs 


The /abrum, or upper lip, is attached to the margin of the clypeus, 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 Gnaphalocnemis 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. Incertain species of Aulacocyclinae e.g. Ceracupes fronticornis and C. austen?) 
this ridge instead of being indistinct is very pronounced. 


Lower Surface 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 concavity 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. The pair 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. 


Pyrothorax. 


The whole upper surface of the prothorax is covered by the pronotum. 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 readily 


' The scars on the mentum of these genera were unfortunately confused with primary scars in my 


1912 preliminary paper. 


1914. ] F. H. GRavELY: An Account of the Oriental Passalidae. 187 


produced by friction however, that but little reliance 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 iittle 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 frosternum, 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, in front of and behind the sockets (coal 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, or 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. 


Mesothorax. 

The scutellum, or median dorsal plate of the mesothorax, 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 


| 


188 Memoirs of the Indian Museum. [Vou. III, 


importance, are the mesothoracic 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 mesothoracic epimera are small plates which do not appear to be of any 
taxonomic importance. 

The mesosternum 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. If 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 to be as fully divided from the 
median part as they are from the mesosternum and perhaps represent the trochan- 
tine 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 L-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 
piece behind this patch is usually found to be more or less completely fused to the 


1914. ] F. H. Gravety: An Account of the Oriental Passalidae. 189 


metasternum. ‘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 family; but I have not found it possible to utilize 
it for differentiating species. 

Metathorax. 

The metathoracic episterna and epimera do not appear to present any charac- 
ters of taxonomic importance; but the metasternum, 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 Tvbertoides, 
Episphenus 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 srarely 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 /ateral 
aveas. ‘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, which 
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- 
cocylinae than in the other subfamilies. 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 


190 Memoirs of the Indian Museum, [Vou. III, 


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) 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 puncturing, 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 Comacupes) 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 bea useful confirmatory generic character, and its extent, 
when present, a confirmatory specific one. 


Legs. 


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 subfamilies 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 
Aulacocyclinae than in the other subfamilies (compare text-figures 1 F and 1G}. 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 occasionally important, at least in 
the genus Taentocerus. 

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 Aulacocyclinae there is often, about 
two-thirds of the way down the outer side of the ¢7bia 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 


Iol4. | F. H. GRAVELY: An Account of the Oriental Passalidae. IQI 


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 Aceraius. 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 genera, Aceraius and Trichostigmus, from their respective allies. 


3. THE CLASSIFICATION OF INDO-AUSTRALIAN PASSALIDAE. 


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 
Aunitulus and Cylindrocaulus, whose affinities with the Aulacocycline genus Ceracupes 
have been established by Arrow (1907, p. 446). Kuwert, who overlooked the 
only known species of the genus Awritulus, defined the Aulacocyclinae only accord- 
ing to the apparent structure of the prosternum 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 frout 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 made here to 
Sharp and Muir’s works on ‘‘ The Comparative Anatomy of the Male Genital Tube in Coleoptera’’ 
(Trans. Ent. Soc. London, r9r2, pp. 477-642, pl. xlii-lxxviii, Passalidae, pp. 579-580, pl. xliv, 
figs. 11-13a). These investigators found that in representatives of the genus Azlacocyclus, 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), Labienus aud Protomocoelus (Gnaphalocneminae), Leptaulax (Leptaula- 
cinae), and of the American genera Proculus and Neleus ‘‘the tezgmen consists of two distinct pieces, 
the basal piece and the lateral lobes.’ 


? 


1g2 Memoirs of the Indian Museum. (Vou. THe 


Part. Section I. | Section IT. 

Mandibles. Plane of the lowest terminal tooth vertical, Plane of the lowest terminal tooth 
the tooth itself not situated well behind the horizontal the tooth itself situa- 
two upper ones, directed forwards when ted well behind the two upper 
the mandibles are open. ones, directed inwards when the 


mandibles are open. 


Upper surface of | Parietal and frontal ridges, and the inner and Parietal and frontal ridges, and the 


head. outer tubercles with the ridges associated inner and outer tubercles with 
with them, all absent. the ridges associated with them, 

present 
Prothorax and its Middle part of prosternum with distinct median | Middle part of prosternum flat or 
appendages. keel which is usually hidden behind by the with median groove or less dis- 
coxae, as the distal ends of these project tinct keel, the whole surface 
vertically from their cavities, their vertical between the coxae exposed, as 


inner faces touching one another in almost the coxae do not project suffi- 
all species. Posterior plate of prosternum ciently from their cavities to meet 
more or less pointed behind. above it. Posterior plate of pros- 

ternum 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. 


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 Cylindrocaulus) 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 single subfamily Aulacocyclinae, 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 Tvistorthus, Taeniocerus, Comacupes 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 
Caultfer 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 .s 2. 
r.¢ Central tubercle very strongly developed, its apex fused with 


anterior margin of head; upper tooth of both mandibles very long 


and slender; middle lower tooth moveable .. Bs .. Ceracupes, Kaup; pp. 212 
& 277. 


1914.] F. H. GRAvELY: An Account of the Oriental Passalidae. 193 


longations of supra-orbital ridges absent ; pronotum not drawn out 
into a bifid protuberance in front; middle lower tooth of mandible 


(? always) immoveable 0 iG a 
Central tubercle absent, head smooth and concave; supra-orbital 


Central tubercle present; supra-orbital tubercles and anterior pro- 
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 ferana 
(es tubercle aig De AG 56 : Bs 4. 
3. Posterior ends of supra-orbital ridges curved fawards to form 
ie a median tooth situated immediately behind central tubercle ; 
central tubercle elongated dorsally ; mesothorax smooth .. [Carlifer, Kaup.] 
Mentum with strong median keel; mesosternum strongly punctured ; 
[ction sterna with at least a few hair-bearing punctures .. Comacupes,' Kaup; pp. 
4: 204 & 267. 
Mentum not keeled; mesosternum at most feebly punctured; abdo- 
minal sterna unpunctured a0 6 = an bic Ke 5. 
a Central tubercle unidentate or tridentate apoE a, .. [Tristorthus,' Kuwert.] 
al Central tubercle bidentate above a a3 oc 6. 
Central tubercle never pedunculate, often without any posterior face, 
anterior face never longer than dorsal, anterior end less highly elevated Taeniocerus, 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 a8 .. Aulacocyclus, Kaup; pp. 
' 211 & 272. 


Front coxae almost contiguous; canthus extending about half way 


across the eye; supra-orbital tubercles flattened, expanded at the 


apex, truncate Auritulus, Zang, p. 279. 


ie Front coxae widely separated; canthus eeitddine all the way across 
Cylindrocaulus, Fairmaire; 


Dp: 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 


the eye; supra-orbital tubercles slender and pointed 


} tubercles of head always simple .. Bc . 2. 
\Mentum with primary scars, except when these are replaced bs 
more or less strongly developed secondary scars*; outer tubercles of 

3. 


head often complex 


! Since preparing this key I have examined the type of Comacupes minor, Heller, and specimens of 
C. foveicollis, Kuw., from Borneo. ‘These have proved to be, in several important characters, transi- 
cera between the genera Comacupes and Tristorthus (see below, p. 267). 

2 I know of no really sharp distinction between these genera. The definition given here involves 
the transference of Taeniocerus deyrollei, and with it I presume T. masterst (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. rvosenbergit more closely than they do any species of Tweniocerus. 

3 This distinction taken by itself is not altogether a satisfactory one, for in one race of Ophrygon- 


194 Memoirs of the Indian Museum. [Vou. III, 


Only three antennal lamellae recognizable when antenna is furled; 


ridge joining inner tubercles absent re te .. PLRURARIINAE; pp. 213 
ai & 270. 
More than three antennal lamellae recognizable when antenna is 
furled; ridge joining inner tubercles present she .. ACERAIINAE; pp. 215 & 
280. 
Supra-orbital and supra-occipital ridges discontinuous .. .. MACROLININAE; pp. 240 
4 & 293. 
Supra-orbita! and supra-occipital ridges continuous! .. Br: ae 2 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 Ae .. GNAPHALOCNEMINAE; pp. 
246 & 297. 
4. Inner tubercles situated between outer tubercles on anterior margin 
Vee head, the middle part of which is formed by the ridge joining 
them: only three antennal lamellae recognizable when antenna is 
furled ne Ac ate a Se .. LEPTAULACINAE; pp. 251 
& 302. 


The first of these subfamilies, the Pleurariinae, contains only one Oriental 
genus, Pleurarius. This genus has been grouped by Kuwert with the genera Ninoides, 
Pertinacides and Epipertinax, 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 
opinion. 

The next three subfamilies comprise between them the Macrolininae of 
Kuwert, together with all the groups which fall under the second number “‘9a”’ 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 cantori 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, however, quite 
unlike that of the secondary scars of any species of Gnaphalocneminae ; and the form 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 Analaches, 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 by the first 
two (see below pp. 213-5). The mentum of the genus Hyperplesthenus, 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 metasternum are, however, fused asin all other genera of the 
Hyperplesthenus group, a fusion which is not known to occur in ary genus of Aceraiinae. See also 
appendix III, p. 326, below. 

' See also p. 183, footnote. 

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


1914. | F. H. GRAVELY: An Account of the Oriental Passalidae. 195 


tum and have the supra-orbital ridges continuous with the supra-occipital ridge. The 
Tiberius here referred to is T. kuwerti, Arrow ( =cancrus, auct.) and I have had to 
create a new genus 7 berioides for the reception of this species, of Chilomazus borealis, 
Arrow, and of a new species (7. austent) 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 ga’’ groups, it is clear that of the grooves and 
depressions found in the mentum of different genera of his Lachinae, 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 them 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 (1g05a, 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—will probably prove to be a mistake. FE. filius, Kuwert, from the 


1 Kuwert omits the scars in his figure of the type of mentum found in this genus. They are men- 
tioned, however, in Percheron’s description of Passalus sexdentatus (= Mastochilus polyphvllus according 
to Kuwert); they are well developed in the specimens in the Hamburg Museum determined by Zang 
M. polyphyilus ; and Kuwert himself mentions in his descriptions of both species of the geaus 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 Mastachilus, see Zang, 1903), p. 418. 
2 Concerning Kuwert’s confusion of the names Analaches and Epilaches, see below p. 283. 


196 Memoirs of the Indian Museum. [ Vou. III, 


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 yet been described. Consequently the genus EF pilaches is not included 
in the following key. The genus Heterochilus, for which Zang (1905), p. 154) has 
suggested the new name Rhipsaspis, does not appear to me to be sufficiently distinct 
for recognition, especially in view of Zang’s notes upon it (1905a, p. 167). The genus 
Ophrygonius, 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*:— 


Inner tubercles separated by a space 4-4 as long as that separating 


outer tubercles ss xc =i se .. Tuiberioides, Gravely ; pp. 
Li 215 & 280. 

Inner tubetcles separated by a space 3-1 times as long as that 

separating outer tubercles a8 : 2 3 2. 

Anterior margin of head symmetrical or bat slightly adymmisteiban 


[toe tight and left outer tubercles simply pointed # ac ot Vee: 
_ Anterior margin of head markedly asymmetrical, left external 
tubercle longer than right, rounded or more or less obliquely truncate 
iets ane AD - a ae ee 50 4. 
Central area of mentum smooth, Seite Be nor depressed near 
J of anterior margin; anterior margin of head quite symme- 


trical xt Sh se 3 as .. Episphenus, Kaup; pp. 
217 & 28r. 

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 be oi mits .. Chilomazus, Zang;,pp. 218 
& 281. 

Frontal ridges represented by fine 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 5. 

I‘rontal ridges neither represented nor accompanied by grooves ; 

antennae with only four pubescent lamellae; lateral grooves of elytra 

broad, their punctures transversely linear .. a .. Ophrygonius, Zang; pp 
227 & 285. 


1 I have since been able to confirm these two suggestions by reference to the type of E. infantilis 
and cotypes of F filius; see below, p. 283. 

2 For a modification of the views expressed 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 order in which the species would have been described. 


1914. | F. H. GraveLy: An Account of the Oriental Passalidae. 197 


Sides of elytra hairless, unpunctured except in the grooves .. Basilianus, Kaup;. pp. 
220 & 282. 
5., Sides of elytra clothed with hair, lateral ridges more or less exten- 
sively punctured especially near the shoulder ade he .. Aceraius, 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 distiaction 
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 allit 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 :—(1) Macrolinus wrus and diuvenbodei from Celebes ; 
(2) M. sikkimensis, nicobaricus, and andamanensis ; (3) the Ceylon forms; and 
(4) the Malaysian and Philippine forms with MW. swlciperfectus trom 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 ga’’ groups of Kuwert’s classification, 7.c. all except the genera 
Chilomazus (=Laches), Epilaches, Heterochilus, Bastlianus, and Acerawus. It is most 
abundantly represented in the Far Eastern part of the Indo-Australian 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 
represented. ' 

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 allied, and that the former occupies the same position in the fauna of the 
Australian 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 J have also, through the kindness of Prof. Lampert, been enabled to examine the type specimen 
of the genus Hyperplesthenus, Kuwert, which is preserved in the Stuttgart Museum. Without reference 
to this specimen I could not have determined the characteristics of the genus. 


198 Memoirs of the Indian Musewm. (Vor. Tig; 
mandibles. 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, forma 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 


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

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 ‘‘ Kaupiolus Group,’’ which includes all genera so characterised, does not 


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


IgT4.] F. H. GrAvety: An Account of the Oriental Passalidae. 199 


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 
places.' 

When thus arranged the genera fall into six groups (each enclosed, in the diagram, 
by a dotted line), 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 peculiar 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 
similar.’ 

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 Pyotomocoelus 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 .. a ne ot a 
Primary scars never present; surface of head without deep or 


iS) 


sharply defined anterior depressions, or dentition of right mandible 
more reduced than that of left .. 50 ere oe alg . 9. 


! I follow Zang (1905), p. 227) in supposing the lateral and intermediate areas of the metasternum 
to be fused in the genus Pelopides ; but Zang himself had not seen a specimen (1995, p. 316). See 
also following footnote. 

2 It is, for instance, very possible that Protomocoelus and Pelopides have sprung more directly thar 
is here indicated from Kaupiolides, which they resemble in the structure of the metasternum more tha ! 
they do Analaches-Cetejus. 

8 Perhaps always; but in the most highly speciaiized forms they are often very small and obscure. 
In all the more highly specialized forms, and also in some less 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 Gnaphalocnemis, 


200 Memoirs of the Indian Museum. [Vou. III, 


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 fused; surface of head usually 
rough ; dentition always complete Se -» (Kaupioloides Group) 3. 
Outer tubercles at most as far from one another as fe anterior 
angles of head; often asymmetrical, always so when lateral and iuter- 
mediate areas of metasternum are fused; surface of head smoother, 


tN 
Ta 


polished ; dentition complete or reduced... a .. (Protomocoelus Group) 6. 
( Anterior intermediate and lateral areas of metasternum distinct 
a from one another... 36 : AG se Pca 


\Aptetios intermediate and lateral areas of rnetaseeraies hia) .. [Kaupioloides, Gravely *). 
Secondary scars ieebly developed (sometimes absent), transverse, 
never extending back as much as half way to posterior margin of 


mentum .. te : 5 BP .. [Episbhenoides, Kuwert *]. 
Secondary scars distinct, yaiteble ice eS 


ing in an acute angle 56 Ar “5 - .. [Mastochilus, Kaup]. 
Secondary scars represented by a single long transverse crescentic 

or semicircular groove, crossing middle-line about halfway between 

anterior and posterior margins of mentum .. =e .. [Pharochilus, Kuwert §]. 


Anterior intermediate and lateral areas of metasternum distinct 
from one another*; dentition complete, normal a 
| Antericr intermediate and lateral areas of metasternum fused ; 
6.4 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 
tooth hs ae ar a0 sh oe ae ot anos 


ie 


| Secondary scars represented by a pair of very small grooves, meet- 


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 Protomocoelus 
group can easily be recognized by the form of the anterior margin of the head (see Kuwert, 1896, pl. 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 
Kauptoloides seems to be almost exactly intermediate between the two groups in these 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 Hyperplesthenus group; in this 
respect the genus Kawpiolotdes 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 Profomocoelus group. 

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

* Concerning the genera Episphenoides and Pharochilus see also Heller, 1910, p. 17. ‘The general 
shape of the scars of the former 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 Mastochilus, rather than 
of that characteristic of the genus Episphenoides. 


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


1914. | F. H. GRAVELY: An Account of the Oriental Passalidae. 201 


straight or lightly curved so as to contain not less than 120°; ridge 
joining inner tubercles usually straight and not strongly crested ; 
pronotum usually very strongly punctured in anterior angles; 
posterior angles of lower side of prothorax usually not very hairy.. [Cctejus, 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 


Upper margin of left mandible, between upper tooth and base, 
very hairy os! 50 a6 a6 -. [Analaches, 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 C . [Pelopides, Kuwert]. 
Left outer tubercle much longer than right; a ridge speonaide Be 
hind deep excavation from left inner tubercle towards middle of 


supra-orbital ridge .. “it a0 4s .. [Protomocoelus, Zang (= 
Pelops, Kaup)]. 


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 oe . ae ae LO: 
All six lamellae of antennae long and rather siemens mandibles 
sometimes normal, more often with anterior lower teed absent or 
fused at base with lowest terminal tooth, especially on left side ; 
secondary scars semi-circular or Y-shaped, more or less transverse, 
often more or less distinctly continuous with one another across 


‘E of antennae usually all rather short and stout, never more 


middle line {3 ers a oe .. (Gonatas Group) 17. 

Lateral and intermediate areas of metasternum fused ; mandibles 

normal on both sides .. as aye 5c .. (Hyperplesthenus Group) 
TI. 

) Lateral and intermediate areas of metasternum distinct ; anterior 

lower tooth of right mandible smaller than that of left, usually 

absent .. ae Se a - = 3: a LAR 


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 Protomocoelus; 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 Analaches or of Protomocoelus ; but which cannot be deter- 
mined before the mandibles and metasternum are described after examination and not, as in the above 
key, by inference. So TI prefer to retain the name for the present. Zang (1905), p. 227) who had also 
not seen a specimen (1905, 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. gravidus, Kuwert, from 
Mindanao, ought probably to be transferred to some genus of the Gnaphalocnemts group on account of 
its toothed labrum. 


202 Memoirs of the Indian Museum. (Vou. IIT, 


f Central tubercle of head free, directed forwards os .. (Aurelius, Kuwert']. 
= Central tubercle of head normal ee ae ei ae ave eg tee 
Secondary scars widely separated 3 He re 3 


Secondary scars not widely separated, directed enards so that 
their posterior ends nearly meet in middle-line without reaching 
I2., 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 A uh .. [Hyperplesthenus, Ku- 
; wert *}. 


markedly asymmetrical *; pronotal scars hairless ie .. [Kaupiolus, Zang* 


bec normal; outer tubercles variable, simple or complex, not 
(=Vellejus, Kaup)]. 


13% 
z Antennae with lamellae of last four joints abnormally long and 


slender ; left outer tubercle broadly truncate, right more or less 
pointed but no longer than left; pronotal scars hairy .. .. ([Labienus, Kaup]. 


| 

Fe tubercles simple, very variable, acute or truncate, highly 
asymmetrical (that of the right side being always the larger) or 
almost (? sometimes quite) symmetrical Bx or .. [Plesthenus, Kaup (= Em- 
bryulcus, Zang *)]. 

At least one of the outer tubercles complex, or truncate with an ad- 

ditional tubercle between itself and anterior angle of same side of 

head AS - ae BG es .. (Gnaphalocnemis Group) 
15. 

w-shaped depression much as in Omegarius; anterior lower tooth 

of right mandible small but distinct a a .. Parapelopides, Zang ; 


pp. 246 & 297. 


Secondary scars united in middle-line behind, together forming an 
Secondary scars completely separated as a rule, anterior lower tooth 


of right mandible absent a a an Bc oe Son aOR 
Outer tubercles equal ® ae Be at .. Trapezochilus, Zang; 
pp. 247 & 297. 
16. Outer tubercles unequal, that of right side always larger than that 
of left oe +6 de es 54 .. Gnaphalocnemis,' Heller ; 
pp. 248 & 208. 


| The genus Aurelius, specimens of which I have myself examined, possesses all the characteristics 
of the Hyperplesthenus group. 

2 The genus Hyperplesthenus 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, 
Kuwert, without any separate generic definition. An examination of the type specimen has enabled 
me to define the genus more precisely, and to find that Labienus gracilis, Heller (Abh. Mus. Dresden 
xiii, 3, p. 16), of which I have also seen the type, is really a Hyperplesthenus. See Mitt. Naturhist. 
Mus. Hamburg XXX, 1913, p. 105. 

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

* Kaupiolus trigonophorus, Zang, as Zang himself suspected, is not a true Kawpiolus, and I have 
made it the type of a new genus Kawpiolotdes (see above, p. 200). 

6 Zang considers the name Plesthenus to be preoccupied by Plesthenes, Stal. 

’ Not always quite symmetrical in structure however; so it is possible that the genus Trapezochilus 
will eventually have to be merged in Gnaphalocnemis. 

1 Kuwert (1896, p. 229) states that there is always a depression between the left outer tubercle 


r9r4.] F. H. GRAVELY: An Account of the Oriental Passalidae. 203 


Secondary scars curved oid a SH | AKG) 
Z i Secondary scars Y-shaped, forming Peres a W- ced ee 
sion ais oe ea AE ..  Gonatas, Kaup, pp. 250 
| & 299. 
Secondary scars approximate, forming together an w shaped de- 
18.¢ ( pression 510 . 6 Be 36 .. [Omegarius, Kuwert!.] 
secondary scars iticip separated fs ae .. [Tatius, Kuwert!.] 


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 (1905¢, p. 106) has split it up intotwo. 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 asa 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 Leptaulav as restricted by Zang, and those belonging 
to the last-mentioned apparently belong to his new genus Leptaulacides ; 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. Ihave, therefore, recogiized the genera as distinct, only when 
describing the species before me,” and have dealt with both under the old name Leptaulax 
(as defined by Kaup and Kuwert) in the sabsequent synonymmic revision of the 
Oriental Passalidae as a whole. The three accepted genera of Leptaulacinae may be 
distinguished as follows : — 

Sie of elytra hairless ie if cab BS ae ae 2. 
Sides of elytra hairy e as ; Trichostigmus, Kaup, pp. 
\ 261 & 310. 


and sapere ridge in this genus. ‘his character is not found in any other accepted genus of the 
group but it is shared by ‘‘ Pelopides’’ gravidus, Kuwert, which must be transferred, in all prob- 
ability, to this group (see Zang, I905a, p. 316 and 1905), p. 227). ‘‘ Pelopides’’ gravidus 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 sometimes found on opposite side of a single speci- 
men. The two genera have therfore been united in the following pages. 


204 Memotrs of the Indian Museum. [Vor. III, 


- 


( Parietal ridges of head short and laterally truncate, being separated 
from supra-orbital ridge by a deep hollow; insects strongly and 
| more extensively punctured = ae 5c .. Leptaulax, Kaup, pp. 251 
a4 & 302. 
Parietal ridges long, laterally continuous with supra-orbital ridges ; 
insects sparsely and less extensively punctured te .. Leptaulacides, Zang. 


4. LIST OF SPECIMENS IN THE INDIAN MUSEUM COI,LECTION, WITH 
NOTES ON THOSE IN CERTAIN OTHER COLLECTIONS.’* 

Localities are recorded as briefly as possible in this list. For further geographical 
information 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 Ephisphenus 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. 


Subfamily AULACOCYCLINAE. 
Genus COMACUPES, Kaup. 
Comacupes cavicornis, Kaup. 
1. C. CAVICORNIs, Kaup,s. str. 
Regd. No. SS SS" Sinkep Island Moti Ram. 
Description.—Length 23-25 mm. Labrum about twice as wide as long, anterior 
margin slightly concave, angles rounded, sides slightly convergent behind. Upper 
tooth of 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 preceding page. 
* A list of these collections will be found above, p. 178. 
This has since been confirmed; see p. 252. 


1914.] F. H. GRAVELY: An Account of the Oriental Passalidae. 205 


of which it is a little 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. Pronotum 
without strong punctures except in the marginal groove, which is not interrupted 
before ot 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. 
Mesothoracic 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 covae, 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, fn. var. 
/ 


Pl. xi, figs. 11a: 
Regd. No. "oe" 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. cavicormis, s. sty,, 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. sty.. and that found in the following variety. 


3. C. CAVICORNIS, var. LAEVICORNIS, Kaup. 
PE xl, figs. 2—-2a. 


9478 
Regd. No. = Malacca ? 
9244 9410-5 9417-20 
1 1 1 Seat 
MBS 9430 9432.3 9437-8 Wontar { Stoliczka bequest, 
»? Sintie &.4 Gee iy pie € 
5 u : \ J. Wood-Mason, ete. 
9445-5 9480 6381-2 
1 Ly qd 


9440-2 


1 ? J. Wood-Mason. 


206 Memoirs of the Indian Museum. [Vou. IIT, 


Description.—Length 23-27 mm. Differs from C. cavicornis, s. sty., only in having 
the upper surface of the /ead 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 me/asternum more sparsely and less 
extensively punctured behind. 


Comacupes stoliczkae, n. sp. 
Pl. xi, figs. 3-34. 


Regd. No." 3 (type) Johore, Malay Peninsula J. Wood-Mason, ete. 


8785 


ss a Sinkep Island Moti Ram. 


This species is intermediate between C. cavicornis and C. masoni; but it closely 
resembles C. cylindraceus 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- 
vaceus 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. cylundraceus 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 Taentocerus 
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. E/ytra sometimes with 
some small punctures along the lower side of the anterior half of the tenth rib; 
otherwise as in C. cavicornis. 


1914. ] F. H. GRAVELY: An Account of the Oriental Passalidae. 207 


Comacupes masoni, Stoliczka. 


Pl. xi, figs 4-4a. 
Regd. No. “ (ty pe') Johore, Malay Peninsula J. Wood-Mason. 


Description.—Length 30 mm. Labrumas 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. Pyvonotum with anterior angles a 
little more prominent, sides slightly divergent behind; enlarged areas of anterior 
marginal groove rather abruptly terminated at their outer ends. Prothrovax ventrally 
as in C. stoliczkae. Mesothorax, metasternum, 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). 


Pl. x1, figs. 5-6a; text-fig. 2. 


(377 6384 AS . 
Regd. No. 5; 47 Sinkep Island Moti Ram. 
: 
2611 ag 
ai Klingkang, Borneo Sarawak Museum. 


ss 5 

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. cormocerus; and the series of specimens before me shows 

ea the species to be so variable that I am inclined to think it 
possible that C. masoni, Stoliczka, may ultimately have to 

Hew eeees roti be referred to it also. . 

C. cylindraceus, from Description.—Length 24-28 mm. Differs from C. stoliczkae 
the left side x 4. and C. masont 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 pl. xi, figs. 5a and 6a, and text- 


TEXT FIGUER 2. 


: : ; : ae 9449 
' 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. 


208 Memoirs of the Indian Museum. [Vou. IIT, 


fig. 2'. The form of the anterior parts of the marginal groove of the pronotum 
and the puncturing of the abdominal sterna are both extremely variable. 


Genus TAENIOCERUS, Kaup. 
Taeniocerus bicanthatus (Percheron). 


Pl. xi, figs. 7-70. 
Regd. No. “¥ ? J. Wood-Mason. 
9477 6362-5 6385 
¥A, 1 lt lt 

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. Mentwm 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 go°, 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. Pronotum smooth, 
polished ; anterior margin almost straight, anterior angles rectangular, pointed but 
not prominent ; sides parallel, posterior angles much rounded, posterior margin some- 
what convex; marginal groove punctured, incomplete in the middle-line in front, 


Johore, Malay Peninsula ? 


' 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 probably from Johore. So Zang’s suggestion (195a, p. 105) that Bornean specimens are 
distinguished hy a broad horn such as is figured by Kaup, and Malayan ones by a longer horn, can no 
longer be maintained. 


1914. | 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 obliquely 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 pygmezus, Kaup. 
Pl. xi, figs. 88a. 
Regd. No. ee ‘Malacca ”’ U 


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

Description.—Length 15-16 mm. Labrum 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 mentum 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 about 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. Pyronotwm polished, anterior margin 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-line 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 


210 Memoirs of the Indian Museum. (Vor, Ts 


behind this patch; mesothoracic 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 any such tooth. Lateral grooves of e/ytra distinctly punctured. 


Taeniocerus bicuspis, Kaup. 


Pl. xi, figs. g-ga. 


Regd. No. =“ - > Sikki Stoliczka bequest, Asiatic 
gus 875 esha Society, ete. 
+ 
8129 Gopaldhara, Rungbong Valley, Dar- H. Stevens (coll. W. K. 
za 9 jeeling District Webb). 
_ _ Chongtang, Darjeeling District J. Wood-Mason. 


» “ue Upper Tista, 4000 ft. F 


i — Sureil, Darjeeling District A. Alcock. 
ud aa N. E. Frontier G 
a oe Dafla Expedition 
x oe Dafla, Camp 9 . H. Godwin-Austen. 
= oot Sas Dikrang Valley | 
+ oe Harmutti, base of Dafla Hills 
Be mas a Kobo, Abor Country, 400 ft. 
sal Rotung, Abor Country, 1400 ft. Noy. 1911—Feb. 1912. 
vs ees Upper Rotung, Abor Country, 2000 ft. S. W. Kemp. 
sree Upper Renging, Abor Country 
4 = ae Cachar J. Wood-Mason. 


Mr. Stevens’ collection includes a specimen from Silonibari at the base of the 
North Lakhimpur Hills in Assam. 

Description.—Length 20-22 mm. Labrum asin T. bicanthatus. 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 consequently 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-orbital 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 


1914. ] F. H. GRavELY: An Account of the Oriental Passalidae. 211 


in an angle of go° 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 go° (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. Scwtellum without (or 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 behind ; 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 
smooth. 

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


Genus AULACOCYCLUS, Kaup. 


Aulacocyclus andrewesi, n. sp. 
Pl. xi, figs 10—10a. 

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_I-shaped structure with a minute denticle on the connecting ridge. Mentum 
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 


212 Memoirs of the Indian Museum. [Vor,. III, 


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 
orizontal part when seen from the side, but only about two-thirds as thick when 
seen from above, extremity concave. Pvonotum 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; mesothoractc 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 legsarmed with a small spine 
on the outer side about two-thirds down, those of the third pair with this spine 
obsolete. 


Genus CERACUPES, Kaup. 
Ceracupes fronticornis (Westwood). 
Pl sa 5 fie 22. 


Regd. No. “_ Gopaldhara, Rungbong Valley. Darjeeling District. H. Stevens (coll. W. K. Webb). 


Description.—Length 22°5 mm. Differs from C. austent 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 go’. 


Ceracupes austeni, Stoliczka. 
Pl. xi, figs 11—-I1a. 


6358 Sb7 
l4 Is 
6356-7 SS 


lt Is 


Regd. No. Datla, Camp 9 


Dikrang Valley 


” 


BBL (Ly pe) | Naga Hills H. H. Godwin-Austen. 
>> 14 é 


6359-60 


» T Manipur 


Description.—Length 20-25 mm. Labrum 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 


W4s2 


| ‘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 polished; 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 §-shaped in profile, being convex (and usually very obtusely 
angular) behind and concave in front, long narrow postero-dorsal surface granular, 
antero-ventral surface flat and very strongly granular except immediately below the tip 
which is smooth. Supra-orbital ridges tooth-like, apical angle 60°-go°, 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 go°. 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 
ot indistinctly punctured. T7biae of middle legs with I-3 spines on the outer side 
before the apex, those of the hind legs with at most one such spine. All furrows 
of elytva distinctly punctured, ribs unpunctured. 


Subfamily PLEURARIINAE. 
Genus PLEURARIUS, Kaup. 
Pleurarius brachyphyllus, Stoliczka. 
Pl. xi, figs. 13-134. 


9317 6385 Beddome. 


Reg. No. ie (paratypes) ! \ S. India Stoliczka. 
a ? Madras Museum. 


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


214 Memoirs of the Indian Museum. [Vor. IIT, 


Regd. No, Puddutottam,  <Ana- 
malais, 4200 ft. | 
1900 Ry sas / In rotten wood. 
” iy Anamalais, 4000 ft. .. » T. Bainbrigge Fletcher. 
us Anamalais, a. 5500 ft. 
be 108 Kalyana, Pandal, Anamalais, 3000 ft. 
7 est (paratype) ' Nilgiri Hills Madras Museum. 
Fe “ Kulattupuzha, W. base of W. Ghats, | 


mh N. Annandale. 
Travancore fh 


This species is represented in the collection of the Madras Museum by two 
specimens from the timber forest, Cochin State; and in that of the Bombay Natural 
History Society by a specimen from Palur presented by G. Lund. 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 355-45 mm. Labrum 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 fead 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 9o°; 
parietal ridges broad, extending straight outwards or slightly forwards. Pvronotwm 
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. Stoliczka and Mr. J. Wood-Mason’’), *\* 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. we which was presented by Stoliczka. 
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 oni, And No. ae 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. 


Ior4. | F. H. GRAVELY: An Account of the Oriental Passalidae. 215 


scattered punctures; mesothoracic episterna punctured above and along anterior 
margin; mesosternum without scars, finely and more or less extensively striato- 
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. Flytra hairless, innermost groove of each almost entirely smooth, 
remaining grooves strongly punctured. 

Habits —Dr. Annandale informs me that this species is probably not gregarious. 
Mr. Bainbrigge Fletcher sent me specimens apparently obtained together with 
colonies of Basilianus, 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). 
Pip fie 7: 


9103 9302-8 


Regd. No. | a Penae *¢< Sikkim 7? Stoliczka bequest, J. Wood 
( Fre aeh Wen ow Mason, ete. 
- wm Peshoke, Darjeeling District, a. 3000 ft. EF. Hannyngton. 
op — Manipur H. H. Godwin-Austen. 


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 mentum with slight (often almost 
imperceptible) hollow in middle; lateral parts punctured evenly all over asa rule. 
Upper surface of jead 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 


216 Memoirs of the Indian Museum, [Vou. III, 


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 plates of under side of prothorax as in 
preceding species. Mesothorax polished above, but dull below; scutellum smooth; 
episternum with normal oblique band of punctures. Mesosternum with or without 
very indistinct scars. Mctasternuwm 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. 


Tiberioides austeni, n. sp. 
Pl. xi, figs. 15-15a. 


Regd. No. [2 SS (type) Dafla Expedition H. H. Godwin-Austen. 


Bs “ Dikrang Valley ! H. H. Godwin-Austen. 
i: “et Kalek, Abor Country, 3800 ft., 29-xii-1r S. W. 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 brachyphyllus ; 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.”’ 


2 This prominence may be indistinctly marked off by a vague groove behind (No. ue shows this best 


among our specimens) and such specimens form a natural transition to the aberrant “* Chilomazus”’ 
borealis, Arrow (1907, pp. 467-8), from the Naga Hills. 


1914. | 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. 
Episphenus moorei, Kaup. 
Pl. xi, fig. 16—16a. 

Regd. No. “@ 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. ea 
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. Wandzbles 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-orbital 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. Pyvonotwm 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 Scutellum 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.”’ 


218 Memotrs of the Indian Museum. (Vor. III, 


roughened scars; surface of mesosternum bordering these scars on inner side very 
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. 
Piexi e077: 

Regd. No. ~ (type) Bulutota,! Ceylon (May 1907) Colombo Museum. 

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

Description.—Length 315-33 mm. Differs from E. mooret 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. 
Pl. xi, figs. 18-194. 


1. KE. comprToni, Kuwert, s. str. 
Regd. No. “= Hakgala,? Cent. Proy., Ceylon E. E. Green. 


7s 
5307-8 


a Ceylon British Museurn. 


” 


The Bombay Natural History Society’s collection includes a specimen from 
Haputale, Ceylon. 

Description.—Length 39-42 mm. Anterior margin of /abrum 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 


| 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 Pelmadulla Road. 10 miles beyond the 
Rakwana Rest House in Kolonna Korale, Ratnapura District, Province of Sabaragamuava. The 
altitude is not recorded, but is I believe about 4oo0 ft.’’ 

2 Mr. Green informs me that the altitude of the Hakgala Botanical Gardens is 5500 ft. 


1914. | F. H. Gravety: An Account of the Oriental Passalidae. 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. T,eft 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 limb 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 asin Episphenus moore, 
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 tule 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 of abdo- 
minal sterna almost smooth. Grooves of elytra normal, punctured ; shoulders slightly 
hairy. 
2. KE. COMPTONI var. FLACHI (Kuwert). 


Regd. No. me (13 spirit specimens ). | Pavipalla yacovoie 7 Cent: | 


1918-14 3737-38 


td 19 19 


? Prov., Ceylon, in rotten N. Annandale. 
(pinned). if 


wood (13-15-x-I9I11) 
Patipolla, a. 6000 ft., Cent. 
| Prov., Ceylon; in rotten 


9479-80 9483 

— —— x F. H. Gravely. 

18-18 | wood (first week in July, ee: 
IgI0) 

é oe 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. *® 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 
live in pairs; but, as they are gregarious, satisfactory proof of this is not very easily 


220 Memoirs of the Indian Museum. [Vor. III, 


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. COMPTONT, 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 EL. comptoni var. flachi from Patipolla. 
The other two are very unequal in size, and the larger of them is much smaller 
than the smallest specimen of the series from Patipolla. Until a larger series is 
available it is impossible to be certain how these Bulutota 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 Episphenus 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, Stoliczka. 
Pl. xi, figs. 20-20b. 


Regd. No. -_ “ ae (all paratypes) S. India. ; F. Stoliczka, ete. 
fe — Trichinopoly Father Honoré. 
- peo N. Kanara H. 1. Andrewes?. 
y s1e7-170 2 ; Mifviris i Director, Forest School, Dehra 
{ Dun; H. L. Andrewes 
af 16TeS Ootacamund Col. Beddome. 


Anamalais, S. India, ca. 5000\ nee 
, 'T!’. Bainbrigge Fletcher. 
ft., 21-i-12 f 


Tp ES 100 105 1987 Anamalais, 4000 ft., 23-i-12 
‘ SU } 2 SY ee 


, “ Anamalais, ca. 5000 ft., 21-i-12 
7 _ Anamalais, ca. 5500 ft. , 21-i-12 ge aie age 
oe met “= | Puddutottam, Anamalais, 
} ‘Od 1044 1985 1080-50 (4200 ft., 22-i-12 
» ae ws Kalyana, Pandal, 3000 ft., 
ae T. Bainbrigge Fletcher, 
“i: me Monica Estate, Anamalais, 
hoea ft eaters hey Bainbrigge Fletcher. 
i ae Bababudin Hills, Mysore, oe Aten i 
Agoo-s00o fia cae Tee I. Bainbrigge Fletcher. 
a _ High Range, ts eae ree 
May, 1891 : 
' Collected by T. R. Bell. * This has since been confirmed; see below, p. 28r. 


* Adults and larvae in spirit. 


1914. | F. H. GRAVELY: An Account of 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 119 specimens in drawing up the following description of 
this variable species. 

Description.—Length 25-41 mm. Labrum hairy, about as long as broad, sides 
parallel or slightly convergent behind, angles rounded, the left usually somewhat 
natrower and more prominent than the right, anterior margin concave. Mandibles 
with anterior margin of upper tooth (which tcoth 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 
oiten 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. Mentwm 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. Left outer tubercle directed inwazds, outer margin 
straight 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 over 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. Pvonotum 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 Memoirs of the Indian Musewm. [Vo1. III, 


sively punctured 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 Episphenus comptont; posterior area of prosternum very rarely bearing long 
erect hairs. Scwtellum 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. 
Mesothoracic episterna polished and punctured above and in front, finely roughened 
across posterior angles. Surface of mesosternum 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 punctured, especially in small specimens. All grooves 
of elytra of normal width; lateral grooves much more strongly punctured than dorsal 
ones. 

Habits.—Mr. Bainbrigge Fletcher writes of the series of this and the following 
species that he collected in the Anaimalai 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 differences 
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 you 
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 Pleurarius brachyphyllus, the 
logs were usually soft and quite rotten.”’ 


Episphenus neelgherriensis (Percheron). 
Pl. xi, figs. 21-214. 
Regd No, 15:6 9879-0680 & Tein ( Col. Beddome, 


16 18 


British Museum. 


iS) 
Oo 


1914. ] F. H. GRavELY: An Account of the Oriental Passalidae. 2 


5501-8! . . 
Regd. No. 45> Bababudin Hills P ee. 
Ns T. Bainbrigge Fletcher. 
Rs sai Coorg 
a mee Malabar 2 
ae High Range, Travancore Trivandrum Museum. 


We 18 
3460-70 
9 “19 


Coonoor, ca. 6000 ft. (Sims Park, 
| Jungle, and Stillbrook Garden) 

3 ‘ere Ootacamund Col. Beddome. 

: Ootacamund, 7500 ft. and 7800 ft., 


— 


sp. B. Seymour Sewell. 


6500! 6669 7500 
2? yb Ww 4 


‘’. Bainbrigge Fletcher. 

\ December, 1912 

-Anamalais (Kalyana, Pandal, 3000 

| ft.; Paralai Estate, 3600 ft.; Monica 

see / Estate, 4ooo ft.; Puddutottam, 4200 /T. Bainbrigge Fletcher. 
{ 


2? at) 
| 't.; ca. 5500 f{t., in rotten wood; all 

January 1912) 

9681 ? ? 


2? 1S 

Description.—Length 24-29 mm. Differs from specimens of FE. indicus 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 flat, wider in front than behind, not sharply defined ; anterior 
angles of head of about 120°, never prominent ; apical angle of supra-orbital ridges 
more pronounced ; /ateral plates of lower side of prothorax somewhat smoother and 
more highly polished towards the front; posterior plate of prosternum usually hairy, 
often punctured. Puncturing of sclfellum variable in extent, but on a somewhat 
different plan from that usually found in £. indicus, the strongest punctures being 
found in the posterior half of the plate which is often punctured almost all over 
metasternum less markedly hairy than is usual in EF. indicus. Posterior part of hind 
coxae smoother. 

Habits.—Mr. T. Bainbrigge Fletcher tells me that ‘‘ Most [specimens found at 
Ootacamund in December, 1912] 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. 


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


224 Memotrs of the Indian Museum, (Vor. IIT, 


Genus OPHRYGONIUS, Zang. 
Ophrygonius cantori (Percheron), s. str. 
I. O, CANTORI (Percheron), s. str. 
Pl. xi, figs. 22-22¢. 
Regd. No. ae Lambatach, Jaunsar Forest Research Institute. 


9105-9111 9118 9115-7 


( Stoliczka bequest, Asiatic 


iterate et “Sikkim”? ¢ 
“4 (ren ee) | Society’s collection, ete. 
‘A “ws Tista, Darjeeling District E. P. Stebbing. 
3 we Sureil, Darjeeling District A. Alcock. 
1” me = Kurseong, Darjeeling District 1 a oe 
- oe Gopaldhara, Rungbong Valley, H. Stevens (coll. W. K. 
Darjeeling District Webb). 

en ~<— Dikrang Valley, Dafla Hills H. H. Godwin-Austen. 

Gre ey eee ? E. I. Company’s Museum, 


ae Se 8 lt t 
G&G, 


I have also seen specimens from the British Museum collection from 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 from 
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 Episphenus 
indicus of similar size and E. neelgherriensis except in the following particulars. 
Anterior margin of /abrum 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 Episphenus 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 less 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; supra-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. Pyvonotum almost invariably punctured and hairy to some extent in the 
posterior angles and on the sides, frothorax beneath much as in Episphenus neelgher- 
riensis. Scutellum entirely smooth, hairless, unpunctured, and highly polished with 
exception of usual patch in middle of anterior margin ; mesothoracic episterna polished 
throughout, punctured above and close to anterior margin; mesosternum 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 
Metasternum 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. 


ae 


Regd. No, “S* Khasi Hills, Assam H. H. Godwin-Austen. 
se ov Teriaghat, Khasi Hills H. H. Godwin-Austen. 
9651 ? ? 


ae 1s 
In the British Museum collection there are two specimens from the Ruby 


Mines District of Burma. 

Description.—Length 27-29 mm., otherwise as in O. cantori,s. str. ‘The difference 
in size is an extremely small one, but appears to be constant in specimens from the 
Khasi Hillsand Burma. One of the specimens from the Ruby Mines has the mentum 
sculptured 7lmost as in the following sub-species. 


3. O, CANTORI, subsp. DUNSIRIENSIS, n. var. 
Rid fers2o5 
Regd. No. “@ (type) we Dunsiri Valley, Assam H. H. Godwin-Austen. 


18 ls 


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 Indian Museum. (Vou. III, 


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 O. cantori in the following characters 
only. Lamellae of antennae 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 ead very slender, almost parallel-sided, simply 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-FIGURE 3. 


A. Ophrygonius birmanicus, head x 4. 
B. Ophrygonius singapurae, head x 4. 


Ophrygonius singapurae, 0. sp. 
Text-fig. 3B. 


Described from a single specimen from Singapore in the collection of the 
K6nigliches 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 imaequa- 
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. wallaceit. The lowest terminal tooth 
of the mandibles 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 Acervaius, 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 O. inaequalis; the left outer tubercle 
is intermediate in form between that of O. cantori and O. birmanicus ; the parietal 
ridges are somewhat short; the supra-occipital ridge is united to the supraorbital 


tor4. | F. H. Gravety: 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 
occuts. The sides of the pronotum are closely punctured below the scars throughout 
their whole length; the Jateral plates of the ventral side of the prothorax are some- 
what rougher and more punctured in front than in O. birmanicus, which this species 
resembles in all other respects. 


Ophrygonius inaequalis (Burmeister). 


Pl. xii, fig. 24-242. 


: 120 «9122-4 9311-3 
1 1 1 
Regd. No. er Malacca Stoliczka bequest. 
9519 3750 
herp akc 
96, j 
i un Perak Moti Ram. 
p44. 
e a Johore ? 
6439-40 9672-6 9861 eee ak 
Pe ee aad Sinkep Island Moti Ram. 


Description.—Length 24-27 mm. Tamellae 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 natrowed 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 Memotrs of the Indian Museum. | [Vor..nue 


marginal groove incomplete across the middle both before and behind, its anterior 
extremities merged in a pair of somewhat triangular strongly punctured areas, 
behind each of 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 /ateral plates of lower side of prothorax somewhat spatser 
than in O. cantor: ; 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. Metasternum 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 
thickly 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 e/ytra wide, with more 
or less transversely linear punctures ', dorsal grooves normal; shoulders slightly hairy. 


Genus ACERAIUS, Kaup. 
Aceraius wallacei (Kuwert). 


Pl. xii, figs. 26-260. 


Regd. No. a 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. Labrum 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 mentum usually with sunken 
semi-circular area along anterior margin and three or four large punctures behind, 


' See also below, p. 285, footnote. 


1914. | F. H. GRAVELY: An Account of the Oriental Passalidae. 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 asin A. borneanus except that the pronotum is less punctured at the sides while 
its median grooveis obsolete, and that the mesosternum is polished, throughout or 
nearly so, except in the long narrow scars. Metasternum also as in A. borneanus 
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. borneanus. 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.—Length 38°5-42 mm. Labrum 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, mandibles 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; e/ytra as in following. 


Aceraius moschleri Kuwert. 


Pl. xii, fig. 33. 


283623 


Regd. No. “4, Sinkep Island Moti Ram. 


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

Description.—Length 36-42 mm. Differs 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 4. perakensis. 


| 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 Memoirs of the Indian Museum. [Vor IIT, 


Aceraius illegalis, Kuwert, s. str. 
Pl. xii, figs. 32-32a. 


Regd. No. “ ae Johore J. Meldrum; ? (purchased). 


Ihave also examined other specimens, including one from Kuching and one from 
between Ulu Madahit and Limbang which are preserved in the Sarawak Museum. 

Description.—Length 39-43 mm. Labrum as in A. wallaceit. amellae of 
antennae somewhat short and thick, first one pilose only round margin of anterior 
face. Mandibles as in A. wallace’. 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 A. wallacet. 
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 (Illiger). 


Pl. xii, figs. 27-274. 


OIBR-71 eng. 
Regd. No, =" == Johore J. Wood-Mason, ete. 
us Sees S0N6-50 Sinkep Island Moti Ram. 
a aan Kuching, Sarawak, 
Borneo Sarawak Museum. 
326: ° 
3 a Limbong, Borneo 
ae ? ? 


” 19 


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

Description.—Length 29-34 mm. T,amella of fifth joint of antennae long and pilose, 
the fourth joint sometimes furnished with a short lamella devoid of pilosity. Labrum 
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. Mentwm 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. 


1914. | F. H. GrAvELY : An Account of the Oriental Passalidae. 231 


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. wal/acei; 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 scufellwm asin A. wallacet; mesothoracic 
episterna with a patch of fine hair-carrying punctures in posterior angles as in A. illegals ; 
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. 


Aceraius grandis (Burmeister). 
I. A. GRANDIS subsp. HIRSUTUS, Kuwert. 


Pl. xii, figs. 28-29. 


Regd. No, “2*8 %6# .. Darjeeling - Stoliczka bequest, 
J. Wood-Mason. 
9259-64 S545 | Dr. J. Anderson, 
af ae ba j sim . T. Atkinson. 
Me L,. Mandelli. 
99 = waa Bhutan ? 
» - Harmutti, Base of Dafla Hills 
” eo Dikrang Valley : 
Anat yeh ae H. H. Godwin-Austen. 
» = SE Dunsiri Valley 
a a Duffla Expedition 
set N.-E. Frontier ? 


! hat from Serai in the Sarawak Museum Collection. 


232 Memoirs of the Indian Museum. [Vor III, 


Regd. No. = Kobo, Abor Country, 400 ft. 
% mu. Janakmukh, Abor Country, 600 ft. 
u 0 feo specimens) Rotung, Abor Country, 1400 ft. + SIs Kemp! 
2 = au Upper Rotung, Abor Country, 2000 
ft. i 
f ses Kalek, Abor Country, 3800 ft. 
E sate Cachar J. Wood-Mason. 
29 9151 9133 9249-51 : 
28 3 , : "2 “ eee A. Stoliezka bequest, 
» | a2 ues 3288-01 J * 8 S. E. Peal, Moti Ram. 
Wi? 670) 704 709 | . 
i } page hal rl "’ ) Khasi Hills Shillong Museum. 
Laid aoe 
2 Eres “h .. Tavoy Pie Ste -- Museum Collector. 
Apri Between ‘Thingannyinaun and 
lu a. ni mae ¥ =o 4 8 F. H. Gravely. 
: \. Sukli, Dawna Hills, 900-2100 ft. 
fs —, .. Sukli, Dawna Hills, c. 2100 ft. .. FE. H. Gravely. 


Mr. Steven’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. Lakhimpur 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 um, and Cambodia. 

Description.—Length 33-49 mm. Labrum as in A. leavicollis. Tamellae of 
antennae much as in 4. leavicollis in small specimens,’ often proportionally shorter and 
thicker in large ones. Mandibles also muchas in A. 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. Mentumas in A. leavicollis. Anterior angles 
of head much more prominent than in A. deavicollis 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 asin 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. /eavicollis, 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 
found. 


* In one specimen from Sikkim (No. =) 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. 


1914. | F.-H. GRAVELY: An Account of the Oriental Passalidae. 233 


in large specimens. Prothorax and mesothorax as in A. laevicollis, but patch of fine 
punctures in posterior angles of mesothoracic 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 elytva 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, 
Deli, Tengger Mountain, Bankalan, Bandjermasin, and N. Borneo. 

Description.—Length 39-52 mm. ‘his 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 elytya 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.” 
Pits ses sO: 

The Sarawak Museum collection includes one specimen from Sarawak; and the 
Berlin 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. hirsutus; 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. 


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

2 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. 


234 Memotrs of the Indian Museum. [Vo1,. III, 


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


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

Description.—Length 48 mm. ‘his species differs from A. grandis var. rectidens 
only in the presence of a stout upwardly directed tubercle on the canthus, just in 
front of the eye. 


TEXT-FIGURE 4. 
A. Aceraius occulidens, head x 4. 
B. Aceraius tricornis, head x 4. 
C.  Aceraius alutaceosternus, left mandible from the side x 4. 
D. Aceraius minor, head x 4. 
FE. Aceraius aequidens, head x 4. 


Aceraius laniger, Zang. 


Two specimens of this species, both from Mt. Kina Balu, c. 5000 ft., are pre- 
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 inclined to be less so; ridge separating lateral and intermediate 
areas of metasternum obtuse, rough ; tenth rib of e/ytra closely punctured and hairy 
at shouldets 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. 


1914. | F. H. GRAVELY: An Account of the Oriental Passalidae. 22 


Ww 
On 


Aceraius kuwerti, Zang. 
Plexi fie. 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 part-of 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 A. Janiger and A. kuwerti chietly 
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; tight outer tubercle of head quite as well developed as in A. laniger, simply 
pointed in profile. Metasternum and elytra variable, resembling those of A. /aniger 
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 mesosternum are almost always distinct. 


Aceraius pilifer (Percheron). 
Pie xiiefionss. 
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 


* But for size and this characteristic the species might easily be confused with 4. perakensis. 


236 Memoirs of the Indian Musewm. [Vor. III, 


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 Antennae never very long and slender; right outer 
tubercle of head somewhat more truncate in profile than in /aevimargo in fresh 
specimens, the suture distinctly bent downwards; scars obsolete on mesosternum ; 
anterior part of tenth rib of e/ytva always punctured and hairy. 


A. alutaceosternus, Kuwert. 
Pl. xii, figs. 34-34a; text-fig. 4C, p. 234. 

Regd. No. = Penang F. Stoliczka ? 

This specimen has been labelled Aceraius emarginatus by Stoliczka, 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 Taiping 4000-5000 ft., in the collection 
of the Deutches Entomologisches Museum. 

Description.—Length 35°5-36°5 mm. Labrumasin A. laevicollis. Mandibles also 
as in that species, but with convexity behind left upper tooth exceptionally large. 
Lamella of fifth joint of antenna short and devoid of the close pubescence with which 
the five succeeding lamellae are clothed. Mentum 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 A. borneanus, but shorter 
than in A. wallacet, triangular as seen from above, imperfectly forked at apex in 
profile. Prothorax much asin A. wallace’. Mesothorax as in A. tllegalis 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. tllegalis. 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. 
Pl. xii, figs. 36-362. 


Regd. No, “5 2 Catiaen ( Stoliczka bequest, J. Wood- 
U Mason. 
et “ee (type) Dikrang Valley H H. Godwin- Austen. 
S262 «. . - . 
ay = §Sirpo River, Nr. Renging, 
19 = M. Cy. 
/ Abor Country ape onie: 
2 “ Naga Hills Robert, through J. Wood- 


Mason. 


' 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. 


1914. ] 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. Left 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. alutaceoslernus ; 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 A. wallacer, the latter 
perhaps a little more distinctly punctured laterally. Sides of etytra clothed with 
short hair, seventh rib usually more or less thickly, ninth always thickly punctured 
throughout, tenth with hair-bearing punctures in front. 


Aceraius assamensis, Kuwert. 
Pl. xii, figs. 37-374. 


Regd. No. pete Cherra Punji H. H. Godwin-Austen. 
33 ea Manipur » 
é “_ Assam British Museum. 
7070 7082 2859 > MacClelland, E. I. Com- 


23 32 eo? 19 


pany’s Museum, etc. 

Description.—Length 31-36 mm. Differs from dA. himalayensis in the following 
respects only: mentum sparsely or not at all puncturedin middle ; inner processof 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 iar from apex of lower portion as from right inner tubercle; scutellum 
as in A. grandis; posterior parts of hind coxae, and scars of abdominal sterna less 
smooth ; seventh rib of e/ytva with or without punctures at anterior or posterior end 
only. 

Aceraius tavoyanus, n. nom. 
Pl. xu, figs. 38-384. 

This species is undoubtedly the same as that from Burma, erroneously described 

by Kuwert in 1898 under the name fz/ifer, Percheron. 


Regd. No. * Sy 5 Tavoy Museum Collector. 


There is a specimen from Siam in the British Museum. 


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


236 Memotrs of the Indian Museum. [Vox. ITI, 


Description.—Length 30-33 mm. Differs from A. assamensis in the following 
characters only: left outer tubercle slender, its distal angles more produced; median 
groove of pronotum entirely obsolete as.in A. wallacet; posterior parts of hind coxae 
finely rugose; abdominal sterna as in A. hima/ayensis ; seventh rib of elytra punctured 
at posterior end only in the four specimens before me, as it was in Kuwert’s 
specimens also. 

Aceraius helferi, Kuwert. 


Pl. xii, figs. 39-394. 
Beer feri 
Regd. No. | so Pe ‘Se sian: J. Wood-Mason. 
ly 
.s Seu Upper Tenasserim J. Wood-Mason. 
2828-9 2831 
AN B Misty Holl 1 Sukli : 
» =S* (adult and lar- Been ate et ooo ora eee tage 
19 y 
\ - : ae Dawna Hills, 2100-2500 ft. 
vae 1n spirit) 
Peet ‘ . Between Thingannyinaung and ) : 
” 1” «1 i Sukli, Dawna Hills, goo-2too ft. $3 F. H. Gravely. 
3743 Rangoon Bombay Natural History 


Society. 

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

Description.—Length 31-40 mm. Differs from A. favoyanus in the following 
characters only: left outer tubercle, though not quite constant either in size or shape, 
normally 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. 
Pl. xii, figs. 25-250. 

Regd. No. = Borneo Sarawak Museum. 

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

! Present throughout in all the three Rangoon specimens, but none of the Tenasserim 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 Tenas- 
serim form, but eveu in specimens found within a few hundred yards of such typical ones they may be 
present at the posterior end exactly as in A. ¢avoyanus. 


TQI4. | F. H. Gravety: An Account of the Oriental Passalidae. 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 24-28 mm. Anterior margin of /abrum 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, 
defined 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 parallelsurfaces ; its counter- 
part on the right side minute. Mentwm 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, stich 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. Pronotum 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 coxae. Scutellum 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 rule not further for- 
wards; central area smooth and polished, as is also ridge separating lateral from ante- 
rior intermediate areas. Posterior parts of hind coxae smooth or nearly so. A bdomi- 
nal sterna indistinctly punctured in scars, the second with a few indistinct, but 
larger, hair-bearing punctures as well in fresh specimens. F/yiva hairy, closely punc- 
tured at sides over whole surface from seventh rib onwards. 


240 Memotrs of the Indian Museum. [Vor Ii 


Aceraius minor, n. sp. 
Text-fig. 4D, p. 234. 


Described from five specimens from Taiping, 4000-5000 ft., in the Kgl. Zool. 
Museum in Berlin. 

Description.—23-25°5 mm. Resembles A. borneanus 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 
sears ; 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 ; mentum 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 A. borneanus. 


Subfamily MACROLININAE. 
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 distinguishing one species from 
another. 


Ig14. | F. H. GRAVELY: An Account of the Oriental Passalidae. 241 


Macrolinus nicobaricus, n. sp. 
Pl. xiii, figs. 40o—40a. 
9092-9 9101 9277 
ile io ees 
9279-81 516-7 430-2 


1 6 ic 
Regd. No. ; pe rt &( 14 Nicobar Islands F. Stoliezka, 
| ie ype) 5 ‘| UF. A. de Roepstorff. 


9847-50 
18 
6981-2 


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 anyextent. 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 oblique 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. Mesosternum 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 polished with exception of a 


1 T find by looking up the ao that this Fearne is of doubtful authenticity; it appears to ae the 
only record of the species outside the Nicobars and is probably incorrect. 


242 Memoirs of the Indian Museum. [Vor IIT, 


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 sparsely or closely. lytra slightly hairy at the shoulders; very 
strongly punctured in the lateral grooves, more weakly in the dorsal ones of which 
the innermost are scarcely punctured at all. 


Macrolinus andamanensis (Stoliczka). 
Pl. xili, figs. 41-41a. 


mses! (paratypes) 
vee? (paratypes) 
9165 
(paratype) 

Regd. No. ; Rr, ; ) Andaman Islands Stoliczka bequest, J. Wood- 
—— (paratypes) Mason, ete. 
4069-70 4072-4 3713-20 
ar) 5 ~~ 10 
6602-8 6428-9 9860 

i 14 1s 
= Moulmein ! Museum Collector. 


Alls appear to be paratypes. Nos. 2465 and 24678 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 frontalridges (which in this species 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 
very strongly marked median groove whose surface is finely roughened, but more 
usually this is obscure or absent; I have not seen it in B. nicobaricus, but it may 
very likely be found occasionally in that species also. 

' T find by looking up the register that this record is of doubtful authenticity, so as it appears to 
be the only record of the species outside the Andamans it is probably incorrect. 

2 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 difference 
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. 


IgI4. | F. H. GRAVELY: An Account of the Oriental Passalidae. 243 


Macrolinus sikkimensis (Stoliczka). 
I. M. SIKKIMENSIS, Stoliczka, s. str. 
Pl. xiii, figs. 42-42a. 


Regd. No. ° (type) ** Sikkim ’?! F. Stoliczka. 
692 706 6405-6 : : . ; 
i a Ge Sa Khasi Hills Shillong Museum. 
z “ Cachar J. Wood-Mason. 
es eo Harmutti, base of Dafla Hills 
eee Dikrang Valley \ 
i Biss Nie eenttontn ‘% Jn _H. H. Godwin-Austen. 
bs SS Se SS Dafla Expedition | 
9865-88 yey at 
‘ = Camp 9, Dafla Expedition 
ef sas-8 Kebo, Abor Country, 400 ft. | 
a S14 Janakmukh, Abor Country, 600 ft. S. W. Kemp. 
19 | Pp 
3142-5 
4s = Rotung, Abor Country, I400 ft. 
r» = Naga Hills Moti Ram. 


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 ridgesas in M. nicobaricus; 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 scarsalso. Mesosternum usually 
more polished 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. Metasternum 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 
grooves. 

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 favoyanus 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. 


“= (type) 


' 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. 


244 Memorrs of the Indian Museum. [Vousrit: 


2659 


1v 


Regd. No. Between Misty Hollow and Sukli, F.H Gravely. 


Dawna Hills, 2100-2500 ft. 
2645-58 


Between Thingannyinaung and Sukli, F. H. Gravely. 


Dawna Hills, goo-2100 ft. 
2660 


” 19 


Dawna Hills, Tenasserim F. H. Gravely. 


I have also seen specimens from Carin Cheba, 3000-3500 and 4000-5000 ft., in 
the collection of the Deutsches Entomologisches Museum. 

Description.—Length 29°5-33'5 mm. Differs from M. sikkimensis, s. str., 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. ®}4®) 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 gradual 
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 215 mm.' Differs from M. rotundifrons only in its smaller 
size, and in the extremely coarse puncturing of all the grooves of the e/yira, the 
dorsal grooves of M. crenatipennis being as coarsely punctured as the lateral ones of 
M. rotundifrons. 

Macrolinus rotundifrons (Kaup). 


Pl xi os Ag. 


Regd. No. “ Kandy, Ceylon } ESBS Eco 
N. Annandale. 

» “ee Peradeniya, Ceylon cee: 
F. H. Gravely. 
r ~ Ceylon F. H. Gravely. 


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

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 tidge between outer tubercles. Pyonotum 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: An Account of the Oriental Passalidae. 245 


Mesosternum 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). 
PAF ait eae 


Reed. 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). Pyonotum unpunctured but (? always) with 
amore or less obscure depression in place of the strongly punctured area behind 
anterior extremities of marginal groove. Scars of mesosternum 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 jead almost vertical to the general surface. In the former 
respect, and in its locality, it agrees with Kaup’s description of M. webert, 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 ? 


asi 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. Labrwm punctured and hairy, anterior 
margin straight, angles rounded. All six lamellae of antennae very 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. Mentwm 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 


246 Memoirs of the Indian Museum. [ Vox, Ait 


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. Pyonotum 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. Pyrothorax beneath punctured and hairy at the sides. Scutel- 
‘um smooth and polished ; mesothoracic episterna punctured above and along anterior 
margin ; mesosternum 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 
backwards 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. Metasternum 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. 


Subfamily GNAPHALOCNEMINAE. 
Genus PARAPELOPIDES, Zang. 
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. Differs 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 very obtuse, middle denticle rarely 
distinguishable. Only extreme lateral angles of scutellwm 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. Metasternum 
with at least a few punctures in the posterior intermediate areas. 


i) 
BI 
3G 


Igr4.| F. H. GRAVELY: An Account of the Oriental Passalidae. 


Genus TRAPEZOCHILUS, Zang. 
Trapezochilus nobilis (Kuwert)'. 
Pl. xiii, fig. 48; text-fig. 5, C-D 


Regd. No. “5 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 


Wl an = a ia 


) ae Sf) 


B. 
TEXT-FIGURE 5. 


A. Parapelopides symmetricus, anterior margin of head x : 

B. 5 right mandible x 4. 

Cy Trapezochilus nobilis, anterior margin of head x 4. 

Fe right mandible x 4. 

E. Trapez zochilus respectabilis, 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 qecumeninty: or absent. Lateral alates of lower surface 


; This species ee aeoved not to be distinct fein elie mete Bea ie names ae to them here 
must both be discarded in favour of dorsalis, Kaup (see below, pp. 297-8) 


248 Memoirs of the Indian Museum. (Vor. III, 


of prothorax smooth and hairless in front, punctured and very hairy further 
back, often rugose behind the coxae. Scwutellum 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, Sumatra, in the collection of the 
Deutsches Entomologisches Museum. 


( 


TEXT-FIG. 6. 
Gnaphalocnemts simplex, 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 metasternum 
sparsely punctured in one specimen. 


Genus GNAPHALOCNEMIS, Heller. 
Gnaphalocnemis simplex, n. sp. 
Text-fig. 6. 
Described from a single specimen from Perak in the British Museum collection. 


Description.—Length 31 mm. Head, with its appendages, as in G. monticulosus, 


' See foot-note on previous page. 


IQr4. | F. H. Gravety: 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 polished. 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. burmeistert. 


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

Described from specimens from Deli, Pedong, Peinan, Soerian Plantation (nr. 
Solok), Battak Mountain, Kepahiang, Redjang-Iebong, 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 
collection. 

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


Gnaphalocnemis monticulosus (Smith). 
Plo xii; figs 40a. 
I have not seen the specimen which Stoliczka says he received from Sumatra ; 


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. “ve 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 labrum slightly 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 tuberclescomplex, somewhat variable in form (compare 
figs. 4g and 4o9a'). Anterior border excavate between left outer tubercle and front of 


| 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 the Indian Museum. [Vou. IIT, 


left supra-orbital ridge. Anterior angles of head containing about 120°, canthus 
rounded externally ; supra-orbital ridge as in Tyvapezochilus 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 very coarsely but often sparsely punc- 
tured, not hairy. Hind coxae usually smooth. Second abdominal sternum usually 
punctured ‘n front of scars; abdominal sterna otherwise smooth. Elytrva with grooves 
5-7 more or less broad, their flattened polished surface usually more or less completely 
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.). 
Pl. xiii, figs. 50-5o0a. 

Regd No. me Java J. Wood-Mason. 

I have also seen one specimen from the island of Madura which is the property 
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 types in the Hamburg 
collection. 

Description.—Length 45-56 mm. Differs from the preceding in the following 
characters only: upper tooth on both sides often more obtuse, 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 coxvae 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 
punctures. 


Genus GONATAS, Kaup. 
Gonatas germari, Kaup. 
Pl. xiii, figs. 47-474. 
Regd. No. ** Java ; W.S. Atkinson. 
Description.—Length 26°5 mm. Labrum hairy and punctured, anterior margin 
with a deep semi-circular excavation, angles rounded and prominent, sides straight 
and parallel. Antennal lamellae long, sixinnumber. Left mandible somewhat larger 
than right; upper tooth small on both mandibles, rectangular on left, more obtuse 
on right ; two upper terminal teeth partly fused, lower one distinct, meeting them in 
an angle of about 60°; anterior lower tooth rudimentary on left, large but blunt on 


19f4. | F. H. GRAVELY : An Account of the Oriental Passalidae. 251 


right mandible. Mentwm 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 obliquely truncate as seen from the 
side; anterior angle of head containing about 100°; canthus almost equilaterally 
quadrangular. Pvonotum with strong median groove, marginal groove incomplete in 
middle in front, obsolete in middle behind. Pvothorax below polished; 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 ; mesosternum 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. 


Subfamily LERTAULACINAE. 
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. Frontal area of head shorter and broader. Pronotum and abdominal 
sterna more as in L. dentatus, the latter not extensively punctured. Posterior parts 
of hind coxae smooth. Sculpturing of lateral grooves of e/ytva 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. 
Pl. xiii, fig. 51. 


Regd. No, “= Ten miles south of Kuching, Sarawak C. W. Beebe. 


19 


_ 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. 


252 Memoirs of the Indian Museum. [Vor.EiT; 


dentatus, in the following characters only. Upper tooth of mandibles, usually weaker, 
anterior lower tooth simple. Inner tubercles of head perhaps a little more 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 metasterynum 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, of a small smooth spot near 
the posterior margin. Depressed surface of two or three outermost grooves of e/ytra 
dull, the sculpturing somewhat indistinct and worn-looking, especially at the ends ; 
the ninth and tenth ribs narrow throughout. 


Leptaulax dentatus (Fabr.). 
1; (Lk. DENTATUS, Fabr. 5. Ste: 
Pl. xiii, figs. 52-52d. 
Regd. No. 4° “2 s5ts Near Vizagapatam Museum Collector. 
7509-14 


ns ia La-ai Valley D. G. A. Pritchard and 
J. R. Waterfield. — 


surat Kobo, 400 f{t., Abor Country 
S sor Janakmukh, 600 ft., Abor Country 
3077-3103 32662 
*9 19 19 


Rotung, 1300-1400 [t., Abor Country 


s we Upper Rotung, 2000 ft., Abor Country » 5. W. Kemp. 
os mes Renging to Rotung 26co ft. Abor 
Country 
» po Kalek, 3800 ft., Abor Country 
” oe POT N.-E. Frontier ? 
& wen Sibsagar, Assam S. E. Peal and Museum 
Collector. 
» a2 Dafla Expedition 
x ase see > Burroi, base of Dafla Hills , 
622 962-3 : j H. H. Godwin-Austen. 
RET Harmutti, base of Dafla Hills 
$3 “ aad Dikrang Valley, Northern Frontier of - 
Assam 
9505 6421 Pee 
” : co Dunsiri Valley, Assam ( ? 
_—o (HH. H. Godwin-Austen. — 


Te oT Durrang, Assam 


! 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. 


1914. | F. H. GraveELy: An Account of the Oriental Passalidae. 253 


= ay 73 ) Stoliczka bequest, 
Regd. No ec ae ) Bhutan | Dr. Cameron. 
1 T ? 
9150-63 9165-7 9170-1 
1 1 a 
9204 9211 9219-20 
1 1 ey 
sf oe ee aes \ Sikkim Stoliczka bequest, etc. 
9375-7 9487-9 oo 
1 1 
9493 6407-9 
lai eal 
9383 ut 
= Darjeeling (Sikkim in register) F. Stoliczka. 
| 9169 a | ’ Stoliczka bequest, 
1 : . 4 , > 
zi ) Rises hes Assam < Asiatic Soc.’s collection, 
See me) J ( E. T. Atkinson, ?. 
ae a E. Cachar J. Wood-Mason. 
9137 9189-40 9142-9 
\\Saa ‘iigcettena aa Ve ( Stoliczka bequest, 
2” § 937: egu @) 
\ ore (ass US. Kurz. 
861-8711 32722 : 
19 Kawkareik, Tenasserim, c. 300 ft. F. H. Gravely. 
i ae Misty Hollow, W. side of Dawna Hills, F. H. Gravely. 
Tenasserim, c. 2200 ft. 
fe oe Between Misty Hollow aud Sukli, top of F. H. Gravely. 
Dawna Hills, 2100-2500 {t. 
i ws Between Thingannyinaung and Sukli, F. H. Gravely. 


E. side of Dawna Hills, 900-2100 ft. 


9913 872-96 895 . ; 
” a“ a 19 Tenasserim J. Wood-Mason. 
i 5875-6 5879-85 5887 \ 
at) 10 10 = ‘A 
+9 Bs80-92 807 S044 pUPper Tenasserim J. Wood-Mason. 
| 10 19 wennLO Nae 
7720-7726 737 
4 4 4 
7740 7748 7754 
4 4 4 
7838 7891 7897 a ; 
» Gh. care te Tavoy Museum Collector. 
7900 8102 8105-6 
4 ry 
90 8763-4 665 
5 5 


( Stoliczka bequest, 
« Dr. W. Abbay. 
( 2 

( Ve a Masen. 


9168 9381-2 9486 


os ee i Mergui 


9356- 515 960-1 . 
33 eee Johore, Malay Peninsula 
? Museum Collector. 
4491-5 4497 4499-4504 
”? 1 10 10 


2866 Java H. E. Andrewes. 


97 19 


Perak, Malay Peninsula Moti Ram. 


| Specimens selected to show the full range of continuous variation from a series of 47 specimens 


taken from a single colony. No. *” is from the same colony. 


2 Adults and larvae in spirit. 


254 Memoirs of the Indian Museum. ‘[ Vor. III, 


I have also seen specimens from Buxa (Duars) in Mr. Andrewes’ collection; one 
from Silonbari at the base of the Lakhimpur hills in Mr. Stevens’ collection ; and 
specimens in the Dehra Dun collection from Kochugarh, Goalpara District, 14-v-06; 
Kapti, Chittagong Hill Tracts; and Chaduar, Assam, g-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 19°5-37 mm. Labrum 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. 52)-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. Pyvonotum 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 
aud 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, 


1914. | 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 lives under the bark of logs which have 
rotted till their outer parts have lost their strength. 

2. lL. DENTATUS var. GLABRIVENTRIS, 0. var. 


9134 9340 9513-4 


1 1 1 
291) RO S144 y i k b me, .W 1- 
Regd. No. Be a arse Bek Teles Stoliczka bequest, J.Wooc 
3792-3 3727 \ Mason, etc. 
\ t0 10 : 
2867 
” 79 Java 
ee ? H. E. Andrewes. 
» aa: Borneo i 
ae Philippine Isles East India Company’s 


99 3 
Museuin. 


This variety is not a very distinct one, forms intermediate between it and 
the typical form being sometimes found. 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. IL. CYCLOTAENIUS, Kuwert, s. str. 
Pl xi, figs se 


Regd. No, “22 Johore J. Wood-Mason, ? 
A a“ Sinkep I. Moti Ram. 
as a Kuching 
2613 : Sarawak Museum. 
= Penrissen 


2D 19 
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; mesosternum finely roughened 


. 


256 Memotrs of the Indian Museum. [Vor 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. 


2. L. CYCLOTAENIUS subsp. HIMALAYAE, Kuwert. 


Regd. No. = Dikrang Valley 
_ H. H. Godwin-Austen. 


3747-3718 Dafla Expedition 


o 19 
3199-3207 
” Tn 


Renging, 2150 ft ee ee 
j le . Kemp. 


3195-8 
92 ty 


Description.—Length 15-17 mm. Differs from L. dentatus in the following respects 
only. Head and mandibles as in L. humerosus 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. 


Upper Renging, 2150 ft. 


L. macassariensis, Schauf., subsp. anibarbis, Kuwert. 
Plate xiii, fig. 54. 

Regd. No. Penrissen 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°5-23 mm. A convex and rather coarse-looking species, 
closely resembling in general appearance the much smaller form L. cyclotaenius 
subsp. himalayae, from which it differs in the following respects only. Angle of outer 
side of mandible obsolete. Parietal ridges of head more as in L. cyclotaenius ; outer 
tubercles rudimentary, situated about twice as far from anterior angles of head as from 
the long inner tuberc!es. Mesosternum as in L. huwmerosus, but with the scars and 
dull lateral bands less sharply defined. Central area of metasternwm unpunctured. 


Posterior parts of hind coxae polished. 


Leptaulax anipunctus (Zang). 
Pl. xiii, fig. 55. 
( Sansi Gorge, Chinese Frontier, Upper 


Reed. No. 2 , ro. W. Beeb>. 
eee e ( Burma, 4900-8000 ft. \ = 


is 


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 equidistant from one 


! In the specimens I have seen, but not in those described ‘by Kuwert. 


1914. ] F. H. GRAVELY : An Account of the Oriental Passalidae. 257 


another and from anterior angles of head; outer tubercles longer than inner, and 
situated only about 4 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. Scutellum and mesothoracic episterna as in 
L. dentatus and L. bicolor. 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 L. bicolor, the grooves a little broader, however, 
and the punctures correspondingly coarser, but scarcely transverse. 


Leptaulax bicolor (Fabricius). 
I. I. BICOLOR (Fabricius), s. str. 
Pl. xiii, fig. 56. 


1119 : 5 
Regd. No. —; Naga Hills ? 
3194 
59 7 Janakmukh, 600 ft., Abor Country S. W. Kemp. 
1120-2 ~ sine . 
0 7 Dafla Expedition H. H. Godwin-Austen. 
1123-6 5 . . 
== Dikrang Valley, Northern frontier 
’ ry 4 . 
: 19 8 © \ H. H. Godwin-Austen. 
of Assam J 
9506 ors! 
% = Dunsiri Valley, 500 ft., Assam ? 
1127 Fes : 
59 —— Dunsiri Valley, Assam H. H. Godwin-Austen. 
19 j 
9187 9189-93 9195-9 U 
BD 1 1 a 
9200-3 9206-10 9212-8 
29 1 1 in eed. . 
9221-8 9225-8 9230-1 Sikkim Stoliezka bequest, etes 
me 1 1 le 
9234-9 9503-4 1129-31 
1 1 19 
Stoliczka bequest, 
9172-8 9364-66 9494-5 3 . 
» Ti aye 1 Darjeeling J. Wood-Mason. 
? 
2869-72 matey ees 
ne a Nilgiri Hills H. E. Andrewes. 
* = Between Sukli and Misty Hollow, 
Dawna Hills, Tenasserim, 2100- ( F. H. Gravely. 
2500 ft. 
1182 
” 19 y ? 
2614-6, . 
ie = Kuching, Sarawak Sarawak Museum. 
2873-5 : 
AS oe Mt. Kina-Balu, N. Borneo 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 
ridges 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. bicolor. 

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


258 Memotrs of the Indian Museum. [Vou. IIT, 


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. Andrewes’ collection by specimens from Karkur Ghat (1500 ft., May ro1r) and 
Nadgam (2500 ft., Oct. rg1o0) 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. Lakhimpur hills in Assam. 

Description.—Length 16-25 mm. Labrum and mentum much 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. dentatus. Scutellum and mesothoracic episterna also as in that 
species. Mesosternwm 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,but in smaller ones usually roughened at least in front 
by fine close puncturing of varying distinctness, a few large scattered punctures some- 
times present; auterior 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 e/ytva not wider than the ridges between them, the strong punc- 
tures with which they are marked not distinctly drawn out to form transverse 
grooves. 

Habits.—Mr. H. I,. 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 


1914. | F. H. Gravety: An Account of the Oriental Passalidae. 259 


suppose, to keep between the bark and the wood of decaying trees instead of working 
into the rotten wood as the other Passalidae I know do.” 


2. J, BICOLOR var. VICINUS (Perch.). 


iy Pl. xiii, fig. 56a. 
Regd. No. <= Ceylon I. M. Mackwood. 


19 


2622 


Ht = Ganiaduwa, Ceylon ie 
eas - Colombo Museum. 
ss ra Dimbula, Ceylon 
ae oe Reneoon : 
oe ; = 
ie pe Moulmein Museum Collector. 
9175-7 9318-28 9325-32 
1 1 l 
9334-9 9341-51 9353-5 
1 1 I ele 
9496-7 4087-8 4096 | 4 Wdamans ( Stoliczka bequest, 
” a = me d 
: eee ( J. Wood-Mason, ete. 
3721 3724-6 ©8728 
~ Ww im) 10 
3730-2 1137-8 
10 19 
/ 9178 9181 9184-5 \ 
1 1 1 ae 
*9359-62 499 9501 | Nigobars Stoliczka bequest, 
} : ae J. Wood- Mason, ete. 
519 6425 1139 
\ 6 14 19 
2877-8 Java 
19 
aS H. EK, Andrewes. 
719 Batjan 
6397 6: 6399 1141 5 i 
2 Tape cia nota) Sinkep Island Moti Ram. 
7074 Pra : 
9 a Philippine Isles East India Co.’s Museum. 
1140 
rs Ti ? > 
2617 us 5 
=r Kuching, Sarawak Ms 
HF} 2618 B j Sarawak Museum. 
19 orneo 


This form is represented in the Sarawak Museum collection by specimens from 
Kuching and Matang Road, in the Colombo Museum collection by specimens fron 
Gamaduwa 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. Pronotwm less strongly punctured ; large specimens with a few sparce punc- 
tures in anterior angles, a few set very close together in scars and, asa rule, a few 
more scattered near outer margin ; small specimens more densely pungannes especially 
in neighbourhood of scars.' Mesosternum always highly polished, its scars with 
inner wall usually longer and more strongly arched. Puncturing of intermediate areas 
of metaste/num sometimes less distinct or more irregular; central area always highly 
Deseae Lateral Se areas of BU EEEESS steyna more or less obsolete. 


: The Pecrae of the Peeaataran in sinall specimens is very ee that found in eeher 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. 


260 Memoirs of the Indian Museum. [Vou III, 


Leptaulax roepstorfi, Kuwert. 
Pl. xin, fie. 57. 


This species and the next are the most strongly flattened forms known to me. 


3172-87 


Regd. No. at Yambung, 1100 ft., Abor Country 
as Rotung, 1400 ft., Abor Country Is, eee 
= ms Misty Hollow, W. side of Dawna N. Annandale. 
Hills, Tenasserim, c. 2200 ft. 

34 “ Tenasserim ? 

i me es Tavoy Museum Collector 
(9384-7 4098-9 4104 4111 

” js es Lare ie ae 2 J. Wood-Mason, ete. 

5 iy ie shy 


Description.—Length 15-18 mm. Mandibles sometimes without ridge between 
upper tooth and first terminal tooth ; otherwise as in L. bicolor. 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, 
scutellum and mesothoracic episterna as in all preceding species of the genus; meso- 
sternum as in L. anipunctus, the horizontal central area occasionally marked with 
obscure fine punctures. Metasternum 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 stevna with usual anterior 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 elytva 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 
logs. 

Leptaulax planus (lIlliger). 


Pl. xiii, fig. 58. 


Regd. No. Tenasserim ! Museum Collector. 
7 ae Tavoy Museum Collector. 
(a 9391-402 ) 
L 1 ° 
$3 {950s eae j Johore, Malay Peninsula J. Wood-Mason, etc. 
Vie ae 
9509 951) “ce Malacca 2? ? 


” l i 


1 According to Register ‘‘ Hills between Tenasserim and Siam, about lat. 13° 45’, long. g8°.’’ 


1914. | F. H. Gravety: An Account of the Oriental Passalidae. 261 


Regd. No. “332? Borneo Sarawak Museum. 
BS “ “ Sinkep Island Museum Collector. 
A ad 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. Pvonotwm 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 metasternum normally marked thus ; intermediate areas of metasternum 
strongly punctured throughout. Hind coxae polished. Abdominal sterna uniformly 
marked with large shallow punctures quite unlike those found in any other species of 
the genus that is known to me. 


Genus TRICHOSTIGMUS, Kaup. 
Trichostigmus thoreyi, Kaup. 


Described from a single specimen from Middle Luzon, in the Hamburg Museum 
collection. 

Description.—Length 24 mm. Head and its appendages much as in the specimen 
of Leptaulax bicolor shown on pl. 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 scarfs ; 
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. Mesothovacic episterna punctured above and along 
almost whole length of anterior margin. Mesosteynwm smooth except in scars which 
are rugose. Lateral areas of metasterynum narrow, linear, finely roughened, widely 
separated from central area; posterior intermediate areas punctured beside inner 
margin, smooth laterally and behind. Hind coxae and abdominal stevna 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. ° (Vox. III, 


5. THE VARIABLE SPECIES. 


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. They are Leptaulax bicolor (length 13-25 mm.), and Leptaulax dentatus (19°5-37 
min.). 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, Episphenus 
indicus (25-41 mm.), Episphenus comptont (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 (30-4c mm.), Leptaulax humerosus (15-20), Leptaulax planus (11-14 mm.}, 
Pleurartus brachyphyllus (35°5-45 mm.) and Ophrygonius cantori (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 Cevacupes austent (20-25 mm.) and Tiberioides 
kuwertt (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 bicolor, Leptaulax dentatus, Episphenus indicus, Epispenus comp- 
toni, and Aceratus 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 neelgherriensis, 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 comptomi, 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 27°5, one 
30, and one 37 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. 


IgI4. | F. H. GRAvELY: An Account of the Oriental Passalidae. 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 indicus and Aceratus 
grandis sub-sp. ivsutus 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 grandis. 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 Leptaulacinae all 
pointsin the same direction. Lepiaulax 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 


! 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. [Vox III, 


of what has been found. Out of 46 specimens labelled ‘‘ Upper Tenasserim’’ 17 are 
21°5-24'5 mm.long ; 2 are 26°5 and 28 mm. long respectively ; and 27 are 30-37 mm. 
long. Outof 45 specimens labelled ‘‘ Tenasserim’’ 21 are 21-24°5 mm. long, 1 is 
27°5 mm. long; and 23 are 30-35 mm. long. And out of 43 specimens labelled 
‘Sikkim "’ 2 are 19°5 and 20 mm. long respectively, 8 are 21-23 mm. long, and 33 are 
245-31 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. 

Leptaulacides 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 different forms of the species, which must, at 
least provisionally, be treated separately, these specimens have proved not to be nearly 
enough to forma 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 units, 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 taxonomic 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 Passalidae 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 
belongs. 


I9T4. | F. H. GRAVELY: An Account of the Oriental Passalidae. 265 


In the Passalidae, asin the Lucanidae, 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 Iucanid 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 
cvandts 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. 52d). 

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 
(2 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. roepstorffi) the front of the head bears two median protuberances, one above 
the cther; 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 Lucanidae (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 and 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. 


6. THE GEOGRAPHICAL DISTRIBUTION AND SYNONYMY OF THE 
SPECIES OF PASSALIDAE BELONGING TO GENERA FOUND IN 
THE ORIENTAL REGION. 


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 Memotrs of the Indian Museum. [Vor. III, 


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 and 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 known from the Oriental Region 
together with 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 which 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 (ft) 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 Passalids recorded from ‘‘ Sikkim ’’ belong to species found in ‘‘ British 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, 


Z 


1914. | F H. Gravety: An Account 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. 

Subfamily AULACOCYCLINAE. 


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], 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, Kuwert, 1897, p. 284. 
Comacupes ¢ minor, Heller, 1896, pp. 3-4, pl. i, figs. I1-Ila. 

The hairless abdomen and practically unpunctured mesosternum suggest that 
this species may be more closely allied 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. 


calities :— 
Localities 1. C. FOVEICOLLIS, s. str. 
Borneo: Sarawak—Baram River. 

2. Sub-sp. Minor, Heller. 


Celebes: N. coast—Buol. 


Comacupes basalis (Smith). 
Passalus + basalis, Smith, 1852, p. 18, pl. 1, fig. 5. 
Aulacocyclus comatus, Kaup, 18682, p. 9. 
(Aulacocyclus comatus + Leptaulax basalis , Gemminger and Harold, 1868, pp. 968-9). 
Comacupes basalis, Kaup, 1871, p. 19, pl. iii, figs. 34-30. 
(Comacupes basalis, Wytsmann, 1884, p. 329). 
(Comacupes Basalis, Kuwert, 1891, p. 171). 
Comacupes basalis, Kuwert, 1896, pp. 284-5. 
(Comacupes basalis, Zang, 19030, pp. 418-9). 


Locality :— 
Philippine Islands. 


268 Memoirs of the Indian Museum. (Vor, Ti; 


Comacupes cavicornis, Kaup. 
Aulacocyclus cavicornis, Burmeister in Kaup!, + /aevicornis, Kaup, 1868a, p. 6. 
(Aulacocyclus cavicornis + laevicornis, Gemminger and Harold, 1868, pp. 698-9). 
Comacupes cavicornis, Kaup. 1871, p. 19. 
(Aulacocyclus | parryi, Stoliczka, 1873, p. 150). 
(Comacupes cavicornis, Wytsman, 1884, p. 329) 
(Comacupes Cavicornis [part], Kuwert, 1891, p. 171). 
Comacupes } westermanni, Kuwert, 1897, p. 284. 
Comacupes cavicornis, Zang, 1905¢, p. 226. 
Comacupes } cavicornis + var. tborneensis + var. f laevicornis, Gravely, above, pp. 204-205, pl. xi, 
figs. 1-2). 

All the specimens identified by Stoliczka as Aulacocyclus parryt have proved to 
be in reality Comacupes cavicornis var. laevicornis. Zang has pointed out that Kuwert’s 
Comacupes westermanni, not his Tristorthus cavicornis, is identical with Kaup’s Coma- 
cupes cavicornis. 

Localities :— 

Malay Peninsula: Penang; Johore (? all var. /aevicornis). 
Sumatran Islands; Sumatra; Sinkep Island (? all typical form). 
Java (? another form).” 

Borneo Sarawak:—Kuching (? all var. borneensis). 


Comacupes stoliczkae, Gravely. 


Comacupes cylindraceus (part), Kaup, 1871, p. 18, pl. iii, upper unit of fig. 4. 
Comacupes + cylindraceus, Stoliczka, 1873, p. 151. 

?(Comacupes cylindraceus, Kuwert, 1891, p. 171). 

? Comacupes cylindraceus [part] Kuwert, 1897, p. 285. 
Comacupes } stoliczkae, Gravely, above, p. 206, pl. x1, figs. 3-34. 

This species was described by Kaup asa 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. 
Java. 

Kuwert definitely recorded this species from Singapore in 1891; 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 tmasont, Stoliczka, 1873, pp. I5I- 2. 
Comacupes Masont, 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. 


I9r4. | F. H. GraAvELy : An Account of the Oriental Passalidae. 269 


Comacupes } cylindraceus, [part], Kuwert, 1897, p. 285. 
Comacupes t masont, Gravely, above, p. 207, pl. xi, figs. 4-4a. 


Zang has suggested that Kuwert’s C. masoni of 1897 is not Stoliczka’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: Johore; 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 Stoliczka’s account 
of it. 


Comacupes cylindraceus (Perty). 
Passalus cylindraceus, Perty, 1831, p. 36, fig. 3. 
Passalus cylindraceus, Percheron, 1841, pp. 40-1, pl. Ixxix, fig. 4. 
Passalus punctifrons, Hope, 1845, p. 29. 
Passalus cylindraceus, Burmeister, 1847, pp. 516-7. 
(Aulacocyclus cylindraceus [part], Kaup, 18682, 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 vylindraceus, Zang, 1903), p. 419). 
(Comacupes | Masoni, Zang, 1905c, p. 231). 
Comacupes f cormocerus, Zang ; 1905d, pp. 233-5. 
Comacupes = punctifrons, Arrow, 1907, p. 444. 
Comacupes } cylindraceus, Gravely, above, p. 207, pl. xi, figs. 5-6a, text-fig 2. 
Other references to the name cylindvaceus will be found under C. stoliczkae 
(previous page). 
Localities :— 
Malay Peninsula: S. Perak (Telom, 4ooo ft.), Larut Hills, 3300-4300 ft., 
Singapore. 
Sumatran Islands: Sinkep Island. 
Sumatra—Padang ; St. Rambé. 
Java. 


Borneo: Sarawak—Kina-Balu, ca. 5000 {t.; Kuching; Trusan ; Penrissen 4200- 
4500 {t.; 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. 

2 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 from the Malay Peninsula, the anterior part of the 


central tubercle, though much elevated, being angular beneath; it may perhaps belong to a distinct 
local race. 


270 Memoirs of the Indian Museum. [Vor,. III, 


Genus TAENIOCERUS, Kaup, 1871. 
= Aulacocyclus [part], Kaup, 1868; Taentocerus [part], Kaup, 1871. 


This genus includes species found in all parts of the Oriental Region except the 
Indian Peninsula and Ceylon. 
T. platypus, Kaup. 
Aulacocyclus platypus, Kaup, 1868a, p. 5. 
(Aulacocyclus platypus, Gemminger and Harold, 1868, p. 969.) 
Taeniocerus platypus, Kaup, 1871, p. 21. 
Aulacocyclus platypus, Lewis, 1883, p. 341. 
(Taeniocerus platypus, Wytsman, 1884, p. 329.) 
(Taeniocerus Platypus, Kuwert, t891, p. 170.) 
Taeniocerus platypus, Kuwert, 1897, p. 275. 


Localities | :— 
Malay Peninsula: Perak ; Naungchik (Bukit Besar, 2500 ft.).’ 
Sumatra: Deli. 
Borneo: Sarawak. 


Taeniocerus bicanthatus (Percheron). 
Passalus bicanthatus, Guérin-Méneville* in Percheron, 1841, pp. 41-2, pl. xxix, fig. 5. 
?Passalus bicanthatus, Burmeister, 1847, p. 516. 
(Passalus bicanthatus, Smith, 1852, p. 17.) 
Aulacocyclus bicanthatus, Kaup, 1868a, p. 6. 
(Aulacocyclus bicanthatus, Gemminger and Harold, 1868, p. 968). 
Taeniocerus bicanthatus, Kaup, 1871, p. 21, pl. iii, fig. 6. 
Taeniocerus + bicanthatus, Stoliczka, 1873, p. 152. 
(Taeniocerus bicanthatus, Wytsman, 1884, p. 329.) 
(Taeniocerus Bicanthatus, Kuwert, 1891, p. 170.) 
Taeniocerus + bicanthatus + ab. perturbans + imitator, Kuwert, 1897, p. 276. 
Taeniocerus bicanthatus, Zang, 19054, p. 105. 
Taenioverus + bicanthatus, Gravely, above, pp. 208-209, pl. xi, figs. 7-70. 


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 Taentocerus, 


| Borneo is the locality given by Kaup, who meutions 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. 

2 This record is based on a pair of specimens in the Oxford Museum. 

® Percheron gives Guérin-Méneville as the author of this and other names, without giving a reference. 
In the case of Passalus (Episphenus) neelgherriensis, where he gives as reference ‘‘ Rev. Zool.’’, a search 
through all possible volumes of that publication has failed to reveal any mention of the species; and as 
Guérin-Méneville himself wrote (1843, p. 50) two years after the publication of Percheron’s paper 
“Cette espéce a éte découverte par M. Ad. Delessert sur le plateau des Neelgheries. Nous 1’avons 
communiqué 4 M. Percheron, qui l’a fait entrer en notre nom, dans sa Monographie... . ’’ 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. 


1914. |] F. H. Gravety: An Account of the Oriental Passalidae. 271 


is the mesosternum hairy ; whereas this is the case in every species of Comacupes 
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 
Taeniocerus 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. 


Localities : — 
Malay Peninsula: Johore. 
Sumatra: N. E. Sumatra. 
E. Sumatra—Langkat. 
Bornean Islands: Borneo. 
Labuan. 


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, 1868a, p. 5. 

(Aulacocyclus pygmaeus, Gemminger and Harold, 1868, p. 969.) 
Taemocerus pygmaeus, Kaup, 1871, pp. 20-21. 

(Taentocerus + pygmaeus, Stoliczka, 1873, p. 152.) 

(Taemocerus pygmaeus, Wytsman, 1884, p. 32g.) 

(Taentocerus Pygmaeus, Kuwert, 1891, p. 170.) 

Taentocerus f parvus + pygmaeus, Kuwert, 1897, p. 270. 

(Taentocerus pygmaeus, Zang, 1903), p. 419.) 

Taenwocerus + pygmaeus, Gravely, above, pp. 209-210, pl. xi, figs. 8-8a. 


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. 
Sumatra. ! 
Borneo: Sarawak—Kuching. 


! This record is based on specimens in Berlin, not seen in time for description in part 4 of this 


paper. 


272 Memoirs of the Indian Museum. [Vo1,.. TE, 


Taeniocerus bicuspis, Kaup. 
? Passalus bihastatus 2 , Percheron, 1844, p. 13. 
Aulacocyclus bicuspis, Kaup, 1868a, p. 5. 
(Aulacocyclus bicuspis, Gemminger and Harold, 1868, p. 968.} 
Taentocerus bicuspis, Kaup, 1871, pp. 21-2. 
Taentocerus } bicuspis, Stoliczka, 1873, p. 152. 
(Taentocerus bicuspis, Wytsman, 1884, p. 329.) 
(Taentocerus Bicuspis, Kuwert, 1891, p. 170 ) 
Taentocerus bicuspis, Kuwert, 1897, pp. 276-7. 
Taeniocerus tbicuspis, Gravely, above, pp. 210-211, pl. xi, figs. 9-ga. 
Localities :— 


E. Himalayas: Darjeeling Dist.—Sureil; Choongtang; Upper Tista, 4000 ft. ; 
Gopaldhara, Rungbong Valley. 
Dafla Hills—Harmutti, base of Dafla Hills; Dikrang Valley. 
Abor Country—Kobo, 4oo ft.; Rotung, 1400 ft.; Upper 
Rotung, 2000 ft.; Upper Renging. 
Assam: Cachar. 
N. Lakhimpur—Silonbari (base of hills). 

Siam. 

Malay Peninsula. 

Sumatra’. 

Borneo. ' 


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 (1896, 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 aminclined 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. 


[Aulacocyclus deyrollei, Kaup. | 
Aulacocyclus Deyrollei, Kaup, 1868a, p. 7. 
(Awlacocyclus Deyrollei, Gemminger and Harold, 1868, p. 968.) 
Taemtocerus Deyrollet, Kaup, 1871, p. 20. 
(Taeniocerus Deyrollet, Wytsman, 1884, p. 329.) 


1 These records are based on specimens in Berlin, not seen in time for description in part 4 of 
this paper. 


1914. | F. H.Gravety: An Account of the Oriental Passalidae. 273 


(Laentocerus Deyrollet, Kuwert, 1891, p. 170.) 
Taentocerus deyrollet, Kuwert, 1897, p. 275. 
Taeniocerus deyrolle:, Arrow, 1907, p. 447. 


Lacality :— 
Australia: N. Queensland—Port Denison’. 


[Aulacocyclus mastersi, Macleay. | 
* Taeniocerus mastersi, MacLeay, 1871, p. 174. 
(Taeniocerus Mastersti, Wytsman, 1884, p. 329.) 
(Taeniocerus Mastersit, Kuwert, 1891, p. 170.) 
Taeniocerus masterst, Kuwert, 1897, p. 275. 


It is possible, I think, that this species and the last may prove to be identical. 


Locality :— 
Australia: Queensland—Gayndah °. 


[* Aulacocyclus teres (Percheron). | 


Passalus teres, Percheron, 1841, pp. 39-40. 

Passalus teres, Burmeister, 1847, p. 515. 

Passalus teres, Smith, 1852, p. I9. 

Passalus teres, Redtenbacher, 1867, p. 50. 

Aulacocyclus teres, Kaup, 18682, p. 9. 

(Aulacocyclus teres, Gemminger and Harold, 1868, p. 99.) 
Aulacocyclus teres, Kaup, 1871, pp. 15-16, pl. iii, figs. I-1a. 
(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 politus, concerning which see Stoliczka, 1873, p. I 56. 


[* Aulacocyclus tereoides, Kuwert]. 
Aulacocyclus Tereoides, Kuwert, 1891, p. 170. 
Aulacocyclus tereoides, Kuwert, 1897, p. 277. 
Locality :— 
Australia. 
[(* Aulacocyclus aliicornis, Kuwert.| 


Aulacocyclus alicornis + ab. sulcatipons, Kuwert, 1897, pp. 277-9- 
Aulacocyclus alicornis ab. sulcatupons, Zang, 1905C, p. 230. 


' Not St. Denis, Réunion Islands (see Arrow loc. cit.). 

2 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 Museum. [Vou. III, 


Locality :— 
Australia. 
|* Aulacocyclus foveipunctatus, Kuwert. | 
Aulacocyclus fovetpunctatus, Kuwert, 1897, p. 278. 
Locality :— 
Australia. 


|* Aulacocyclus, rotundatoclypeatus, Kuwert. 
Aulacocyclus Rotundatoclypeatus, Kuwert, 1891, p. 170. 
Aulacocyclus rotundatoclypeatus, Kuwert, 1897, pp. 278 and 282. 
Locality :— 
Australia: Cape York. 


[* Aulacoeyclus arcuatoclypeatus, Kuwert. | 
Aulacocyclus Arcuatoclypeatus, Kuwert, 1891, p. 170. 
Aulacocyclus arcuatoclypeatus, Kuwert, 1897, pp. 278 and 282. 
Locality :— 
New Guinea : Fly River. 


* Aulacocyclus fratricornis, Kuwert. |] 
Aulacocyclus Frairicornis, Kuwert, 1891, p. 170. 
Aulacocyclus fratricornis, Kuwert, 1897, pp. 279 and 282. 
Locality :— 
? South Sea Islands or Australia. 


Aulacocyclus rosenbergii, Kaup. 
Lulacocyclu; Rosenbergu + Percheroni, Kaup, 18684, pp. 7 and 8. 
(Aulacocyclus Rosenbergi + Percheront, Gemminger and Harold, 1868, p. 969.) 
Aulacocyclus Rosenbergit, Kaup, 1871, p. 10. 
(Aulacocyclus Rosenbergu, Wytsman, 1884, p. 328.) 
(Aulacocyclus Rosenbergit, Kuwert, 1891, p. 170.) 
Aulacocyclus Rosenberg + Percheront, Blackburn, 1896, p. 234. 
Aulacocyclus rosenbergt Kuwert, 1897, pp. 279 and 282. 
(Aulacocyclus Rosenbergit, Zang. 1903), p. 410.) 
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. rosenbergit. 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. percheront. 


| Kaup gives ‘‘Trisbane,’? but Gemminger-and Harold are presumably right in regarding this 


as a misprint for Brisbane. 


1914. | F. H. GRAVELY: An Account of the Oriental Passalidae. 


[* Aulacocyclus kaupi, Macleay. | 
*Aulacocyclus kaupt, Macleay, 1871, p. 173. 
(Aulacocyclus Kaupi, Wytsman, 1884, p. 328.) 
(Aulacocyclus Kaupi, Kuwert, 1891, p 170.) 
Aulacocyclus Kaup1, Blackburn, 1896, p. 234. 
Aulacocyclus kaupt, Kuwert, 1897, pp. 280 and 282. 


Locality :—- 
Australia: Queensland—Gayndah. 


[* Aulacocyclus edentulus (Macleay). | 


Passalus edentulus, Macleay. 1820, p. 439. 
*Passalus furcicornis, Boisduval, 1835, p. 242. 
Passalus cylindraceus, Percheron, 1835, pp. 103-4, pl. vii, fig. 8. 
*Passalus furcicornis, Dejean,! 1837, p. 195. 
Passalus edentulus, Percheron, 1841, p. 39. 
Passalus edentulus, Burmeister, 1847, pp. 515-0. 
(Passalus edentulus, Smith, 1852, p. 20.) 
Passalus furcicornts, Montrouzier, 1855, p. 29. 
Passalus furcicornis, Montrouzier, 1860, p. 288. 
Passalus furcicornis, Fauvel, 1862, p. 135. 
(Aulacocyclus edentulus, Kaup, 18682, p. 8.) 
(Aulacocyclus edentulus, Gemminger and Harold, 1868, p. 969.) 
Aulacocyclus edentulus, Kaup, 1871, pp. 14-15. 
(Aulacocyclus edentulus, Wytsman, 1884, p. 328.) 
(Aulacocyclus Edentulus, Kuwert, 1801, p- 170.) 
Aulacocyclus edentulus, Blackburn, 1896, 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, pl. 233. 
Locality :— 


Australia. 
[* Aulacocyclus collaris, Blackburn. | 


Aulacocyclus collaris, Blackburn, 1896, pp. 233-4. 
Locality :— 


Australia. 
Aulacocyclus andrewesi, Gravely. 


Aulacocyclus ¢ andrewesi, Gravely, above, pp. 211-212, pl. xi, figs. 10-10a. 
Locality :— 
India: Anamalai Hills, ca. 3500-4000 ft. 


! Gemminger and Harold give d’Urville as the author of this name in Dejean’s catalogue. 


N 
™S 
On 


Memoirs of the Indian Museum. [Vou. ITI, 


\* Aulacocyclus glabriusculus, Kuwert. | 


Aulacocyclus Glabriusculus, Kuwert, 1891, p. 170. 
Aulacocyclus glabriusculus, Kuwert, 1897, pp. 280 and 282. 


Locality :— 
Aru Islands. 
*Aulacocyclus dilatus, Kuwert. 


Aulacocyclus Dilatus, Kuwert, 1891, p. 170. 
Aulacocyclus dilatus, Kuwert, 1897, pp. 281 and 282. 


Locality :— 
Philippine Islands. 


Aulacocyclus parryi, Kaup. 
Aulacocyclus Parryi, Kaup, 1868a, p. 8. 
(Aulacocyclus Parryi, Gemminger and Harold. 1858, p. 969.) 
Aulacocyclus Parryi, Kaup, 1871, p. 16. 
Aulacocyclus parryi, Kirsch, 18774, p. 139. 
(Aulacocyclus Parryi, Wytsman, 1884, p. 328.) 
Aulacocyclus Parreyi, Kuwert, 1891, p. 170. 
Aulacocyclus parryi, Kuwert, 1897, p. 281. 
Aulacocyclus parreyi, Kuwert, 1897, p. 282. 


Localities :— 
? Java. 
Moluccas: Ceram. 
New Guinea: Rubi. 
? Australia. 


Kuwert in 1897 omits all reference to his Javanese record of I8gI, so it may be 
doubted whether this species is really Oriental! 


(* Aulacocyclus perlatus, Kaup.| 
Aulacocyclus perlatus, Kaup, 1868a, p. 7. 
(Aulacocyclus perlatus, Gemminger and Harold, 1868, p. 969.) 
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. 
? Australia. 


The Australian record rests solely on the authority of Kuwert’s 1891 list; it is 


omitted in his subsequent paper. 


' There are specimens in the Stuttgart Museum collection from Java, which perhaps belong to 


this species. 


TQI4.| F. H. GravELy. An Account of the Oriental Passalidae. 


Aulacocyclus aruensis, Kuwert. 


Aulacocyclus Pavreyi var. arwensis, Kuwert, 1891, p 170. 

Aulacocyclus taruensis, Kuwert, 1897, p. 282. 
Localities : — 

Borneo: Sarawak. ! 

Aru Islands. 


[* Aulacocyclus celebensis, Heller.] 
Aulacocyclus celebensis, Heller, 1898, pp. 22-3. 
Locality :— 
Central Celebes: Luhn-Djaladja. 


[* A. variolosus, Kuwert. | 
Aulacocyclus variolosus, Kuwert, 1897, p. 282. 


Locality :— 
New Guinea. 


[*A. pugnax (Fauvel) | 


*Comacupes pugnax, Fauvel. 1903, p. 359. 
Aulacocyclus pugnax, Zang, 1905c, p. 228. 
Locality :— 
New Caledonia. 


Genus _ERACUPES, Kaup, 1871. 
= Aulacocyclus [part], Kaup, 1868. 


The genus Ceracupes is only known from the E. Himalayas, Tibet, 
Upper Burma and Formosa. 


Ceracupes fronticornis (Westwood). 


Passalus fronticornis, Westwood, 1842, pp. 124-5. 
Passalus bihastatus 3, Percheron, 1844, pp. 12-13, pl. cxxxv, fig. 3. 
* Passalus fronticornis, Hope, 1845. 
Passalus bthastatus, Burmeister, 1847, p. 517 
Passalus bihastatus, Smith, 1852, p. 17. 
(Aulacocyclus bihastatus, Kaup, 1868a, p. 6.) 
(Aulacocyclus bihastatus, Gemminger and Harold, 1868, p. 968.) 
Ceracupes fronticornis, Kaup, 1871, p. 17, pl. iii, fig. 2 20 
Ceracupes fronticornis, Stoliczka, 1873, p. 151. 
(Ceracupes frontcornis, Wytsman, 1884, pp. 328-9.) 
(Ceracupes Fronticornis, Kuwert, 1891, p. 171.) 
Ceracupes fronticornis, Kuwert, 1897, p. 273. 


Tonkin , 


' Kuwert has determined as 4. aruensis a specimen preserved in the K6nigliches Zoologisches 


Museum in Berlin, collected by Wallace in Sarawak. 


278 Memoirs of the Indian Museum. (Vor. TIL, 


Ceracupes tfronticornis [ ? part], Zang, 1g05a, p. roo. ! 
Ceratocupes fronticornis, Arrow, 1907, p. 444. 
Ceracupes F fronticornis, Gravely, above, p. 212, pl. xi, fig. 12. 

Zang refers Fruhstorfer’s Nigidius-like Passalid from Ke-Lung in Formosa 
(Fruhstorfer, rg02, 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 :— 

FE. Himalayas: Nepal. 
Darjeeling District—Gopaldhara, Rungbong Valley. 
Tibet. 
Tonkin: Mt. Mauson 2000-3000 ft.; Than 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 Entomolgisches Museum. Its horn is 
more slender and more strongly widened and turned up at the end, and its elytra 
ate 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 
localities. 


Ceracupes arrowi, Heller. 
*?Fruhstorfer, 1902, p. 28. 


? Ceracupes fronticornts [part], Zang, 19054, p. 100. 
Ceracupes Arrowit, Heller, 1911, pp. 256-7, text-fig. 
Ceracupes + arrowt, Gravely, I914a, p. 31. 

Localities :— 
Fomosa: Kosempo; Hoozan;? Ke-Lung. 


Ceracupes austeni, Stoliczka. 
Ceracupes + austent, Stoliczka, 1873, p. 151. 
(Ceracupes Austeni, Wytsman, 1884, p. 329.) 
(Ceracupes Austent, Kuwert, 1891, p. 171.) 
Ceracupes austent, Kuwert, 1897, p. 274. 
Ceracupes } austent, Gravely, above, pp. 212-213, pl. xi, figs. 11-1Ia. 
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 
cornutus (see Zang, loc. cit.). 
* ‘These records are based on specimens in the British Museum. 


19I4.] F. H. Gravety: An Account of the Oriental Passalidae. 279 


Stoliczka recorded this species from the ‘‘ Naga Hills, North-Eastern 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 
sense. 


Genus CYLINDROCAULUS, Fairmaire, 1880. 


Only one species of this genus is yet known; it comes from China. 


*C. bucerus, Fairmaire. 

* Cylindrocaulus bucerus, Fairmaire, 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, 1905¢, pp. 228-9. 
Cylindrocaulus bucerus, Arrow, 1907, p. 446. 

Locality :— 
Western China: Sz-T'schwan—Moupin. 


Genus AURITULUS, Zang, 1905. 


This genus is only known from Japan. 


Auritulus patalis (Lewis). 

Aulacocyclus + patalis, Lewis, 1883, p. 341, pl. xiv, figs. 6-7. 

Auritulus patalis, Zang, 1905¢, pp. 228-6. 

Aurtkulus tpatalis, Arrow, 1907, p. 440. 

Locality :— 

Japan: ‘‘in 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. 


Subfamily PLEURARIINAE. 


Of this sub-family only one genus of two species is known. 


Genus PLEURARIUS, Kaup, 1880. 


This genus is recorded from Sumatra and the Indian Peninsula. 


*Pleurarius pilipes, Kaup. 
Pleurarius pilipes, Kaup, 1868), pp. 1-2. 
(Pleurarius pilipes, G mininger and Harold, 1868, p. 970.) 
Pleurarius pilipes, Kaup, 1871, p. 28, pl. 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. 


280 Memoirs of the Indian Museum. [Vou. III, 


Pleurarius brachyphyllus, Stoliczka. 
Pleurarius ¢ 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 } brachyphyllus, Gravely, above, pp. 213-214, pl. xi, figs. 13-134. 
Localities :— 

Travancore: Kulattupuzha (W. base of W. Ghats). 
Cochin: timber forest. 
Madras Presidency: Malabar. 

Nilgiri Hills. 

Anamalai Hills—Puddutottam, 4200 ft., and other un- 

recorded localities up to 5500 ft. 
S. Arcot or Chengalput—Palur. 


Subfamily ACERAIINAE. 


This sub-family occurs throughout the Oriental Region to which it is probably 
confined. 


Genus TIBERIOIDES, Gravely, 1913 


= Aceraius [part] Kaup, 1868; Basilanus [part] Kaup, 1869; Tiberius [part] 
Kuwert, 1896-8. 


The genus Tiberioides is only known from the E. Himalayas and Naga Hills. 


Tiberioides kuwerti (Arrow). 
Aceraius cancrus, Kaup, 18684, p. 29. 
Aceraius cancrus, Kaup, 18686, p. 4. 
(Aceraius cancrus, Gemminger, and Harold, 1868, p. 972.) 
Bastlianus cancrus, Kaup, 1871, pp. 56-7. 
Basilianus ¢ cancrus, Stoliczka, 1873, p. 160. 
(Bastlianus cancrus, Wytsman, 1884, p. 336.) 
(Tiberius Cancrus, Kuwert, 1891, p. 164.) 
Tiberius t cancrus, Kuwert, 1808, p. 188. 
Tiberius kuwerti, Arrow, 1906, p. 446. 
Tiberioides + kuwerti, Gravely, above, pp. 215-216, pl. 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. 
Bhutan. 
Assam: Naga Hills—Manipur. 
Burma. 


1g 14.] F. H. GRAVELY: An Account of the Oriental Passalidae 


Tiberioides austeni, Gravely. 
Tibertoides + austent, Gravely, above, pp. 216-217. pl. xi, figs. 15-15@. 
Localities :— 
E. Himalayas: Dafla Hills—Dikrang Valley. 
Abor Country—Kalek, 3800 ft. 


Tiberioides borealis (Arrow). 
Chilomazus ¢ borealis, Arrow, 1906, pp. 467-8. 


Locality :— 
Assam: Naga Hills. 


Genus EPISPHENUS, Kaup, 1871. 


281 


Incl. Aceraius [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 Mooret, Kaup, 1871, p. 45. 
(Episphenus Mooret, Wystman, 1884, p 334.) 
(Episphenus Mooret, Kuwert, 1891, p. 165.) 
Episphenus { mooret, Kuwert, 1898, p. 189 
Episphenus ¢ mooret, Zang. 1905a, p. 163. 


Episphenus + moorei + + pearsoni, Gravely, above. pp. 217-218, pl. xi, figs. 16-17. 
Localities :— 


Ceylon: Central Province—Nitre Cave district, c. 1800-3000 ft. ; 
Nalanda. 


Ratnapura District—Bulutota. 


The series of specimens in Berlin and the British Museum prove conclusively 
that the characters by which I supposed E. fearsoni to be distinguished from 


E. mooret (see above, loc. cit.) have no real taxonomic value. 


Episphenus comptoni (Kaup). 
Aceraius Comptont, Kaup, 1868a, p. 28. 
Aceraius Comptont, Kaup, 1868), pp. 3-4. 
(Aceraius Comptont, Gemminger and Harold, 1868, p. 972.) 
Laches Comptonii, Kaup, 1871, pp. 49-50, pl. iv, figs. 5-54. 
(Laches Comptonit, Wvtsman, 1884, p. 335:) 
Laches Comptonti + Flachii + Frustorferi + Puella, Kuwert, rX9r, p. 167. 


. . a . nig ~ n , o 
Laches parallelogrammifrons + fruhstorferi, + puella + flachi + socius + comptoni, Kuwert, 


1898, pp. 328-9. 
Laches + Comptoni, Zang, 19054, p. 163. 


1 See Appendix I, p. 316. 


Episphenus + comptoni + var. + flachi, Gravely, above, pp. 218-219, pl. xi, figs. 18-192. 


282 Memoirs of the Indian Museum. [Vou. III, 


Zang has shown that Kuwert’s species are all of them identical with C. comptont, 
Kaup ; and the specimens I have seen confirm this’. 


Localities :— 
Ceylon: Central Province—Hakgalla; Patipolla c. 6000 ft.; Namunakuli Hill. 
Uva—Haputale. 
Ratnapura District—Bulutota. 


Episphenus indicus (Stoliczka). 
Aceraius Cantort [part] + neelgherriensis [? part], Kaup, 18682, pp. 28 and 30. 
Aceraius Cantor: [part] + neelgherriensts [? part], Kaup, 18682, p. 4. 
(Aceraius Cantori |part] + neelgherriensis, [part], Gemminger and Harold, 1868, p. 972.) 
Bastlianus neelgherriensis [? part], Kaup, 1871, pp. 55-0. 
Basilianus + indicus, Stoliczka, 1873, pp. 159-160. 
(Basilianus neelgherriensis {part] + indicus, Wytsman, 1884, p. 336.) 
(Basilianus Indicus + Certus, Kuwert, 1891, p. 164.) 
Basilianus indicus + stoliczkae + neelgherriensis, Kuwert, 1898, pp. 340-341. 
Basilianus indicus, Zang, 1905¢, p. 223. 
Basilianus + certus + stoliczkae + F indicus, Zang, 1g06a, pp. 180-3. 
Episphenus ¢ indicus, Gravely, above, pp. 220-222, pl. xi, figs. 20-20b. 

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 5. stoliczkae 
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 1898 paper is identical with EF. 
neelgherriensis ; that they are not identical one with another has already been indicated 
by Zang in his synonymy of r1go6. 

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 localities up to 5500 ft. 


' While this paper was still in the press I was able myself to examine Zang’s series in Berlin. It 
contains aspecimen 38 mm. long, so the only distinction found between FE. comptoni var. flachi (32-37 mm.) 
and E. comptont, 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. 


1914. | F. H. GRAVELY: An Account of the Oriental Passalidae. 283 


Nilgiri Hills—Ootacamund ; Gudalur ; Naduvatum. 
Madura—Dindigal. 

Trichinopoli. 

Cuddapah. 


Episphenus neelgherriensis (Percheron). 


Passalus neelgherviensis, Guérin-Méneville in Percheron,! 1841, p. 4, pl. lxxvii, fig. I. 
Passalus neelgherriensis, Burmeister 1847, p. 469. 
(Passalus neelgherriensis, Smith, 1852, p. 6.) 
? Aceraius neelgherriensis [part], Kaup, 18682, p. 30. 
Aceraius ¢ infantilis + + puerilis + ? neelgherriensis [part’, Kaup, 18680, pp. 4 and 6. 
(Aceraius infantilis + puerilis + neelgherniensis [part], Gemminger and Harold, 1868, p. 972.) 
Laches + infantilis + + puerilis + ? Basilianus neelgherriensis [part], Kaup, 1871. pp. 48 and 55-6. 
(Laches infantilis + puerilis + Basilianus neelgherriensis [part], Wytsman, 1884, pp. 335-6.) 
(Basilianus Neelgherriensis + Laches Puerilis + Epilaches Infantilis, Kuwert, 1891, pp. 164 and 167.) 
Analaches } filius + infantiiis + Laches puertlis + Basilianus inaequalis + certus, Kuwert, 1898, 
pp. 336, 338 and 341. 
Basilianus ft binominis, Zang, 19054, pp. 243-4. 
Basilianus } neelgherriensis, Zang, 1905¢, p. 223. 
Basilianus } neelgherriensis + + binominis, Zang, 1906, pp. 179-183. 
Episphenus + neelgherriensis, Gravely, above pp. 222-223, pl. xi, figs. 21-214. 


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 
Laches 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. 


Localities :— 
Madras Presidency : Coorg. 
Malabar. 
Nilgiri Hills—Coonoor, c. 6000 ft.'; Ootacamund, 7800 ft. 
Annamalai Hills—various localities ranging from about 
3000-5500 ft.” 
Madura—Dindigal. 


! See above, p. 270, footnote 3. 


2 For more detailed reference to localities round Coonoor and in the Annamalais see above, p. 223. 


284 Memoirs of the Indian Museum. [Vou. IIT, 


Mysore: Bababudin Hills, 4000-5000 ft. 
Cochin. 

Travancore—High Range. 

Pondicherry. 


Kuwert’s record of ‘‘ Basilianus inaequalis’’ from Malacca doubtless refers to the 
true inaequalis, 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. Basilianus [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 Philippines. It is not known from the Andamans or Nicobars. 


Ophrygonius cantori, Percheron. 

Passalus cantort, Percheron, 1844, pp. 3-4, pl. exxxiv, fig. 2. 
Passalus Cantort, Burmeister, 1847, pp. 468-9. 
(Passalus Cantori, Smith, 1852, p. 7.) 
Aceraius Cantori [part], Kaup, 18682, p. 28. 
Aceraius Cantori [? part], Kaup, 18684, p. 4. 
(Aceraius Cantort, Gemminger and Harold, 1868, p. 972.) 
Basilianus Cantori [part], Kaup, 1871, p. 57. 
Basilianus } Cantorts, Stoliczka, 1873, p. 159. 
(Bastlianus Cantori [part], Wytsman, 1884, p. 336.) 
(Basilianus Cantort, Kuwert, 1891, p. 164.) 
Basilianus cantori + finterrogationus, Kuwert, 1898, pp. 340-1. 
Basilianus + convextfrons, Zang, 1904a, p. 698 footnote. 
(Basilianus cantort, Arrow, 1907, p. 445.) 
Basilianus } Cantori + + convexifrons, Zang, 1909, pp. 178-183. 
Ophrygonius { cantor: + subspp. ¢ convexifrons + + dunsiriensis, Gravely, above, pp. 224-225, 

pl. xi, figs. 22-22a. 

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. QO. convexifrons (Zang) at present seems to be a distinct local race, 
although it differs from the typical form of O. cantori 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. QO. CANTORI, Percheron, s. str. 
K. Himalayas: Darjeeling District—Tista; Sureil; Kurseong; Gopaldhara, 
Rungbong Valley. 


' See Appendix I, p. 316, below. 


1914.] F. H. GravEty: An Account of the Oriental Passalidae. 285 


Sikkim—Gantok. 

Bhutan. 

Dafla Hills—Dikrang Valley. 
Naga Hills: Manipur. 


2. Subsp. CONVEXIFRONS, Zang. 


Assam: Khasi Hills—Shillong ; Teriaghat. 
Burma: Ruby Mines; Sin Im, Bhamo, 6000 ft. 


3. Subsp. dunsiriensis, Gravely. 
) Ni 


Assam: Dunsiri Valley. 


Ophrygonius birmanicus, Gravely. 
? Basilianus cantori [part], Kaup, 1871, p. 57- 
Ophrygonius + birmanicus, Gravely, above, p. 226, text-fig. 3A. 

The specimen from Cambodia which Kaup identified as O. cantort, 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 Passalid fiuna of this peninsula and that of the rest of the continent 
of Asia. 

Localities :— 

Burma: Ruby Mines. 
2 Cambodia. 
?? Malay Peninsula. : 


Ophrygonius singapurae, Gravely. 
>? Basilianus cantori [part], Kaup, 1871, p. 57- 
Ophrygonius + singapurae, Gravely, above, pp. 226-227, text-fig. 3B. 

It is difficult to identify with this species the specimen from the Malay Penin- 
sula which Kaup referred to O. 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 O. 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, pl. I, fig. 4. 


| The type specimen. of Ophrygonius oroletus has elytra with strongly punctured lateral grooves ; 
but the punctures are round, rather than transversely linear as.in the typical O. 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 ovoleius ought perhaps 


to be recognized as a variety. 


286 Memorrs of the Indian Museum. [Vor.. III, 


(Leptaulax orolieus + Aceratus inaequalis, Gemminzer and Harold, 1868, pp. 970 and 972.) 
Leptaulax orolieus, Kaup, 18609, p. 40. 

Basilianus inaequalis, Kaup, 1871, p. 56. 

Basilianus t inaequalis, Stoliczka, 1873, p. 159. 
( Bastlianus inaequalis, Wytsman, 1884, p. 336.) 

(Bastlianus inaequalis, Kuwert, 1891, p. 164.) 

Basilianus f sinkepicus, Kuwert, 1898, pp. 339-340. 

(Basilianus inaequalis, Zang, 1903), pp. 418-9.) 

Ophrygonius } quadrifer, Zauz, 1g04a, pp. 697-700, text-fig. 3. 

Ophryonius } inaequalis, Zang, 19054, p. 192. 

Basilianus sinkepicus, Zang, 1906a, pp. 177-183: 

Ophrygonius + inaequalis, Gravely, above, pp. 227-228, pl. xii, figs. 24-24a. 


Kuwert gives 36 mm. as the length of his Basilianus sinkepicus ; but I have seen 
specimens identified by himself, and found the biggest of them to be only 27 mm. 
long. 

Localities :— 

Malay Peninsula: Penang; Perak; Johore; Singapore. 
Sumatran Islands: Sinkep Island. 
Sumatra: Deli. 
Java. 
Borneo. 
? Philippines. 


The sub-family to which this species belongs is so abundant in the Oriental 
Region right up to Wallace’s Tine and seems to stop so abruptly there, that I am 


inclined to regard Burmeister’s ‘‘ Mollucca”’ record as a misprint for “‘ Malacca ”’ (see 
also Kaup, 1871, p. 56). 


Genus ACERAIUS, Kaup (1868) 1871. 


= Ocythoe [part], Castelnau, 1850; incl. Heterochilus, Kuwert, 1896 — 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 best group character that I have been able to 
find. 

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 criniths, Kuwert, t891, p. 166.) 

Heterochilus + wallacet + crinitus + oculitesselatus, Kuwert, 1898, pp. 334-5: 
Heterochilus t wallacet, Zang, 1905a, p. 167. 

Aceratus t wallacet, Gravely, above, pp. 228-229, pl. xil, figs. 26-26b. 


1914 | F. H. GravELY : An Account of the Oriental Passalidae. 287 


Localities: — 
Malay Peninsula. 
Sumatran Islands: Nias. 
Sumatra—Somgei Lalah in Indragiri. 
Borneo: Sarawak—Kuching. 
British N. Borneo—Mt. Kina-Balu, c. 5000 ft. 


Aceraius perakensis, Kuwert. 
Aceraius perakensis, Kuwert, 1898, p. 308. 


Aceraius + perakensis, Gravely, above, p. 229. 
iocalities:—— 
Malay Peninsula: S. Perak (Telom, 4000 ft.); Taiping 4000~4500 tees atte: 


Hills, 3300-4300 ft.” 
Sumatra: Mt. Singaleng. 


Aceraius moschleri, Kuwert. 


Aceraius moschleri, Kuwert, 1891, p. 163. 
Aceraius maschleri, Kuwert, 1898, p. 344- 
Aceraius + méschleri, Gravely, above, pp. 229-230, pl. xii, fig. 33- 


Localities : — 
Malay Peninsula: Singapore. 
Sumatran Islands: Linga; Sinkep; Sumatra ; Nias. 
Java. 
Aceraius illegalius, Kuwert. 

Aceraius illegalis, Kuwert, 1891, p. 163. 

Aceraius illegalis, Kuwert, 1893, p. 345: 

Aceraius + illegalis, Gravely, above, p. 239, pl xii, figs. 32-324. 


Localities :— 
Malay Peninsula: Johore; Penang. 
Borneo: Kuching ; between Ulu Madahit and Limbang. 


Aceraius laevicollis (Illiger). 


Passalus + laevicollis, Iliger, 1800, p. 103. 
Passalus emarginatus, Weber, 1801, pp. 81-2 
Passalus emarginatus, Fabricius, 1801, pp. 255-9. 
*Passalus laevicollis, Schéuherr, 1806-17, I (3) p- 333: 
* Passalus laevicollis, Dejean, 1837, pp. 194 and 501. 
Passalus emarginatus, Smith, 1852, p. 6. 
Aceraius emarginatus [part], Gemminger and Harold, 1868, p. 972. 
Aceratus + emarginatus [part], Stoliczka, 1873, p. 158. 
(Aceraius emarginatus [part], Wytsman, 1884, p. 336). 
Aceraius Meyeri, Kuwert, 1891, p 163 
Aceraius | meyeri + ¢ pontifex + Fluzonicus, Kuwert, 1898, p. 345: 
Aceraius f laevicollis, Gravely, above, pp. 230-231, pil. xii, figs 27-274. 


BY hill of anything like this altitude in fhe jmimediate 


! De Aunandale informs me that there is n 
neighbourhood of the town of Taiping. 
2 ‘This record is based on specimens in the British Museum. 


288 Memoirs of the Indian Museum. [ Vor. EEE, 


.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 mevert, pontifex, and luzonicus together with all intermediate forms. 


Localities :— 
Malay Peninsula: Perak, Johore. 
Sumatran Islands: Sumatra (Deli, Somgei Lalah, Serdang, Peinan and Solok); 
Nias; Lingga; Sinkep Island ; Billiton. 
Java. 
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, pl. ii, fig. 1. 

Passalus emarginatus, Percheron, 1841, p. 3. 

Passalus grandis, Burmeister, 1847, p. 463. 

Ocythoe emarginatus, Castelnau, 1850, p. 170. 

Passalus grandis, Smith, 1852, p. 5. 

Aceratus grandis, Kaup, 18682, p. 27. 

Aceraius grandis, Kaup, 1868), p. 3. 

(Aceratus grandis, Gemminger and Harold, 1868, p. 972). 

Aceratus f grandis, Stoliczka, 1873, p 158. 

(Aceraius grandis, Wytsman, 1884, p. 336). 

Aceraius Addendus + Rectidens + Hirsutus + Ceylonicus, Kuwert. 1891, p. 163. 

Aceraius addendus + tgrandis + }minutifrons + prosternisulcatus + ceylonicus + thirsutus + trectidens 
+ t magnus + chinensis + molossus, Kuwert, 1898, pp. 342-4. 

Aceratus t+ grandis, Zang, 1904), pp. 184-5. 

Aceraius t hirsutus, Zang, 1905a, p 104. 

Aceraeus } palawanus, Zang, 1905d, pp. 236-238. 

Aceraius + grandis, Gravely, Igt4a, p. 31. 

Aceraius + grandis + var. } rectidens + sub. sp. + hirsutus, Gravely, above, pp. 231-233, pl. xii, 
figs. 28-30. 

Zang (1904b) has shown that A. addendus and minuttfrons are identical with A. 
grandis, at the same time suggesting that A. rectidens, magnus and molussus might 
also prove to be identical with this species. The three last named are probably all 
identical with the var. rectidens described above, 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 §. China, in view of the fact that hirsutus 
is known to occur in Tonkin and Hainan, and so presumably occurs all along the 


1914. | 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. chinensis must provisionally be regarded 
a distinct race. 

Kuwert’s hirsutus and ceylonicus are undoubtedly the same, as Zang has already 
suggested (1g05a, 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. HirsuTuUS, Kuwert. 
E. Himalayas: Darjeeling District -Gopaldhara, Rungbong Valley. 
Bhutan. 
Dafla 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—Sukli, E. side of Dawna Hills, 2100 Et. 


Tavoy. 
Cambodia. 


Tonkin: Mt. Mauson 2—3000 ft. 

Hainan Island. 

Formosa: Kosempo ; Polisha ; ie: hi-ku ; Chip-Chip ; Lake Candidius ; Fuhosho ; 
Hoozan ; Sokutsu iBanctorye ise ; Kankau; Taihorinsho ; Sniclaryor 
Taihorin. 

Philppine Islands: South Palawan. 


Kuwert’s Ceylon record is hardly credible, as Zang has already pointed out 
(19054, p. 104). The genus Aceraius seems to occur neither in Ceylon nor in the 


Indian Peninsula. 
2. A. GRANDIS, Burmeister, s. str.' 
Malay Peninsula. 


Bintang Island. 
Sumatra: Deli. 
Java: Tengger Mountain, Bankalan. 
Borneo: Bandjermasin ; N. Borneo. 


Kuwert records A. prosternisulcatus (=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 eae 2s 


290 Memoirs of the Indian Museum. 


3. Var. RECTIDENS, Kuwert.! 
Malay Peninsula. 


Sumatra: Solok ; Tebing tinggi. 
Java. 
Borneo: Sarawak—Kuching. 


?4. Subsp. CHINENSIS, Kuwert. 
? Southern China. 


Aceraius occulidens, Zang. 
Aceraius f occulidens, Zang, 19054, pp. 190-1. 
Aceratus f occulidens, Gravely, above, p. 234, text-fig. 4A. 
Locality : 
Borneo: Mt. Kina-Balu. 


Acerais laniger, Zang. 
Aceraius }laniger, Zang, 19054, p. IgI-2. 
Aceraius + laniger, Gravely, above, p. 234. 
Locality :— | 
Borneo: Mt. Kina-Balu, c. 5000 ft. 


Aceraius kuwerti, Zang. 
? Aceraius emarginatus [part], Kaup, 1868a, p. 27. 
? Aceraius emarginatus [part], Kaup, 18688, p. 3. 
? Aceraius emarginatus [part], Kaup, 1871, p. 53. 
Aceraius + Kuwerti, Zang, 19034, p. 339. 
Aceraius Kuwertt, Zang, 1905a, p. 189. 
Aceraius + kuwertt, Gravely, above, p. 235, pl. xi, fig. 31. 


[Vor IT, 


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 A. lamigey, 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 A. alutaceosternus. 
Localities :— 
? Java (A. emarginatus, Kaup). 
Borneo: Mount Kina-Balu, c. 5000 ft. 


Aceraius tricornis, Zang. 
Aceratus 7} tricornis, Zang, 19034, p. 339. 
Aceratus tricornis, Zang, 19054, p. 189. 
Aceratus + 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. 


IgI4.| F. H. GraveELy : An Account of the Oriental Passalidae. 291 


Aceraius laevimargo, Zang. 


Passalus emarginatus, Burmeister, 1847, pp. 463-4. 

(Passalus emarginatus, [part], Gemminger and Harold, 1868, p. 982.) 
(Aceratus emarginatus [part], Wytsman, 1884, p. 336. ) 

Aceraius tlaevimargo, Zang, 19054, pp. 244-5. 

Aceraius ¢ laevimargo, Gravely, above, p. 235. 


Localities :— 
Sumatra. 
Borneo: Mt. Kina-Balu, c. 5000 ft. 


Aceraius pilifer (Percheron). 
Passalus pilifer, Percheron, 1835, pp. 23-4, pl. ii, fig. 2. 
Passalus pilifer, 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 + } reticulaticollis + +borneanus + ¢ nicobaricus, Kuwert, 1898, pp. 346-8. 
Aceratus + pilifer, Gravely, above, pp. 235-236, pl. 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 Passalid 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 Aceratus 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. 
Sumatra. 
Java: Tjibodas, ca. 5000 ft.; Preanger, 4—6000 ft. 
Borneo. 


Aceraius alutaceosternus, Kuwert. 


? Aceraius emarginatus [part], Kaup, 18682, p. 2 
? Aceraius emarginatus [part], Kaup, 1868), p. 3. 
? Aceraius emarginatus [part], Kaup, 1871, p. 53. 

Aceraius + emarginatus [part], Stoliczka, 1871, p. 158. 

Aceratus alutaceosternus, Kuwert, 1898, pp. 347-8. 

Aceraius } alutaceosternus, Gravely, above, p. 236, text-fig. 4H, pl. xii, figs. 34-34a. 


Te 


Memotrs of the Indian Museum. [Vor,. III, 


Localities :— 
Malay Peninsula: Penang Hill; Taiping, 4000-5000 ft.; Larut Hills, 3300- 


4300 ft.! 
Aceraius himalayensis, Gravely. 


Aceraius } emarginatus [part], Stoliczka, 1873, p. 158. 
Aceraius } himalayensts, Gravely, above, pp. 236-237, pl. xii, figs. 36-36a. 


Localities :— 
E. Himalayas: Darjeeling District. 
Dafla Hills—Dikrang Valley. 
Sikkim: Pedong. 
Abor country—Sirpo Valley, nr. Renging. 
Assam: Naga Hills. 


Aceraius assamensis, Kuwert. 


Aceraius + emarginatus [part], Stoliczka, 1873, p. 158. 
Aceraius assamensis, Kuwert, 1898, p. 347. 
Aceraius f assamensis, Gravely, above, p. 237, pl. xii, figs. 37--37a. 
Localities :-— 
Assam: Khasi Hills —Cherra Punji. 
Naga Hills—Manipur. 


Aceraius helferi, Kuwert. 


Aceraius Helferi, Kuwert, 1891, p. 163. 
Aceratius pilifer + helferi, Kuwert, 1898, pp. 346-7. 
Aceraius +tavoyanus* + thelferi, Gravely, above, pp. 237-238, pl. xii, figs. 38-39a. 
Localities :— 
Upper Burma: Carin Ghecti, 4000-5000 ft. ; Carin Asciuii Cheba 3500-4000 ft. 
Carin Cheba 3000-3500 {t.; Ruby Mines; Sin Lum, Bhamo 6000 ft. 


Lower Burma: Rangoon. 
Amherst District of Tenasserim.—Dawna Hills between Misty 


Hollow and Thingannyinaung, 900-2500 ft. 
Tavoy District of Tenasserim. 


Siam. 
Tonkin: Mt. Mauson, 2000-3000 ft. 


Aceraius borneanus, Kaup. 


Aceratus ft borneanus, Kaup, 1871, p. 52. 
Aceraius kawptt, Kirsch in Kaup, 1871, p. 52. 
Aceraius percheronti, Kaup, 1871, p. 53. 


' This record is based on 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 Lum and several from the Ruby Mines as well as 


some from various places previously recorded. 


1914. | F. H. GRAvELY: An Account of the Oriental Passalidae. 293 


Aceraius incidens, Kirsch, 1877, p. 28. 

(Aceraius emarginatus [part], Wytsman, 1884, p. 336.) 

(Aceraius Percheronit + Incidens + Kaupt, Kuwert, 1891, p. 164.) 

Aceriaius + percheront + fincidens + + Kaupi + fnanus, Kuwert, 1898, pp. 348-9. 

Aceraius | Kaupit, Zang, 19054, p. 112. 

Aceraius laevicollis, Zang, 1905¢, p. 225. 

Aceraius + borneanus, Gravely, above, pp. 238-239, pl. xii, figs. 25-25) 

The only two differences Kaup was able to find between A. percheronii and 
A. kaupu 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. kaupit, there is no reference to the left, but only to the right, outer 
tubercle being shorter than in A. ¢ncidens, 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. 
Localities :— 
Malay Peninsula: Perak. 
Sumatran Islands: Sumatra—Deli; Indrapura; Tebing tinggi; Peinan; Lang- 
kat. 
Nias—Hili Zobobo. 
Mentawei—Sipora. 
Java: Batavia. : 
Borneo: Kina-Balu, c. 5000 ft.; Sampit; Tandjong; Bandjermasin; Banguey 
Island. 
Phillippines: South Palawan. 


Aceraius minor, Gravely. 
Aceraius + 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. 
Locality :— 
Borneo: Kina-Balu. 
Sub-family MACROLININAE. 


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; Tvberius [part], 
Kuwert r8oI. 


Memotrs of the Indian Museum. [Vor,. III, 


204 
[Macrolinus duivenbodei, Kaup. | 


Macrolinus ¢ Diuivenbodet, Kaup, 18682, p. 19. 
(Macrolinus Duavenbodei, Gemminger and Harold, 1868, p- 970.) 


Macrolinus Duivenbodei, Kaup, 1871, p. 43, pl. iv, fig. 6 
(Macrolinus Duivenbodei, Wytsman, 1884, p. 334.) 
(Macrolinus Duivenbodei, Kuwert, 1891. p. 165.) 
Macrolinus duivenbodei, Kuwert, 1898, p. 184. 


Locality :— 
Celebes: Menado. 

|Macrolinus urus, Heller. | 

(Macrolinus + urus, Heller, 1898, pp. 23-4, pl. i, fig. 26.) 


Locality :— 
Celebes; Mt. Bonthain, 5000-7000 ft. 


Macrolinus nicobaricus, Gravely. 


Basilianus + andamanensis [part], Stoliczka, 1873, pp. 160-r. 


Tiberius Andamanensis [part], Kuwert, 1891, p. 164. 
Macrolinus + nicobaricus, Gravely above, pp 241-242, pl. xiii, figs. 40-40a. 


Localities! :— 
Nicobars. 
?? Burma: Rangoon. 


Macrolinus andamanensis (Stoliczka). 


Basilianus + andamanensis [part], Stoliczka, 1873, pp. 160-1. 

(Bastlianus andamanensis, Wytsman, 1884, p. 336.) 
(Tiberius andamanensis [part], Kuwert, 1891, p. 164.) 
Tiberius andamanensis, Kuwert, 1898, p. 188. 


(Tiberius andamanensis, Zang, 1905a, p. 163.) 
Macrolinus + andamanensis, Gravely, above, p. 242, pl. xiii, figs. 41-41a. 


Localities :— 


Andamans. 
?? Lower Burma: Amherst District—Mcoulmein. 


Macrolinus sikkimensis (Stoliczka). 


Basilianus + sikkimensis, Stoliczka, 1873, pp. 161-2. 

(Basilianus sikkimensis, Wytsman, 1884, p. 336.) 
(Tiberius sikkimensis, Kuwert, 1891, p. 164.) 
Tiberius sikkimensis, Kuwert, 1898, p. 188. 


(Tiberius stkkimensis, Zang, 1go5a, p. 163.) 
Macrolinus + sikkimensis + subsp. + tavoyanus, Gravely, above, pp. 243-244, pl. xiii, figs. 42-42a. 


' There are specimens in Berlin labelled ‘“‘ Andamans, de Roepstorff’’ ; but it is very unlikely, I 


think, that the species really occurs outside the Nicobars. 


1914. | F. H. Gravety: An Account of the Oriental Passalidae. 295 


Localities :— 
1. M. SIKKIMENSIS, Stoliczka, s. str. :— 


E. Himalayas: Darjeeling District—ca. 1500 ft., 2 miles. K. of Punkabari. 
Dafla Hills—Dikrang Valley ; Harmutti (base of hills). 
Abor Country—Kobo, 400 ft. ; Janakmukh, 600 ft.; Rotung, 
1400 ft. 
Assam: Khasi Hills. 
Cachar. 
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 
goo and 2500 ft.; Tavoy. 


Macrolinus crenatipennis, Kuwert. 

Macrolinus + crenatipennis, Kuwert, 1898, p. 185. 

Tiberius crenatipennis, Zang, 1905a, p. 163. 

Macrolinus + crenatipennis, Gravely, above, p. 244. 
Locality :— 

Ceylon. 

Macrolinus rotundifrons, Kaup. 

Macrolinus rotundifrons, [? part], Kaup, 1871, pp. 44-5. 

(Macrolinus rotundifrons, Wystman, 1884, p. 334-) 

(Macrolinus rotundifrons, Kuwert, 1891, p. 165.) 

Macrolinus rotundifrons + singhalensis, Kuwert, 1898, p. 185 

Tibervus trotundifrons + singhalensis, Zang, 19054, p. 163. 

Macrolinus + rotundifrons, Gravely, above, pp. 244-245, pl. xili, fig. 43. 
Localities :— 

Ceylon: Central Province—Kandy ; Peradeniya ; Nalanda ; Patipolla ; Gala- 

gedara. 


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 Waterhouset, Kaup, 1871, pp. 43-4- 

(Macrolinus Waterhouse1, Wytsman, 1884, p. 334-) 
(Macrolinus Waterhousei, Kuwert, 1891, p. 105.) 

Macrolinus Waterhouset, Kuwert, 1898, p. 185. 

(Tiberius Waterhouser, Zang, 1905a, p. 163.) 

Tiberius + waterhousei, Gravely, above, p. 245, pl. xiii, fig. 44. 


2096 Memorrs of the Indian Museum. : {[Vox. III, 


Localities :— 
Ceylon : Ratnapura Distict—Bulutota. 


[Macrolinus sulciperfectus, Kuwert.| 


Macrolinus sulctperfectus, Kuwert, 1891, p. 165. 
Macrolinus + sulctperfectus, Kuwert, 1898, p. 184. 


Locality :— 
South Celebes: Bonthain. 


Macrolinus weberi, Kaup. 
Macrolinus Weberi, Kaup, 1868a, p. IQ. 
(Macrolinus Weberi, Gemminger and Harold, 1868, p. 971.) 
Macrolinus Weberi, Kaup, 1871, p. 44. 
(Macrolinus Weberi, Wytsman, 1884, p. 334.) 
(Macrolinus Weberi, Kuwert, 1891, p. 165.) 
Macrolinus } weberi, Gravely, above, p. 245. 


Locality : 
Philippine Islands. 


Macrolinus latipennis (Percheron). 
Passalus marginepunctatus, Dejean, 1837, p- 194. 
Passalus latipennis, Dup. in Percheron,! 1841, pp. 8-9, pl. Ixxiii, fig. 3. 
Passalus latipennis, Burmeister, 1847, pp. 464-5. 
(Passalus latipennis, Smith, 1852, p. 6.) 
Macrolinus latipennis, Kaup, 1868a, p. 19. 
(Macrolinus latypennis, Gemminger and Harold, 1868, p. 970.) 
Macrolinus latipennis, Kaup, 1871, p. 43. 
(Macrolinus ft latipennis + + weberi, Stoliczka, 1873, p. 155.) 
(Macrolinus latipennis, Wytsman, 1884, Pp: 334-) 
Macrolinus latipennis, Kuwert, 1891, p. 165. 
Macrolinus flatipennis + ab. orbatus + javanus + dissimilis + fraternus + parallelipennis + batesi + 
tweberi, Kuwert, 1898, pp. 186-7. 
Macrolinus + latipennis, Gravely, above, pp. 245-246, pl. 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 case 
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, parallelipennis 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 :— 

Burma. 


' See above, p. 270, footnote. 


1gI4.| F. H. GRAVELY : An Account of the Oriental Passalidae. 


bo 
Ne) 
MN 


Malay Peninsula: Johore. 

Sumatran Islands: Nias—Ombolata. 
Sumatra—Deli. 

Java: Buitenzorg. 

S. E. Borneo. 

Philippine Islands: 5. Palawan. 

Molucca (? = Malacca). 


Subfamily GNAPHALOCNEMINAE. 


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 
Burma. 

Genus PARAPELOPIDES, Zang. 
Parapelopides symmetricus, Zang. 
Parapelopides + symmetricus, Zang, 1904a, pp. 695-7, figs. 1-2. 
Parapelopides symmetricus, Zang, 19054, p. 189. 
Parapelopides + symmetricus, Gravely, above, pp. 2460-247, text-fig. 0A-B. 
Locality :— 
Borneo: Mt. Kina-Balu, c 5000 ft. 


Genus TRAPEZOCHILUS, Zang, 1905. 
= Eriocnemis [part], Kaup, 1871 ; — Phraortes, Kuwert, 18908. 


The genus Trapezochilus occurs in Sumatra, the Malay Peninsula, and the 
extreme south of Burma. 


Trapezochilus dorsalis (Kaup). 

Eviocnemis + dorsalis, Kaup, 1871, p. 41. 

(Eriocnemis dorsalis, Wytsman, 1884, p. 334-) 

(Eviocnemis Dorsalis, Kuwert, 1891, p. 168.) 

Eriocnemis dorsalis + Phraortes generosus + respectabilis, + nobilis, Kuwert, 1898, pp. 323 
& 326. 

Trapezochilus + nobilis + + respectabilis, Gravely, above, pp. 247- -248, text-fig. 5C-E, pl. xiit, 
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 exes 
of the fifth joint bears only a ewelline 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 Prof. 


| Kuwert states that he had only one specimen of T. generosus, ‘but gives both Perak and Sumatra 
as its localities. 


298 Memotrs of the Indian Museum. [Vor. IIT, 


Poulton under whose care they are preserved in the Oxford Museum. ‘These prove 
conclusively that 7. nobilis and T. respectabilis are also indistinguishable from one 
another ; and the type of Eriocnemis dorsalis, Kaup, has proved to belong to same 
species. Consequently none of the names given by Kuwert in this genus can be 
retained. 
Localities :— 
Lower Burma: Tenasserim—Tavoy. 


Malay Peninsula: Perak—Taiping, 4000-5000 ft. ; Telom, 4000 ft. ; Larut Hills, 
4000-4500 ft. 
E. Siamese Malay States—Nawngchik (Bukit Besar, 2500 ft. ; 


and Ban Sai Kau, base of Bukit Besar). 
Sumatra: Deli. 


Genus GNAPHALOCNEMIS,' Heller, rgoo. 
=Ocythoe [part], Castelnau, 1850* 4 Eriocnemis [part], 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 the 
following list. 
Gnaphalocnemis simplex, Gravely. 
Gnaphalocnemis + simplex, Gravely, above, p. 248, text-fig. 6. 
Locality :— 
Malay Peninsula: Perak. 


Gnaphalocnemis burmeisteri (Kaup). 
Eriocnemtis + Burmeistert, Kaup, 1868a, p. 22. 
(Eviocnemts Burmeistert, Gemminger and Harold, 1868, p. 971.) 
Eriocnemis Burmeisteri, Kaup, 1871, pp. 41-2. 
(Eriocnemis Burmeistert, Wytsman, 1884, p. 334.) 
(Eriocnemis Burmeisteri, Kuwert, 1891, p. 168.) 
Eriocnemis burmeisteri + } faberi, Kuwert, 1898, p. 323. 
Gnaphalocnemts + burmeistert, Gravely, above, p. 240, pl. xiii, fig. 49. 

Localities :— 


Sumatra: Somgei Lalah (Indragiri); Soerian Platation (near Solok); Deli; 


Pedong ; Peinan; Battak Mountain ; Kepahiang ; Redjang-Lebong. 

Java. 
Gnaphalocnemis monticulosus (Smith). 

Passalus + monticulosus, Smith, 1852, p. 6, pl. i, fig. 1. 

Eriocnemis tridens, Kaup, 1868a, p. 22. 

(Aceraius monticulosus, Gemminger aud Harold, 1868, p. 972.) 

Eriocnemis monticulosus, Kaup, 1869, pp. 38-9. 


' In addition to the species here mentioned ‘‘ Pelopides’’ gravidus, Kuwert (1891, p. 168; and 
1898, p. 332) from Mindanao perhaps belongs to this genus (see Zang, 19054, p. 316, & 1905¢, p. 227). 

* This name has priority. But it has been so long forgotten in this connection 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. 2 


Eviocnemis monticulosus + tridens, Kaup, 1871, pp. 41-42. 

(Eviocnemis monticulosus, Stoliczka, 1873, P- 155-) 

(Eriocnemis monticulosus, Wytsman, 1884, p 333-) 

(Eviocnemis dispar + monticulosus, Kuwert, 1891, p. 168.) 

Erocnemis + dispar + ignotus + } monticulosus, Kuwert, 1898, p. 324. 

(Eviocnemis monticulosus, Zang, 1903), p. 419.) 

Gnaphalocnemis + monticulosus, Gravely, above, pp. 249-20, pl. xiii, figs. 49-49a. 

Localities :— 

Siam. 


Malay Peninsula : Penang, Tengah Mountain. 


Sumatra: Tandjong Morawa .(Serdang) ; Deli; Solok; Lampung; Medan. 
Borneo : Sarawak—Kuching. 


British N. Borneo—Mt. Kina-Balu. 


Gnaphalocnemis tridens (Wiedemann). 
Passalus + tridens, Wiedeman, 1823, pp. 109-710. 
Passalus laterisculptus, Perty, 1831, p. 37, fig. 2 
Passalus tridens, Percheron, 1835, pp. 24-6, pl. ii, fig. 3 
* Passalus f ortentalis, Dejean, 1837, p. 194. 
Passalus tridens, Percheron, 1841, p. 5. 
Passalus tvidens, Burmeister, 1847, pp. 461-3 
Ocythoe tridens, Castelnau, 1850, II, p. 170. 
(Passalus tridens, Smith, 1852, p. 6.) 
(Passalus tridens, Redtenbacher, 1867, p. 49.) 
Evriocnemis Mniszechi, Kaup, 1868a, p. 22. 
(Eriocnemis Mniszechi + tridens, Geniminger and Harold, 1868, pp. 971-972.) 
Eriocnenis Mniszecht, Kaup, 1871, p 41, pl. iv, figs. 3-34. 
(Eriocnemts  tridens, Stoliczka, 1873, p. 155.) 
(Eriocnemis tridens + Mniszechi, Wytsmau, 1884, p. 334-) 
Eriocnemis Mniszechi + Tridens, + var. Angustior Kuwert, 1891, p. 168. 
Eriocnemis mniszechi + t tridens, + ab. angustior, Kuwert, 1898, pp. 322 and 324. 
Gnaphalocnemts } tridens, Zang, 1904), p. 185. 
Gnaphalocnemis tridens, Gravely, above, p. 250, pl. xiii, figs. 50-50a. 
Localities :— 
Sumatra: Palembang. 


Javanese Islands : Java—Ardjoeno ; Tengger Mountain ; Tjibodas, c. 5000 ft. ; 
Soekaboemi; Preanger; East Java. 
? Moluccas : Amboina.! 


Genus GONATAS, Kaup, 1871. 
= Aceratus [part], Kaup, 1868 Gonatas [part], Kaup, 1871; = Gonatas, 
Rawer, 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. 


! This record rests on “Kuwert’ s authority. 
Region. 


It is the only one yet made outside the Oriental 


300 Memoirs of the Indian Museum. 


[* Gonatas tridentatus, Kuwert.| 
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 schellongt, Kuwert, 1898, p. 314. 
Locality :— 
New Guinea. 


[*Gonatas differens, Kuwert.] 


Gonatas Schellongi var. ? differens, Kuwert, 1891, p. 168. 
Gonatas dtfferens, Kuwert, 1898, p. 314. 


Gonatas differens [incl. G. tibialis, Zang, MS.], Zang, 1905a, p. IIo. 
Localities :— 

Moluccas: Amboina. 

Aru Island. 


New Guinea: Kaiser Wilhelms Land—Finschhafen. 
Bismark Archiplego: New Britain. 


(*Gonatas albertisi, Kuwert.| 


Gonatas Albertisi [Yorkensis Fairm. ?],! Kuwert, 1891, p. 169. 
Gonatas albertist, Kuwert, 18098, p. 314. 


Localities :— 
New Guinea. 


? Bismark Archipelago: New Britain (omitted in Kuwert’s later work). 


Australia : Queensland—Cape York. 


[*Gonatas major, Kuwert.| 
Gonatas Major, Kuwert. 1891, p. 169. 
Gonatus major, Kuwert, 1898, p. 314. 
Localities :— 
Aru Islands. 
New Guinea. 


[*Gonatas novaebritanniae, Kuwert.| 
Gonatas novaebritanniae, Kuwert, 1898, p. 314. 
Gonatas novaebritanmiae, Zang, 18954, pp. 10-1. 


Locality :— 


Bismark Archipelago: New Britain—Kinigunang. 


(Vor. II, 


' Kuwert gives no reference to the place where this name was published, and I have been unable 


to trace it. 


1914. | F. H. GRAVELY : An Account of the Oriental Passalidae. 


Gonatas germari, Kaup. 


Aceraius Germart, Kaup, 18682, p. 30. 

Aceratus Germart, Kaup, 1868), p. 4. 

(Aceraius Geymarit, Gemminger and Harold, 1868, p. 972.) 

Gonatas Germari, Kaup, 1871, p. 51. 

Gonatas + Germart, Stoliczka, 1873, p. 158. 

(Gonatas Germari, Wytsman, 1884, p. 335.) 

Gonatas Germari, Kuwert, 1891, p. 169. 

Gonatas germart + ab. brevis, Kuwert, 1898, p. 315. 

Gonatas + germari, Gravely, above, pp. 250-251, pl. xiii, figs. 47-47¢. 
Localities :— 

awa 

Moluccas: Batchian ; Ceram. 

Ternate. 

Australia: Queensland—lLzard Islands. 


(* Gonatas cetioides, Zang. | 
Gonatas cettoides, Zang, 19054, p. 316. 


Locality :— 
Unknown. 


[*Gonatas peregrinus, Kuwert. | 


Gonatas peregrinus, Kuwert, 1898, 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, pl. cxxxiv, fig. 1. 
? Passalus naviculator [part], Burmeister, 1847, pp. 467-8. 
Aceratus naviculator, Kaup, 1868a, p. 31. 
Aceratus naviculator, Kaup, 1868), pp. 3 & 7. 
Aceratus naviculator, Gemminger and Harold, 1868, p. 972. 
Gonatas naviculatoy, Kaup, 1871, pp. 50-1. 
Gonatas naviculator, Stoliczka, 1873, pp. 157-8. 
Gonatas naviculator, Kirsch, 1877), p. 141. 
Gonatas naviculator, Kuwert, 1891, p. 160. 
Gonatas naviculator, Kuwert, 1898, p. 315. 
(Gonatus naviculator, Arrow, 1907, p. 445.) 


Localities :— 
Java. 
Philippines. 
Moluccas: Batchian; Ceram; Amboina. 
Papuan Islands: New Guinea—Dore. 


301 


302 Memoirs of the Indian Museum. [Vor. III, 


Mafoor. 

Mysore. 

Jobi. 
Australia. 

|*Gonatas altidens, Heller. | 
Gonatas altidens, Heller, 1910, pp. 15-16, pl. I, fig. 13. 
Locality :— 

New Guinea, c. 3000. 


Subfamily LEPTAULACINAE, 
This subfamily occurs throughout the tropical parts of the Indo-Australian region. 
Genus LEPTAULAX, Kaup, 1868. 
incl. Leptaulacides, 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 unable 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, both 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 [ndo-Australian region 
inhabited by Passalidae. 

Leptaulax beccarii, Kuwert. 
Leptaulax beccarit, Kuwert, 1891, p. 189. 
Leptaulax } beccarti, Kuwert, 1898, pp. 294-5. 
Leptaulax + hwmerosus [part], Zang, 19054, p. 112. 
Leptaulax + beccarit, Gravely, above, p. 251. 
Locality :— 


Sumatra. 3 
Leptaulax humerosus, Kuwert. 


Leptaulax humerosus, Kuwert, 1891, p. 189. 

Leptaulax humerosus, Kuwert, 1898, pp. 289 and 294. 

Leptaulax humerosus, Zang, 1904), p. 185. 

Leptaulax + humerosus [part], Zang, 1905a, p. 112. 

Leptaulax humerosus, Gravely, above, pp. 251-252, pl. xiii, fig. 51 


Localities :— 
Sumatran Islands: Sumatra—Battak and Solok Mountains. 
Nias. 
Java: Tengger Mt.; Ardjoeno; and an unrecorded locality at an altitude of 


5000 ft. 
Borneo: Sarawak—Kuching; 10 miles south of Kuching. 


1914. | F. H. Gravety : An Account of the Oriental Passalidae. 303 


Leptaulax anna, Zang. 


Leptaulax + anna, Zang, 19054, p. 316. 
Locality :— 
Sumbawa. 


Leptaulax timoriensis (Percheron). 


Passalus timoriensis, Percheron, 1841, pp. 19-21, pl. Ixxviii, fig. 1. 
(Leptaulax timoriensis [part], Wystman, 1884, p. 332-) 

2? Leptaulax dentatus var. Timorensis, Schaufuss, 1885. pp. I 16-7. 
Leptaulax + glabricollis, Kuwert, 1898, pp. 293-4. 
Leptaulax timortensis, Zang, 1905¢, p. 223. 


Locality :— 
Timor. 


Leptaulax dentatus (Fabricius). 


Passalus dentatus, Fabricius, 1792, p 241. 
Passalus dentatus, Fabricius, 1801, p. 256. 
Passalus dentatus, Weber, 1801, p. 82-3. 
Passalus quadridentatus, 1826, Sturm, p. 182. 
' Passalus dentatus, Percheron, 1835, pp. 66-7, pl. 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. Stnith, 1852, pp. 17 and 20. 
Passalus dentatus, Redtenbacher, 1867, p. 49. 
Passalus dentatus, Kaup, 1868a, p. 14. 
(Leptaulax dentatus, Gemminger and Harold, 1868, p. 969.) 
Leptaulax timoriensts + dentatus, Kaup, 1871, pp. 33-34- 
Leptaulax + dentatus + } bicolor [part] Stoliczka, 1873, p. 155- 
(Leptaulax timoriensts [part] + dentatus, Wystman, 1884, p. 332.) 
Leptaulax dentatus + ? var. Timorensis, Schaufuss, 1885, pp. 186-7. 
Leptaulax ? Ribhei + Saigonicus + ? Dentatus + ? var. Bornensis + 2? var. Indicus + Timortensis + 
Darjeilingi, Kuwert, 1891, pp. 188-190. 
Leptaulax ? abdominibarbatus + malitiosus + submedus + saigonicus + tindicus + ?ab. insignis + 
timoriensis + exterris + + darjeelingt + + celebensis + f dentatus + mixtus +tnterponendus + fF bor- 
nensis + tribbei + ? geminatus, Kuwert, 1898, pp. 293-300. 
Leptaulax + malttiosus, Zang, 1904), p. 185 
Leptaulax + tonkinensis + + humerosus [part], Zang 19054, pp. 102-4 and 112. 
Leptaulax + dentatus, Gravely, 19144, p. 31. 
Leptaulax + dentatus + var. ¢ glabriventris, Gravely, above, pp. 252-255, pl. xiii, fig. 52-52d. 
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. abdomint- 
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 differs 
from L. abdominibarbatus are just those in which the specimens I have identified as 


304 Memoirs of the Indian Museum. [Vou,. III, 


L. ambarbis differ from L. dentatus.' Kuwert’s descriptions of L. ribber and L. gemmi- 
natus leave it uncertain whether the specimens from which they were drawn up 
belong to L. dentatus or to L. novaeguineae, 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. glabriventris may 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. 
Localities :— 
Madras Presidency: Nr. Vizagapatam. 
?W. Himalayas: Mussoorie. 
FE. Himalayas: Darjeeling District. 
Buxa. 
Bhutan. 
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.; Renging to Rotung, 
2600 ft.; Kalek, 3800 ft. 
Mishmi country—Beside the La-ai, a tributary of the Kalem 
River. 
Assam: Kochugarh, Goalpara District; Chaduar; Sibsagar; Dunsiri Valley ; 
Silonbari, base of N. Lakhimpur Hills; E. Cachar. 
Chittagong Hill Tracts: Kapti. 
China. 
Formosa: Kosempo; Chikutoge; Taihorin; Taihorinsho; Hoozan; Fuhosho ; 
Polisha ; Chip-Chip ; Sokutsu, 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; Singapore 
Sumatran Islands: Sumatra; Nias. 
Java: Ardjoeno. 
Borneo: Matang; Kuching. 
Philippines: ? Mindanao. 
Sumbawa’. 


' It appears from the collection of the Deutsches Entomologisches Museum, that Zang applied the 
name abdominibarbatus to a form of L. bicolor which I am unable to recognize as distinct. 
* There are specimens from this island in the Hamburg Museum. 


T914.] PF. H. Gravety: An Account of the Oriental Passalidae. 


305 
Celebes. 
Amboina. 
New Guinea. 
Australia. 


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. novaeguineac; 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 list. Concerning the 
Mussoorie record see above, p. 254. 


Leptaulax cyclotaenius, Kuwert. 
Leptaulax cyclotaenius, Kuwert, 1891, p. 188. 
Leptaulax himalayae + cyclotaenius + + angustifrons, Kuwert 1898, pp. 285-6. 
Leptaulax { cyclotaenius + subsp. ¢ himalayae, Gravely; above, pp. 255-256, pl. xiii, fig. 53. 
This species is very near the last, and may prove to be nothing more than a 
variety of it. aa 
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 L. cyclotaenius, s. str. 
Localities :— 
I. I. CYCLOTAENIUS, Kuwert, s. str. 
Malay Peninsula: Johore ; Perak; Singapore. 
Sumatran Islands : Sinkep. 
Borneo : Sarawak—Kuching ; Braang; Matang Road (near Kuching); Mt. 
Penrissen. 


2. Sub-sp. HIMALAYAE, Kuwert. 
K. Himalayas: Dafla 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 + papua + ambarbis, Kuwert, 1898, pp. 291 and 293. 

Leptaulax macassariensis subsp. + anibarbis, Gravely, above, p. 256, pl. 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. 


306 Memotrs of the Indian Museum. (Vor. III, 


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. I,, MACASSARIENSIS, Kuwert, s. str. 
South Celebes : Macassar. 
New Guinea. 


2. subsp. ANIBARBIs, Kuwert. 


Borneo : Sarawak. 


Leptaulax anipunctus (Zang). 
Leptaulacides t anipunctus, Zang, 1g905a, pp. 234-5. 
Leptaulax + anipunctus, Gravely, above, pp. 256-257, pl. xiii, fig. 55. 
Localities :— 
Assam. 
Burmo-Chinese Frontier: Sansi Gorge, 6000—8o000 ft. 
Upper Burma: Catchin Cauri.' 


[* Leptaulax novaeguineae, Kuwert. | 
Leptaulax Novaeguineae + ? conseguens, Kuwert, 1891, pp. 188-9. 
Leptaulax novaeguineae + ab. morator + hansemanni +? consequens, 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 
ribbet and geminatus differ in any constant character either from this species or from 
L. dentatus. But the two last are much larger than any kuown specimen of 
L. novaeguineae 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. bicolor; 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 dentatus and bicolor. 
Localities :— 
? Batchian (L. consequens). 
New Guinea. 


' There is a specimen from this locality in the Kgl. Zool. Museum in Berlin. 


TOT 4. F. H. Gravety: An Account of the Oriental Passalidae. 07 
OT 4. | 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). | 
Trichostigmus glaber, Kirsch, 1877), pp. 139-140. 
(Trichostigmus glaber, Wytsman, 1884, p. 331.) 
Leptaulax glaber, Kuwert, 1891, p. 188. 
Leptaulax glaber, Kuwert, 1898, p. 292. 
Leptaulacides } glaber, Zang, 1905¢, pp. 227-8. 
Leptaulacides } glaber + | pulchellus, Arrow, 1907, pp. 406-7. 


L. pulchellus from New Guinea is apparently a distinct variety, for Arrow was 
able to compare his type with several specimens of the typical glaber. 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 trom Jobi 
is much to be desired. 


Localities :— 
Molluccas: Batchian. 
Papuan Islands: New Guinea; Jobi. 


Leptaulax bicolor (Fabricius). 


Passalus bicolor, Fabricius, 1801, p. 256. 
* Passalus bicolor, Schénherr, 1896-17. 
Passalus bicolor, Percheron, 1835. pp. 69-70; pl. v, fig. 3. 
* Passalus bicolor + innocuus, Dejean, 1837, p. 195. 
Passalus bicolor, Percheron, 1841, p. 21. 
Passalus vicinus, Hope in Percheron, 1844, p. 11. 
Passalus bicoloy + vicinus, Burmeister, 1847, pp. 478-9. 
(Passalus bicolor + vicinus, Smith, 1852, pp. 17 and 20.) 
Leptaulax Eschscholtzi + bicolor, Kaup, 1868a, pp. 14 and 16. 
Leptaulax bicolor [part] + Eschscholtzt, Gemminger and Harold, 1868, p. 969. 
Leptaulax Eschscholtzi + bicolor, Kaup, 1871, pp. 323- 
Leptaulax + bicolor [part], Stoliczka, 1873, p. 155- 
Leptaulax bicolor, Kirsch, 1877), p. 140. 
(Leptaulax Eschscholtzi + bicolor [part], Wystman, 1884, p. 332.) 
Leptaulax bicolor, Schaufuss, 1885, p. 186. 
Leptaulax Manillae + Malaccae + ? Consequens + + Batchianae + Bicolor + }Cicatrosus + Abdomint- 
sculptus + Differentispina + Incipiens + Eschscholtzii + Aurivillit + Separandus,' + vats. 
Medius and Maxillonotus, Kuwert, 1891, pp. 188-190. 


1 Since drawing up this synonymy I have seen a specimen determined by Kuwert as L. separandus 
and found it to be L. dentatus, not L. bicolor. As this is the only instance in which I have not found 


308 Memoirs of the Indian Museum. (Vou. III, 


Leptaulax + malaccae + manillae + dindigalensis + | batchianae + ¢ roepstorfi [part] + } cicatrosus + 
? consequens + + subsequens + ¢ bicolor [part] + insipiens + sumatrae + }calcuttae + abdomint- 
sculptus + | geminus + sequens +  niae + ¢ eschscholtzi + f aurivillit + tenasserimensts + evidens + 
+ differentispina + sebarandus'! + maxillonotus + medius + ab. divaricatus, Kuwert, 1898, 
pp. 286-300. 

Leptaulacides t rugulosus + t Fruhstorferi + ¢ Eschscholtzi + } barbicauda + | Andamanarum + tpala- 
wanicus + } anaulax + + Nietneri, Zang, 19054, pp. 100-2, 106-9, 164-7, 232-4, 235-6, 246-7. 

Leptaulacides vicinus, Arrow, 1907, p. 445. 

Leptaulacides + papauanus + } analis, Zang, 1906). 

Leptaulax f bicolor, Gravely, 19144, p. 31. 

Leptaulax + bicolor + var. ¢ vicinus, Gravely, above, pp. 257-259, pl. xiii, figs. 56-56a. 

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. dentatus 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, isa 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. bicolor, 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 able 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. roepstorfi, to specimens 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 probable that it ought 
to go. ‘ 

'! See footnote on previous page. 

2 It should, however, be pointed out here that this character is not equally strongly developed 
in ail 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. maxillonolus aud medius, names which appear in the above list as synonyms of L. bicolor. 


Tord. | F. 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—Janahmukh, 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. 
Andamans. 


Nicobars. 

Formosa: Kosempo; Polisha. 

Siam. 

Tonkin : Than-Moi ; Mt. Mauson, 2000~3000 ft. 

Cambodia. 

Malay Peninsula. 

Sumatran Islands: Sumatra ; Sinkep ; Nias. 

Java, 4,000 ft. 

Borneo : Sarawak—Kuching ; Matang Road; 4th mile, Rock Road ; Lingga 

(on the Batang Lupar River). 

British N. Borneo.—Mt. Kina-Balu. 

Philippines : Manilla; S. Palawan. 

S. Celebes. 

Moluccas: Batchian. 

Papuan Islands ; New Guinea ; Jobi. 

Australia. 


I have omitted from the above list of localities Kuwert’s ‘‘ ? Old Calabar,’’ as it 
is 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. 
Leptaulax { planus [part], Stoliczka, 1873, p. 155. 
Leptaulax } roepstorfi [part] + planicollis, Kuwert, 1898, pp. 288-290. 
Leptaulax { roepstorfi, Gravely, above, p. 260, pl. xiii, fig. 57. 


Localities :— 
? Bengal: Calcutta. 
E. Himalayas: Abor County—Yambung, 1100 ft. ; Rotung, 1400 ft. 
Lower Burma: Tenasserim—Misty Hollow, W. side of Dawna Hills, Amherst 
District, c: 2200 ft. Tavoy-.. 


Andamans. 


310 Memoirs of the Indian Museum. [ Vou. III, 


Leptaulax planus (lIlliger). 
Passalus planus, Iliger, 1800, p. 104. 
(Leptaulax bicolor [part], Gemminger and Harold, 1868, p. 969.) 
Leptaulax + planus [part], Stoliczka, 1873, p. 155. 
(Leptaulax bicolor [part], Wytsman, 1884, p. 332.) 
(Leptaulax planus [part], Kuwert, 1891, p. 188.) 
Leptaulax + planus, Kuwert, 1898, p. 285. 
Leptaulacides + planus, Zang, 1904), p. 185. 
Leptaulax + planus, Gravely, above, pp. 260-261, pl. xiii, fig. 58. 
Localities :— 
Lower Burma : Tenasserim—Tavoy. 
Malay Peninsula ; Johore. 
Sumatran Islands: N. E. Sumatra—Serdang. 


fave Sinkep Island. 


Borneo : Sarawak—Kuching ; Mujang ; Sadong; Matang Road; Paku. 


Genus TRICHOSTIGMUS, Kaup. 
This genus appears to be restricted to Celebes and the Philippines. 


Trichostigmus thoreyi, Kaup. 
Leptaulax Thoreyi, Kaup, 1868a, pp. 13-14. 
(Leptaulax Thoreyi, Gemminger and Harold, 1868, p. 970.) 
Trichostigmus Thoreyi, Kaup, 1871, p. 31. 
(Trichostigmus Thoreyt, 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. 187. 
Trichostigmus Ursulus, Kuwert, 1891, p. 187. 
Trichostigmus ursulus, Kuwert, 1808, p. 283. 
Locality :— 
South Celebes (according to the labels on the type specimens; Schaufuss gives 
no definite record). 


ZOOGEOGRAPHICAL RESULTS. 


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. 


1914. | F. H. GRAVELY: An Account of 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 Ophrygonius 
cantort 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 Awlacocyclus 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- 
drocaulus 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 dis- 
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 Leptaulacinae 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, excépt the Andamans and Nicobars. 
The distribution of the different genera belonging to it is peculiar. In Ceylon we 
find the somewhat rare symmetrical species Episphenus mooret, 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 
Episphenus comptoni. In the Indian Peninsula we find two almost equally common 
but more markedly asymmetrical species of Episphenus, E. indicus, and E. neel- 
gherriensis. ‘Tue former has the anterior angles of the head prominent, and resembles 
Epis ohenus comp ont 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, Tiberioides, 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 


oD See Bice belon: p. 330. 


312 Memotrs of the Indian Museum. [Vou.-TLy; 


Islands we find the genus Aceraius, of which one species, A. grandis, resembles Epis- 
phenus comptont and Episphenus 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. Finally, in the 
greater part, if not the whole of this area, we find the genus Ophrygonius, apparently 
less common, and nearly related to the genus Aceratus. 

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 A cerazinae, 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 may 
be inferred that Episphenus indicus, which differs from FE. neelgherriensis in having 
the anterior angles of the head prominent, in its very variable size, and in its 
somewhat more markedly 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. neelgherriensts. 

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 highly 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 angtes 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 specializa- 
tion increasing in response to a fresh environment, is precluded by the occurrence of species allied to 
Episphenus moorei in Australia (see below, p. 315). 


1914. | F. H. GraveLy: An 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 the Gangetic 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 Ophrygonius and 
Aceraius, which inhabit the Oriental Region west of the Bay of Bengal, Aceraius 
grandis has alone developed all the characteristics of adominant 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 vectidens 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 
Ophrygonius, 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 genus 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 specialized species found in 
different parts of the Oriental Region, to find that the genus A cevaius 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. Aceravus 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 allied rather to that of 
the Archipelago, and from which two representatives of this fauna—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 ofthe islands east of 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.' 


|G. germari and naviculator 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. 


314 Memoirs of the Indian Museum. [Vou. III, 


E.HIMAL AYAS 
BORNEO 


CELEBES TO NEW CALEDONIA 


INDIAN 


= 151 
PENINSULA el 


AY AI 
\ ST lh ( "a ) 


— - } ; r eran 
PROTOMOCOELUS: GNAPHALOCNEMIS: © ite et om. \ ' GONATAS, 


/ACERAIUS GRANDIS. 
TATIUS OMEGARIUSieun! 


EP1SPHENUS INDICUS. 


fred | 


H\ Cf | 
| TRAPEZOCHILUS. 


fees 7 
‘ACERAWUSior nen srr 


PLESTHENUS.’ 
A 


a 


fr, | s\ 
ie! 


EPISPHENUS. 
NEELGHERRIENSIS. 


\, “ 7 Q\—A- 2 0 
(Rao < = 
| ANALACHES| f PELOPIDES) 
| ceTesus PARAPELOPIDES 
ASYMMETRICAL SPP. 


MALAYSIA 


(| (A, — 
Ws 
ANALACH - 
| NALAGHES. HYPERPLESTHENUS. 
SYMMETRICAL SPP. 


pace 


Copuavconius. 


| EPISPHENUS | 
COMPTON}. 


V Ao 


(| es, Z 
x a> — 
AURELIUS. KAUPIOLUS.  vagienus. | 


[ae pao Ee Ce ee 
TIBERIOIDES. \ 


| PHAROCHILUS. | 
| MASTOCHILUS. | 
lEPISPHENOIDES 


CEYLON || AUSTRALIA 


pe 
EPISPHENUS MOORE!) 


D Bagchi, del, 


TEXT-FIGURE 7. 


Diagram illustrating the evolution of the different types of asymmetry found in the Aceraiinae and 
Gnaphalocneminae, 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:—(1) a species of E-pisphenoides 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 Gnaphalocnemis 
which has been recorded from Amboina; and (4) the genus Plesthenus, whose anomalous distribution (in 
Australia and Celebes) calls for further study. The genus Kaupiolotdes is in some ways intermediate 
between the Australian genera and Cetejus-Analaches 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 Protomocoelus than is here indi- 
cated. The particular structures whose various modifications the diagram is designed to illustrate are 
shown by thickened lines. The figure of Plesthenus is from Kaup; those of Pelopides and Tatius are 
from Kuwert; that of Aurelius is partly from Kuwert, partly from memory; and that of the sym- 
metrical species of Cefejus-Analaches entirely from memory. 


To14. | F. H. GRAVELY: An Account of the Oriental Passalidae. ahi 


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'—Episphenus moorei 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: (1) a single species—pectinigera, Heller—from New Guinea, of the other- 
wise purely Australian genus Episphenoides; and, (2) a few species of the closely allied 
genera Cefeyus and Analaches. All the remaining extra-Australian symmetrical species 
are specialized in other ways. For instance, in the whole of the Hypferplesthenus 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 Awrelius, 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 inthe 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 any species of the genus 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. exxiii-cxxv 


316 Memoirs of the Indian Museum. [Vor. III, 


the more highly specialized species have been able to establish themselves on all’ the 
islands on one side or other of ‘‘ Walllace’s Line’’ and even in a few instances (Gonatas 
spp. and perhaps Guaphalocnemis 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. 


7. APPENDIX I.—A REVISED CLASSIFICATION OF THE ACERATINAE. 


In the key to the genera of Aceraiinae given on pp. 196-7 of the present 
paper the genera Basilianus, Ophrygonius, 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 
group. . 

Not only do the genera Bastlianus (old sense), Ophrygonius and Aceraius grade 
one 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 mooret, 
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 
head. : 

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 cantort, a species in which the head is highly asymmetrical, but the 
mandibles 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 gap 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 


1914. | F. H. GrRAvELY: 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 Ophrygonius. 

The monospecific genus Chilomazus is almost as closely related to the monospecific 
genus Episphenus on the one hand, and to the genus Basiianus 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 Ceteyws &nd Analaches. 

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 Ofhrygonius 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 aequidens 
and A. minor are allied. 

Definitions, based on the structyre 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 Aceratus wallacet 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 :— 


Inner tubercles separated by a space 4—3 as long as that 


separating outer tubercles .. er re ..  Tiberioides, Gravely, 
Inner tubercles separated by a space }—1I times as long as that pp. 215 & 280. 
aoe outer tubercles . ‘ ae ae ahs se hte 5 
Left outer tubercle acute, site 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 

tooth smaller than left in the more highly asymmetricalspecies. Episphenus, Kaup, 

Left outer tubercle always larger than right, directed more or pp. 217 & 281. 
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 


318 Memoirs of the Indian Museum. [Vou. III, 


( A little hair present on elytra at shoulders only; dentition 
always complete, left anterior lower tooth rarely enlarged. Ophrygonius, 


By Elytra hairy at sides also; right lowest terminal and anterior Zang, pp. 224 & 284. 
"| lower teeth almost always rudimentary; left anterior lower 

tooth almost always enlarged ae Sas .. Aceraius, Kaup, 
( pp. 228 & 286. 


8. APPENDIX II.—KEYS FOR THE DETERMINATION OF SPECIES 
OF ORIENTAL PASSALIDAE. 


In most of the following keys Oriemtal species only are dealt with. Of the 
seven genera occurring in both the Oriental and Australian Regions five (Comacupes, 
Macrolinus, Gnaphalocnemis [?], Leptaulax 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. 


Mesosternum strongly punctured all over, abdominal sterna 
with at least a few hair-bearing punctures in fresh specimens. . ye ae 2 
2 Mesosternum unpunctured except at sides, abdomen unpunc- 
tured and hairless ie Be és . C. foveicollis, p. 267. 
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 axe ae 3: 
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 


above 20 J nee 5: 
( Central éuberale pe tenses (atnreeee as in anes .. C. basalis, p. 267. 
\ Central tubercle not pedunculate se A a; he aA 4 
Central tubercle broad and strongly excavate above .. C. cavicornis, pp. 204 & 268. 
Central tubercle narrow, at most longitudinally grooved above C. stoliczkae, pp. 206 & 268. 
Whole anterior end of central tubercle raised well above supra- 
orbital ridges, not truncate or concave in front .. .. C. masont, pp. 207 & 268. 
5/ Central tubercle less elevated, truncate or concave in front, 
| very variable .. Ap Be aie .. C.cyclindraceus, pp. 207 & 
269. 


' 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 Passalidae. 319 


Genus Taenioers, Kaup. 


( Upper surface of central tubercle about twice as long as broad, 

; flat, punctured, bordered by very distinct horse-shoe-shaped 

/ ridge which is open in front : 
Upper surface of central tubercle releaves onder as a rule, 
less flattened, unpunctured, marginal ridge often indistinct or 

pee across middie-line behind as well as in front 

{ Anterior tibiae very broad. 

| Anterior tibiae slenderer 


Anterior margin of canthus meeting side of head a considerable 
distance behind anterior angle; external angle of canthus 
obtuse : A : 0 

Anterior margin of oadthius meeting Side of head a very short 
distance behind anterior angle; external angle of canthus 
sharper 


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 : dic ae 
Apical portion of central tubercle more é strongly bent over for- 
wards, never upwardly directed, at least the extreme distal 
portion of the upper margin bent a little downwards 
Dorsal grooves of elytra unpunctured .. : 
Dorsal grooves of elytra very distinctly binceaneay as in fhe 
the two following species es oe 
Central tubercle as seen from above not or seeeie broader in 
| front than behind : ; 
Central tubercle seen from above mnie broader in front tian 
eee ‘ : & 
(see portion of central fibercie very short, not bent down- 
wards as a whole ant fie be 
Apical portion of central tubercle much longer, very slightly 
te distinctly bent downwards as a whole 


Genus Ceracupes, Kaup. 


Apex of horn formed by fusion of central tubercle with ante- 
I, rior margin of head bifid 
Apex of this horn acute oi 
( Upper surface of this horn roughly r panatial sided dufguationte. 
\ Uper surface of this horn much broader in front than feuinidl 


Genus Cylindrocaulus, Fairmaire. 


Only one species, C. bucerus, p. 270. 


Genus Auritulus, Zang. 
Only one species, A. patalis, p. 270. 


T. bicanthatus, pp. 208 & 270. 


T. platypus, p. 270. 


T. pygmaeus, pp. 209 & 271. 


T. bicuspis, pp. 210 & 272 


NO 


Oo 


A. rosenbergti, p. 274. 


A. dilatus, p. 276. 


A. andrewst, p. 211 & 275. 
A. aruensis, p. 277. 


A. parryi, p. 276. 


i) 


C. austent, pp. 212 & 278. 
C. arrowt, p. 278. 
C. fronticornis pp. 212 & 277. 


Memoirs of the Indian Museum. 


Genus Pleurarius, Kaup. 


[Vor IIT, 


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. brachyphyllus , from India (above, pp. 213 and 280), are 
the only species of the genus that have yet been described. 


Genus Tiberioides, Gravely. 


{ Lateral grooves of elytra narrow, normally punctured 

, \ Lateral groves of elytra broad, their punctures transversely linear 
Mentum with a protuberance on anterior margin, somewha* as 

i Episphenus comptoni, and a strong transverse ridge a little 
further back - ne es 

| Mentum without any such ridge or protuberance . . 


Genus Episphenus, Kaup. 


Anterior margin of head symmetrical; anterior margin of men- 
;/ tum not depressed or grooved oe 
Anterior margin of head more or less asymmetrical ; anterior 
eee of mentum more or less depressed or grooved 
Anterior margin of head not very strongly asymmetrical as a 
| rule; anterior margin of mentum strongly grooved on either 
2 side of a strong median tubercle (occasionally paired) 
Anterior margin of head strongly asymmetrical; mentum 
ee any strongly marked tubercle 
{ Anterior angles of head not prominent 


3 
l Anterior angles of head more or less prominent 


Genus Ophrygonius, Zang. 

35 At least five well developed and pubescent antennal lamellae 
*) Antennal lamellae very short, only four of them pubescent 

Anterior lower tooth of both mandibles normal, conical and 
‘| acute an 

Anterior lower tooth of both mandibles very ‘ween more 
Vee flattened dorso-ventrally ae : 

Left outer tubercle moderately stout, directed more or less 
,) forwards 5 : 
Left outer tubercle very aieaaen siete sfunele inwards 


Genus Aceraius, Kaup. 


T. kuwerti, pp. 215 & 280. 


T. borealis, p. 281. 
T. austent, pp. 216 & 28r. 


E. moorei, pp. 217 and 281. 


iS} 


E. comptoni, pp. 218 & 28r. 


Ww 


E. neelgherriensis, pp. 222 
283. 
E. indicus, pp. 220 & 282. 


O. inaequalis, pp. 227 & 285. 
3: 
O. singapurae, pp. 226 & 285. 


O. cantort, pp. 224 & 284. 
O. birmanicus, pp. 226 & 285. 


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 very 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 


1914. | F. H. GraveLy: An Account of the Oriental Passalidae. 321 


have seen in each species has been carefully 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 We 56 o 
Upper tooth of both mandibles eee lowest terminal tooth 
of both mandibles normal; anterior lower tooth of right man- 
dible well developed, that of left side not always abnormally 
large 50 : He ae 18. 
No convexity of upper margin of left A PCrNES 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, 27a, 28), 29) 3¢ 
Upper tooth of left mandible always simple, set ina hollow in 
front of a convexity of the upper margin from which it is dis- 
tinctly separated at base (see fig. 25a, pl. xii; & 4C, p. 234) ig avs T4. 
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. 26a, 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. 280) 56 oC on se 4. 
Anterior angles of the head obtuse, never prolonged forwards ; 
apex of upper tooth of left mandible always more or less dis- 
tinetly bifid in unworn specimens (see especially fig. 31) a0 aie 50 To. 
Right outer tubercle simple, elongate, acute (figs. 26, 32, 
. | 33)5 anterior angles of head truncate or at most moderately 
prolonged oe = Se 5: 


~ ———— ee 
Oo bo 


4( Anterior angles of head at see muaersect eratdaged’ 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 oc - 36 8. 
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 oD .. A. wallacet, pp. 228 & 286. 
| Proximal lamellae of antennae shorter; inner margin of oS 
outer tubercle oblique throughout Se : oes = 6. 
/ Apex of right outer tubercle truncate or imperfectly bifid in 
profile, the suture distinctly bent downwards; anterior angles 
of head not prominent; mentum punctured sparsely or not at 
all in middle (left upper tooth very variable) SC .. A. perakensis, pp. 229 & 287. 
| Apex of right outer tubercle simpiy pointed in profile, suture 
not bent downwards; anterior angles of head more or less dis- 
tinctly prominent; mentum more or less strongly punctured 


throughout .. ; q : os ae a 7: 
Anterior angles of head not very rominent tip of right outer 
tubercle as in A. pevakensis, not bent outwards (fig. 33) .. A. miéschleri, pp 229 & 287. 


peel angles of head more prominent; tip of right outer 
tubercle bent somewhat abruptly outwards Bn .. A. illegalis, pp. 230 & 287. 


322 Memoirs of the Indian Museum. [Vor III, 


Canthus without any upwardly directed tubercle .. Se ae S3 =O; 
A stout erect tubercle arising from dorsal surface of canthus 
") immediately in front of eye Hs .. A. occulidens, pp. 234 & 290. 


At most 34 mm. long; anterior angles of head saree more 
prominent than in A. illegalis, somewhat variable; apparently 
confined to the Malay Peninsula and Archipelago, and the 
/ Philippines as =. e .. A, laevicollis, pp. 230 & 287. 
At least 33 mm, long, specimens ren the Malay Peninsula 
and Archipelago always much bigger than this; anterior angles 
of head often strongly produced forwards, especially in small 
specimens atc ot ac : -. A. grandis*, pp. 231 & 288. 
Specimens at least 43 mm. long; right outer abeiele sdunied 
: or obsolete except in one species always over 47 mm. long .. ; x va 
\ Specimens at most 40 mm. long; right outer tubercle always 
more or less distinct and pointed ae ae Be + fu 13. 
/Right outer tubercle distinct; ridge between lateral and inter- 
aaa areas of metasternum obtuse, rough; teuth rib of 
elytra punctured and hairy anteriorly .. ae .. A. laniger, pp 234 & 290. 
Right outer tubercle more or less obsolete; ridge between 
lateral and intermediate areas of metasternum acute, polished ; 


10 


tenth rib of elytra usually hairless and unpunctured sie ang Se me 
Supra-orbital ridges and inner tubercles normal; right outer 
tubercle not completely absent Se .. A. kuwerti, pp. 235 & 290. 


) Supra-orbital ridges with the apical angle very ational deve- 
\ loped, and situated on inner side of, rather than behind, the 
anterior truncation; left inner tubercle situated on inner side 
of base of left outer tubercle; right outer tubercle missing .. A. dricornis, pp. 235 & 290. 


' These distinctions are much more definite in reality than might be a7 | 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 A. Jaevicollis 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. hirsutus, Kuwert. 

B. Southern race, confined to the Malay Peninsula and Sunda 
Islands. Seventh rib of elytra never punctured; hair very 
variable in length, usualiy longer than in the northern 
race. Occurring in two forms:— 

a. Anterior angle of left side of head at least moderately 


prolonged and curved inwards fo . A. grandis, Burmeister, s. str. 
Anterior angle of left side of head at most bderately 
prolonged, directed outwards or forwards .. var. vectidens, Kuwert. 
?C. Chinese race with elytra like those of the southern race .. sub-sp. chinensts, Kuwert 


[Mr. Arrow tells me that Burmeister’s type resembles rectidens; in this case the form here and 
throughout this paper called grandis, s. str., becomes var. addendus, Kuwert.] 


1914. | F. H. GRAVELY: An Account of the Oriental Passalidae. 323 


At least 34 mm. long; very variable in structure; right outer 
tubercle simply pointed in profile in unworn specimens; scars 
Bs on mesosternum almost always distinct a: .. A. laevimargo,' pp. 238 & 291. 
At most 32 mm. long; less variable; right outer tubercle 
slightly truncate in profile in perfectly unworn specimens; 


scars on mesosternum obsolete ac ae . A. pilifer, pp. 235 & 291. 
fee part of tenth and whole of eighth ribs of elytra un- 
+} pons ae 09 50 ae oe 15. 


iGeeeatn to tenth ribs of ne tra (inclusive)punctured throughout A. borneanus, pp. 238 8& 292. 
onvexity of upper margin of left mandible very high and 
strongly curved—usually much higher than in auy other 
species; left outer tubercle rather slender; right outer tuber- 
15/ cle more or less truncate in profile .. .. A.alutaceosternus pp. 236 & 291. 
ee of upper margin of left mandible fitodetately high 
and strongly curved; left outer tubercle somewhat stouter; 
right outer tubercle variable 3h 
Free portion of left outer tubercle distinctly longer than Broads 
iF right outer tubercle often very broadly bifid in profile .. A. helfert, pp. 238 & 292. 
eae portion of left outer tubercle more or less square; right 
outer tubercle not very broadly bifid in profile .. ae se bis ity 
=) Right outer tubercle distinctly bifid in profile .. .. A. assamensis, pp. 237 & 292. 
7) Right outer tubercle at most imperfectly bifid in pro‘le .. A. himalayensis, pp. 236 & 292. 
‘Auterior 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 


10. 


10 


r8¢ elytra unpunctured ac yee tes 3¢ A. minor, pp. 240 & 293. 
Anterior lower tooth of left mandible scarcely larger than that 
of right; s:venth to tenth ribs of elytra (inclusive) somewhat 


sparsely punctured throughout i 5 A. aequidens, pp. 240 & 293. 


Genus Macrolinus, Kaup. 


Inner tubercles, and anterior part or whole of frontal ridges, 


ost 3c be 2c 3t me ZF 
) Frontal ridges complete and well developed, inner tubercles 
( more or less distinct as vs ne 50 * a a 
Apex of central tubercle approximately rectangular in profile, 
{aivected upwards, not overhanging .. - . WM. urus, p. 294. 
Apex of central tubercle acute, directed ferred somentint 
‘ae ae +s ee -. M. duivenbodet, p. 294. 
Ridge joining iuner eahercies separated from anterior margin 
e head throughout its whole length by a more or less concave 
3 surface AG a8 ae 4. 
ae joining inner uteveles closely approximated to anterior 
margin of head either in middle or throughout 6. 


( Lateral grooves of elytra narrow, punctures normal pas ee fe Gy 
\ Lateral grooves of elytra broad, punctures transversely linear M. sikktmensis, pp. 243 & 294. 


' Or the slightly larger A. pevakensis (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. /aevimargo belongs. 


324 Memoirs of the Indian Museum. 

pee tubercles slender in profile, truncate; ridge joining 
inner tubercles concave 

Outer tubercles stouter and distinctly bifid in orks ariel 

joining inner tubercles straight 3A re 


Inner tubercles situated distinctly behind anterior margin of 
head AC ts "a 5 A 
Inner tubercles situated on (or vertically above) auterior margin 
of head 


Grooves of elytra coarsely punctured, lateral grooves almost 
f broad as intervening ridges, their punctures very coarse 
7\ indeed ate Bs ws vs 

i ooves of elytra less cbaisely punctured, lateral grooves 

much narrower than intervening ridges 

Ridge between inner tubercles convex, evenly curved through- 
eee anterior angles of pronotum strongly punctured 

Ridge between inner tubercles straight throughout almost its 
ieee length; anterior angles of pronotum unpunctured 

Third lamella of antennae distinctly shorter than fourth; 

median groove of pronotum distinct, complete .. : 
9) Third lamella of antennae not distinctly shorter than Fourth: 
median groove of pronotum obsolete as oe 
Tip of second lamella not reaching line joining tips of first and 
ae when antenna is furled AC 
Tips of all six lamellae arranged in a straight line moan antenna 
is furled fe “hs Se He 


Genus Parapelopides, Zang. 


Only one species, P. symmetricus, pp. 246 and 297. 


Genus Trapezochilus, Zang. 


Only one species, T. dorsalis, pp. 247 and 297. 


(Vor. III, 


M. nicobaricus, pp. 241 & 294. 


M. andamanensis,pp. 242&294. 


M.crenatipennis, pp.244 & 295. 
8. 
M. rotuntifrons, pp. 244 &295. 
M. waterhouset, pp. 245 &295. 
M. sulciperfectus, p. 296. 
10. 
M. webert, pp. 245 & 296. 


M. latipennis, pp. 245 & 206. 


Genus Gnaphalocnemis, Heller. 


Left outer tubercle consisting of a single, somewhat slender, 
obliquely truncate process; right outer tubercle consisting of 
) asimilar but broader and slightly bifid inner process, together 
\ with smaller pointed outer and middle processes oe 
Both tubercles consisting of three denticles more or less fused 
together, the middle one sometimes obsolete 


Lateral grooves of elytra all narrow, simply punctured 
Grooves 5-7 of elytra more or less broad; each containing a 
polished flattened band which is marked by a single row of 
punctures, and defined on either side by a more less distinct 
roughened line, with which the punctures may be to some 
extent confluent 


ce simplex, pp. 248 & 298. 


2 
=. 


G. burmeistert, pp. 249 & 298. 


ue 


1914. | F. H. GRAVELY: An Account of the Oriental Passalidae. 325 


Grooves 5-7 of elytra somewhat variable in width; the poste- 
{io part of groove 8 rarely wider than the anterior part, 


3 never as wide as groove 7 .. oA . .. G.monticulosus, pp. 249 & 208. 
| Grooves 5-7 of elytra always very broad; the posterior part 
of groove § like them ays oe BE .. -G. tridens, pp. 250 & 299. 


It is possible that ‘‘ Pelopides”’ gravidus, Kuwert, whose position is uncertain, 
may belong to this genus. It differs 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. ; 


Left mandible less clougated and more curved; lamellae of 


antennae longer and slenderer si ee .. G. germari, pp. 250 & 301. 
I : : ; 

Left mandible longer and less curved; lamellae of antennae 

shorter and stouter a ae ne .. G. naviculator, p. 301. 


Genus Leptaulax, Kaup. 


J Depressed surface of two or three outermost grooves of elytra 
dull, the sculpturing somewhat worn-looking 

| Elytra polished throughout > ae a Be * 4. 
af more or less distinct polished siberate formed out of each 


iS) 


of the transverse ridges in lateral grooves of elytra .. L, beccarit, pp. 251 & 302. 
No such tubercles present : 3 see Be 4.6 3. 
‘Pronotum flatenned, densely panetused laterally, sides practi- 
cally straight .. 50 oe se .. L. humerosus, pp. 251 & 302. 
3, Pronotum convex; punctured only in marginal groove, in aud 
ee round scars, and (usually) in anterior angles; sides dis- 
eee curved .. ae 5 BG ea ania p. 803: 
/ Inner tubercles at most twice as far ftom one another as from 
outer tubercles; puncturing of lateral grooves of elytra some- 
what variable, but normally transverse toa marked degree in all 
species ; intermediate areas of metasternum closely punctured, 
unpunctured band along outer margin absent or quite narrow SF a 5 
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 


aS 
ee eee 


along the outer margin 8. 
Inner and outer tubercles daprowimavely equal, ene former at 
most twice as far from one another as from the latter 6. 
5, Inner tubercles much longer than outer, the later almost 
| obsolete; the former at least twice as far from one another as 
Ge the latter a as i .. L. macassariensis, pp.256 & 305. 


!T have no specimens of L. novaeguineae, L. obtusidens, or L. glaber before me, and cannot be cer- 


tain that these characters are correctly noted here in their case. 


326 Memoirs of the Indian Museum. [ Vou. III, 


ees chee unpunctured, except in scars and marginal grooves L. timoriensis, p. 303. 

Sides of pronotum more or less thickly covered with strong 

punctures AC an 0 se ai Sc =i a. 
,( Central area of metasternum unpunctured Ae .. L. dentatus, pp. 252 & 303. 
\ At least a few strong punctures on central area of metasternum L. cyclotaentus, pp. 255 & 305. 
{ Puncturing of abdominal sterna very variable; punctures, 


when present, of the usual type ae : A oe 30 Q: 
Abdominal sterua covered all over with soiidmhat obscure, 
| broad, shallow punctures aye os ie .. L. planus, pp. 260 & 310. 
f Posterior intermediate areas of metasternum punctured ae 50, ro. 
\ Posterior intermediate areas of metasternum unpunctured .. L. Wien: pp. 260 & 309. 


Central part of mesosternum sharply defined 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 
' the posterior margin, where they curve strongly outwards; the 


1 
i \ whole of the mesosternum outside these lines depressed and 
granular; only the central part smooth a8 L. antpunclus, pp. 256 & 306. 
Mesosternal sears smaller, not occupying the whole of the si: ie 
of the mesosternum te ai ne Il. 
{ Lateral grooves of elytra moderately broad Cp amekares slightly 
I1*\ transverse Sic Ge oe .. L. novaeguineae, p. 306. 
Lateral grooves of elytra narrower, punctures rounder 53 He 30 I2. 
{ Frontal ridges not reaching anterior margin of head 5 c a I3- 
\ Frontal ridges reaching anterior margin of head .. oa AE aieolor: pp 257 & 307. 
Posterior intermediate areas of metasternum finely Sidetared 
fee inner angle; elytra uniformly coloured ae .. L. obtusidens, p. 307. 


| Posterior intermediate areas of metasternum strongly and 
confusedly punctured ; elytra reddishin front and black behind L. glaber, p. 307. 


Genus Trichostigmus, Kaup. 


‘About 24 mm. long; a few punctures in anterior angles of 
ronotum; marginal groove of pronotum broad and deep and 
t=) 


I coarsely punctured xe oe a .. I. thoreyi,Kaup; pp. 261&310. 
_ About 17 mm. long; surface of pronotum unpunctured except 
(close to scars; marginal groove very fine or almost obsolete... T. wrsulus, Schaufuss; p- 310. 


g. APPENDIX III—THE GENUS TARQUINIUS, AND A REMARKABLE 
NEW GENUS FROM NEW GUINEA. 


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- 
eee genera outlined above is of peculiar interest. It appeets, moreover, to 


' It is quite possible that this character is of less value tae in this case it at ee seems to 
be, and that these two species are not really distinct. 

* This distinction is by no meaus a satisfactory one; but the position of L. novaeguinede, as ex- 
plained above (p. 306), has not yet been properly determined. 


1914. | F. H. GRAvELY: An Account of the Oriental Passalidae. 324 


throw light on the position of that most aberrant of all Indo-Australian Passalids, 
Tarquinius paradoxus, Kuwert, concerning which I have not hitherto dared to hazard 
an opinion. 

Pseudepisphenus, n. gen. 

Lamellae of antennae six in number, all moderately long. Mandzibles 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. Mentumasin 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. I,ateral 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. Wollaston during the 


Utakwa River Expedition in Dutch New Guinea, and preserved in the British 
Museum. 


“ki 


TEXT-FIGURE 8. 


A. Pseudepisphenus 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 245 mm. The lamellae of the antennae are moderately long and slender, 
the last three much longer than the first three. The /abrum 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 mentwm 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 jad is highly polished, and entirely smooth 
except for afew punctures in front of the parietal ridges. Its general form is shown in 


328 Memoirs of the Indian Museum. [ Vor. ais 


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 pronotwm 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 lower side of the 
prothorax bear punctures and hair distributed as usual in the family, but not so 
thickly or extensively as sometimes. ‘The scutellwni is very indistinctly punctured all 
over; the mesothoracic episterna are coarsely punctured above and along the anterior 
margin; the mesosternum is highly polished throughout, except in the deeply im- 
pressed semi-lunar scars, and even these are scarcely dull. The lateral areas of the 
metasterynum 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 fosterior coxae are smooth. The scars of the abdominal sterna are 
more or less roughened and punctured. The e/ytra 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 Hypferplesthenus. 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 perplexus. 

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 possibility of the absence of primary scars from the mentum in this group 
has been pointed out above (p. 194, footnote); andit 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 Targuinius. ‘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 


Ig14.| F. H. GRAVELY: An Account of the Oriental Passalidae. 329 


Tarqumus than in Pseudepisphenus, i.e. that the peculiarities which have appeared 
in Pseudepisphenus tend to be accentuated in Targunius. 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 Leptaulacinae 
are the only known Indo-Australian forms to which Targuinius 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 
Leptaulaci..ae should not be included as well. Whether these several series are to rank 
as subfamilies or 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 
forms. 

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 ina 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 .. 58 fe .. Pseudepisphenus, Gravely. 
Inner tubercles situated on anterior margin of head as in Lep- 

taulax ; outer tubercles symmetrical .. fe .. Tarquinius, Kuwert. 


330 Memorrs of the Indian Museum. [Vou. IIT, 


The evolution of asymmetrical Leptau/ax-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 Leptau/ax 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 have not yet had time fully 
to establish themselves there. 

Whether the high degree of specialization thus indicated in Tarquinius will prove 
sufficiently advantageous to its possessor 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 Pseudepisphenus perplexus we havea species, obviously 
of Australian and not Oriental extraction, whose mentum is indistinguishable from 
that of an Oriental species. Its Australian ancestry 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 Tarquinius. ‘Two so- 
called species, ‘‘ Laches’’ infantilis and puerilis, 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. 


10. APPENDIX IV—SUPPLEMENTARY CATALOGUE OF SPECIMENS IN 
THE INDIAN MUSEUM COLLECTION. 


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. 


1914. | F. H. GraveLy : An Account of the Oriental Passalidae. 


Subfamily AULACOCYCLINAE. 
Genus COMACUPES, Kaup. 
Comacupes cylindraceus (Perty). 


Reed. No. a Telom, S. Perak, 4000 ft. Oxford Museum. 


20° 20 


Comacupes foveicollis, Kuwert. 


7295-6 OAS 
Regd. No. ae Baram R., Sarawak British Museum. 


Genus TRISTORTHUS, Kuwert 
Tristorthus tricuspis (Kaup). 


Regd. No. 3” Yahoué, New Caledonia K. Zool. Mus., Berlin. 


20 


Genus TAENIOCERUS, Kaup. 


Taeniocerus platypus, Kaup. 


1372-4 


Regd. No. a0 Deli, N.E. Sumatra K. Zool. Mus., Berlin. 
Genus AULACOCYCLUS, Kaup. 
Aulacocyclus deyrollei, Kaup. 
Regd. No. ** Ballarat Djst., Victoria Deutsches Entom. Mus. 
Aulacocyclus sp. 
Regd. No. = Australia ? 
Genus CERACUPES, Kaup. 
Ceracupes arrowi, Heller. 
Regd. No. = Kosempo, Formosa Deutsches Entom. Mus. 
Genus PURITULUS, Zang. 
Auritulus patalis, Lewis. 
Regd. No. “* (cotypes) Yuyama, Japan British Museum. 


Subfamily ACERAIINAE. 
Genus OPHRYGONIUS, Zang. 


Ophrygonius cantori subsp. convexifrons, Zang. 


7283-92 : , ae 
Regd. No. 20 Sin Lum, Bhamo, 6000 ft. British Museum. 
7293—4 
» = Bhamo 


” 


Oo 
Oo 
iS) 


Regd. No. 


Regd. No. 


Regd. No. 


29 


Memoirs of the Indian Museum. [VoL 


Ophrygonius inaequalis (Burmeister). 


= Singapore British Museum. 
es Penang a 
Genus ACERAIUS, Kaup. 
Aceraius perakensis, Kuwert. 
= Taiping, 4000-4500 ft. K. Zool. Mus. Berlin. 
ms Telom, S. Perak, 4000 ft. Oxford Museum. 
Aceraius laevicollis (Illiger). 
4386-7 j W. Sumatra (‘Bandar Buat b.| | ’ 
20 i Padang Schoede S. G.’’) ' K. Zool. Mus., Berlin. 
ee Delhi, Sumatra - 
“ Java Deutsches Entom. Mus. 
“ Bandjermasin K. Zool. Mus., Berlin. 
~ S. Palawan Deutsches Entom. Mus. 
= Luzon K. Zool. Mus., Berlin. 
Aceraius grandis (Burmeister), s. str. 
= Sumatra K. Zool. Mus., Berlin. 
a Tengger Mountain, Java Deutsches Entom. Mus. 
ao ee K. Zool. Mus., Berlin. 
a ; foe Deutsches Entom. Mus. 
at Bandjermasin K. Zool. Mus., Berlin. 
A. grandis var. rectidens, Kuwert. 
oe Deli, Sumatras K. Zool. Mus., Berlin. 
“s Java 3 
ae (A. Magnus, > Deutsches Entom. Mus. 


Kuwert det.) 


A. grandis subsp. hirsutus, Kuwert. 


“ Catchin Cauri Deutsches Entom. Mus. 
— Mt. Mauson, 2-3000 ft. Tonkin K. Zool. Mus., Berlin. 
1“ Tonkin Deutsches Entom. Mus. 
eee Hainan K. Zool. Mus., Berlin. 
o Fuhosho, S. Formosa os 

a Kosempo, Formosa Deutsches Entom. Mus. 
ner Polisha, Formosa K. Zool. Mus., Berlin. 
M512 4458 


Sokutsu, Formosa Deutsches Entom. Mus. 


2 «20 


pies 


Regd. No. 


Regd. No. 


Regd. No. 


F. H. Grave.y: An Account of the Oriental Passalidae. 


Aceraius kuwerti, Zang. 


a Mt. Kina-Balu, c. 5000 ft. Deutsches Entom. Mus. 

sake ? 

20 : ” 

cu Mt. Kina-Balu, 4500 ft. Sarawak Museum. 
Aceraius laevimargo, Zang. 

wee Mt. Kina-Balu, c. 5000 ft. Deutsches Entom. Mus. 
Aceraius pilifer (Percheron). 

“ne Preanger, Java, 4000-6000 ft. Hamburg Museum. 

a ‘Tjibodas, c. 5000 ft. K. Zool. Mus., Berlin. 

“re Tjibodas Hamburg Museum. 

oe Borneo H. E. Andrewes. 

as Mt. Kina-Balu, 4500 ft. Sarawak Museum. 
Aceraius helferi, Kuwert. 

_ Sin Lum, Bhamo, 6000 ft. British Museum. 

Po Rangoon ss 
Aceraius borneanus, Kaup. 

“— Mt. Kina-Balu, c. 6000 ft. Deutsches Entom. Mus. 

4AST 


S. Palawan 


20 we 


Aceraius minor, Gravely. 


M5 (coty pe) 


20 


Taiping, 4000-5000 ft. K. Zool. Mus., Berlin. 


6214-6 
20 


Telom, S. Perak, 4000 ft. Oxford Museum 


Subfamily MACROLININAE. 
Genus MACROLINUS, Kaup. 


Macrolinus crenatipennis, Kuwert. 


8° (cotype”) Ceylon K. Zool. Mus., Berlin. 


Macrolinus sulciperfectus, Kuwert. 


= (cotype) Bonthain, Celebes K. Zool. Mus., Berlin. 


Subfamily GNAPHALOCNEMINAE. 
Genus EPISPHENOIDES, Kuwert. 


Episphenoides quaestionis (Kuwert). 


4431 


Sie Sidney Deutsches Entom. Mus 


| Larva, pupa and adult in spirit. 
2 Kuwert says that he only saw one specimen ; but he gives the lengths of two. 


Oo 


Oo 


Regd. 


Regd. 


Regd. 


Regd. 


Regd. 


Regd. 


Regd. 


Regd. 


Regd. 


Regd. 


No. 


No. 


No. 


No. 


Memoirs of the Indian Museum. (Vor. III, 


Genus MASTOCHILUS, Kaup. 
Mastochilus polyphyllus (Macleay). 


= Australia K. Zool. Mus., Berlin. 


Genus PHAROCHILUS, Kaup. 
Pharochilus sp. 


Australia ? 


Genus KAUPIOLOIDES, Gravely. 
Kaupioloides trigonophorus (Zang). 


4425 
20 


Toricelli Mts., Kaiser Wilhelmsland Deutsches Entom. Mus. 


Genus CETEJUS, Kaup. 


Cetejus grabowskyi, Kuwert. 


oe Stephansort, Kaiser Wilhelmsland —_K. Zool. Mus., Berlin. 
Genus PROTOMOCOELUS, Zang. 
Protomocoelus heynei (Kuwert). 
ares : Kaiser Wilhelmsland, New Guinea K. Zool. Mus., Berlin 
Genus KAUPIOLUS, Zang. 
Kaupiolus moluccanus (Percheron). 
4 Moluccas Deutsches Entom. Mus. 
Kaupiolus compergus (Boisduval). 
a Kapaur, Dutch New Guinea K. Zool. Mus., Berlin. 
Genus LABIENUS, Kaup. 
Labienus, sp. 
-_ ? K. Zool Mus., Berlin. 
= Bukana, Huon Gulf, German New us 35 
Guinea 
Genus HYPERPLESTHENUS, Kuwert. 
Hyperplesthenus gracilis (Heller). 
HS (cotype) Toricelli Mts. Kaiser Wilhelmsland Deutsches Entom. Mus. 


Genus PARAPELOPIDES, Zang. 


Parapelopides symmetricus, Zang. 


= Mt. Kina-Balu, c. 5000 ft. Deutsches Entom. Mus. 


1914. | 


Regd. 


No. 


. No. 


I’. H. GRavELy: An Account of the Oriental Passalidae. 


Se) 
W 


o7T 


Genus TRAPEZOCHILUS, Zang. 
Trapezochilus dorsalis (Kaup). 


a Taiping, 4500-5000 ft. K. Zool. Mus., Berlin. 

ee Bukit Besar, 2500 ft. 

sae Telom, 4000 ft. } Oxford Museum. 

“we Deli, Sumatra Deutsches Entom. Mus. 
Genus GNAPHALOCNEMIS, Heller. 
Gnaphalocnemis burmeisteri (Kaup). 

aa Padang, W. Sumatra Deutsches Entom. Mus. 

“ Deli, Sumatra 5 

a“ Peinan, Sumatra K. Zool. Mus., Berlin. 

“ Kepahiang se 

Gnaphalocnemis monticulosus (Smith). 

= Deli, Sumatra K. Zool. Mus., Berlin. 

ae Sumatra of 

= Sarawak ai 
Gnaphalocnemis tridens (Wiedemann). 

= java Deutsches Entom. Mus. 

a Asia ' 56 

a“ Tjibodas, Java K. Zool. Mus., Berlin. 

Genus OMEGARIUS, Kuwert. 
Omegarius minimus, Kuwert. 

ut Samberi, Dutch New Guinea K. Zool. Mus., Berlin. 

“ Kaiser Wilhelmsland, New Guinea 5 

“ie N. Bucht Squally I., New Britain Hamburg Museum. 

Subfamily TARQUINIINAE 
Genus TARQUINIUS Kuwert. 
Tarquinius parodoxus, Kuwert. 
“ Wandesi, New Guinea Deutsches Entom. Mus. 
Subfamily LEPTAULACINAE. 
Genus LEPTAULAX, Kaup. 
Leptaulax humerosus, Kuwert. 

= W. Sumatra Deutsches Entom. Mus. 

a Ardjoeno, Java ” 

4457 


=r Tengger Mountain, Java » 


336 


Memoirs of the Indian Museum. 


[Vor. ITI 


Leptaulax anna, Zang. 


Regd. No. “i (cotype) Sumbawa 


Hamburg Museum. 


Leptaulax timoriensis (Percheron). 


Regd. No. Timor 


K. Zool. Mus., Berlin. 


L. dentatus (Fabricius) s. str. 


Regd. No. “2 Mt. Mauson, 2000-3000 ft. 
23 — Fuhosho, Formosa 
1463 


” a] Polisha, Formosa 
4460 4162 hit 


at 29 2 2 


Sokutsu, Formosa 


K. Zool. Mus., Berlin. 
Deutsches Entom. Mus. 


? 


L. dentatus, var. glabriventris, Gravely. 


Reed. No. ‘44 


20 


Deli, Sumatra 


K. Zool. Mus., Berlin. 


Leptaulax cyclotaenius subsp. himalayae, Kuwert. 


K. Zool. Mus., Berlin. 


K. Zool. Mus., Berlin. 
Deutsches Entom. Mus. 
K. Zool. Mus., Berlin. 
Deutsches Entom. Mus. 
Hamburg Museum. 

K. Zool. Mus., Berlin. 


Hamburg Museum. 


” 


Regd. No. 4i° Mt. Mauson, 2000-3000 ft. 
Leptaulax bicolor Fabricius) s. str. 

Regd. No. “3>5 4 Mt. Mauson, 2000-3000 ft. 

5 “we Kosempo, Formosa 

i. “es Polisha, Formosa 

4s a Taiping, 4000-5000 ft 

- — St. Rambe, Sumatra 

md air Preanger, Java, 4000-6000 ft. 

es = = Tyjibodas, c. 5000 ft. 

a = Tjibodas, Java 

i “ey Banguey Island 

Leptaulax bicolor var. vicinus (Percheron). 
4758-65 


Regd. No. 


20 


11, 


Mt. Kina-Balu, 4500 ft. 


Sarawak Museum. 


SUMMARY AND CONCLUSIONS. 


1. Classification. 


The Indo-Australian Passalidae have been divided into two main sections (pp. 


IgI—1Q2). 
Cylindrocaulus and Auritulus 


The first section contains the Aulacocyclinae of Kuwert, together with 


It has been regarded as a single subfamily, but the 


two last-named genera and Ceracupes may 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, 


1914. ] F. H. Gravely: An Account of the Oriental Passalidae. 337 


Kaupioloides, Protomocoelus, Hyperplesthenus, Gnaphalocnemts, Plesthenus, Gonatas, 
Tarquinius and Leptaulax groups. Of these groups the first three and the last two appear 
to beof 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. 107, 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 Leptaulacinae ; 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 all 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 test 
by reason of their remarkable variability, in size especially ; and certain structural 
variations are corellated with variation in size, much as in the Lucanidae (pp. 262— 
265). These species are Episphenus comptont, Episphenus indicus, and Aceraius 
evandis 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 Leptawlax 
dentatus and Leptaulax bicolor which are the dominant species of Leptaulacinae 
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 Leftaulax-like form, 


1 As regards the Gnaphalocneminae see Gravely, 1914 (0). 


338 Memoirs of the Indian Museum. [Vo1r. IIT, 


Tarquinius, from an asymmetrical ancestor allied to the Gnaphalocneminae, 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 Aulacocyclinae 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 
Macrolininae the reverse is the case (pp. 311 & 313). 

The discontinuous distribution of the genus Awlacocyclus suggests that the 
Aulacocyclinae were once more abundant towards the west than they are now. 
Although the Macrolininae 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 specialization 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 


T914. | 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 
mainland. 

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. 
SS iss 15) E 

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 Aceralinae, and two to the Leptau- 
lacinae (p. 262). 

Of the first three, one (Episphenus comptonz) is confined to Ceylon, one (Epis- 
phenus 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 widely 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 ; Tiberioides, 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 
(Taentocerus bicuspis, p. 211; Pleurarius brachyphyllus, p. 215; Tibertordes austent, 
pp- 216-7 ; Episphenus compton, pp. 219-220; Episphenus indicus, p. 222 ; Episphenus 
neelgherriensis, p. 223; Aceraius grandis, sub-sp. hirsutus, p. 233; Aceraius helfert, 


340 Memoirs of the Indian Museum. [Vor. EIT, 


p. 238; Macrolinus sikkimensis, p. 243; Macrolinus rotundifrons, p. 245; Leptaulax 
dentatus, p. 255; Leptaulax bicolor, pp. 258-9; and Leptaulax roepstorfi, 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 Lamellicornia 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 voepstorfi, 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 (Leptaulax 
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, Aceraius 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 (Episphenus 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. 223). 


VIII.—LIST OF PUBLISHED PAPERS DEALING WITH PASSALIDAE 
BELONGING TO GENERA FOUND IN THE ORIENTAL REGION. 


Papers marked with an asterisk (*) are not available in Calcutta. 


1792. Fabricius, J. C. ‘‘ Entomologia Systematica emenda et aucta.’’ Vol. I 
(Hafniae, 1892), pp. 240-1. 
* 1800. Illiger, K. ‘‘ Vierzig neue Insecten aus der Hellwigischen Sammlung in 
Braunschweig’ Wiedemann Archiv fiir Zool. u. Zoot. (Berlin and 
srunswick 1800) I (2), pp. 103-150, and II, pp. 229-230. 
* 1800. Erichson in Wiedemann Archiv fiir Zool u. Zoot. I (1800). 


1914. | 


T8or. 
I8or. 


F. H. GRAvELY: An Account of the Oriental Passalidae. 341 


Fabricius, J.C. ‘‘ Systema Eleutheratorum II’’ (Kiliae, 1801), pp. 255-6. 
Weber, F. ‘‘Observationes Entomologica’’ (Kiliae, 1801). 


* 7806-17. Schonherr,C. J ‘‘Synonymia Insectorum,oder: Versuch einer Syno- 


123. 


1826. 


1842. 


1843. 


1844. 


1847. 


1850. 


1852. 


1855. 


nymie aller bisher bekanten Insecten; nach Frabricii Systema 
Eleutheratorum geordnet ’’ (Stockholm, 1806-17). 

Wiedemann, C. R. ‘‘ Zweihundert neue Kafer von Java, Bengalen, und 
dem Vorgebirge der Gutten-Hoffiniing.’’ Zool. Mag. II (1), p. 109. 
Macleay, W. S. ‘‘Catalogue of Insects collected by Captain King, 
R.N.’’ Appendix to P. P. King’s ‘‘ Narrative of a Survey of the 
Intertropical and Western Coasts of Australia performed between the 

years 1818 and 1822.’’ Vol. II (London, 1826). 

Sturm, J. ‘‘Catalog meiner Insecten-Sammlung,’’ I (Niirnberg, 1826). 

Perty. ‘‘Obs. Nonnulae Coleopt. Indiae Orientalis.’’ 

Boisduval, J. B. A.D. de. ‘‘Faune Entomologique de 1!’Ocean Paci- 
fique. II. Coléoptére et autres Ordres’’ in ‘‘ Voyage de I’ Astrolobe 
....pendent....1826-9 sous le commandent de M. J. Dumont 
qd’ Urville etc 

Percheron, A. ‘‘ Monographie des Passales (Paris, 1835). 

Dejean, P. F. M. A. ‘‘ Catalogue de la Collection de Coléoptéres de M. 
le Baron Dejean,’’ 3 ed. (Paris, 1837), pp. xiv, 503. 

Percheron, A. ‘‘ Révision critique et Supplement 4 la Monographie du 
Genre Passale, premiére partie.’ Mag. Zool., Insectes, 1841. PI. 
Ixvit-Ixix. 

Westwood, J.O. ‘‘Insectorum novarum Centuria.’’ Ann. Mag. Nat. 
Hist. VIII, 1842, pp. 123-125. 

Guérin Méneville, F. E. ‘‘Animaux Articulés’’ in ‘‘ Souvenirs d’un 
Voyage dans l’Inde executé de 1834 4 1839 par M. Adolphe Deles- 
sert,’’ Pt. II (Paris 1843), pp. 33-98. 

Percheron, A. ‘‘ Monographie des Passales, Second Supplement.’’ Mag. 
Zool., 1844, pl. cxxxiv-cxxxv. 

Hope, F. W. A. (Westwood) ‘‘ A Catalogue of the Lucanoid Coleop- 
tera in the collection of....F. W. Hope, ete.”” (London, 1845), p. 31. 

Burmeister, H. ‘‘Handbuch der Entomologie,’’ V (Berlin, 1847), pp. 
461-510. 

Castelnau, F. L. de L. de. ‘‘ Histoire Naturelle des Animaux Arti- 
culés.”” Insectes Coléoptéres par M. le Compte de Castelnau, II 
(Paris 1850), pp. 178-9. ; 

Smith, F. ‘Nomenclature of Coleopterous Insects in the Collection of 
the British Museum. Pt. VI Passalidae’’ (Ijondon, 1852). 

Montrouzier, le Pére. ‘‘ Essai sur la Faune de 1’Ile de Woodlark ou 
Moiou.’’ Ann. Sci. Phys. et Nat. d’ Agric. et d’Industrie (Soc. Imp. 
d’ Agric. etc. de Lyon, 1855) VII (1), pp. 1-114. 


Vs 
NO 


1860. 


1868. 


1868. 


1869. 
1871. 


I88o. 
1883. 


1884. 


1885. 


1887. 


1890. 
1891. 
1896. 


1896. 


Memoirs of the Indian Museum. [Vou. III, 


Montrouzier, le Pére, ‘‘ Essai sur la Faune Entomologique de la Nouvelle- 
Calédonie (Balade) et des Iles des Pins, Art, Lifu ete.’’ Ann. Soc. 
Ent. France (3) viii, pp. 229-308. 

Fauvel, A. ‘‘Coléoptéres de la Nouvelle-Calédonie recueilles par M. E. 
Déplanche 1858-60.’’ Normandie, Soc. Linn. Bull. VII, pp. 120-185. 

Redtenbacher, L. ‘‘ Coleoptera’’ in ‘‘ Reise der Osterreichen Fregatte 

‘Novara’; Zool. II’’ (Vienna, 1867}, 249 pp., 5 pl. 

Gemminger and Harold. ‘‘Catalogus Coleopterorum’’ II (Munich, 
1868), pp. 968-972. 

Kaup, J. (a) ‘* Prodromus zu einer Monographie der Passaliden.”’ 

Coleopterologische Hefte, III, 1868, pp. 4-32. 
(b} Ditto. Coleopterologische Hefte, IV, 1868, pp. 1-7. 

Kaup, J. Ditto. Coleopterologische Hefte, V, pp. 38-40. 

Kaup, J. ‘‘Monographie der Passaliden.’’ Berlin Ent. Zeitschr. 15 
Jahrg. Supplement, 125 pp., pl. iii—vu. 

Macleay, W. ‘‘ Notes on a Collection of Insectes from Gayndah (Cole- 
optera) pt. III.” Trans. Ent. Soc., N.S. Wales IT, 1871, pp. 159-205. 

Stoliczka, F. ‘‘A contribution towards a Monograph of the Indian 
Passalidae.’’ J.A.S.B. XLII (II), 1873, pp. 149-162. 

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. 

Fairmaire, L,., in Le Naturaliste, 1880, p. 164. 

Lewis, G. ‘‘Iucanidae of Japan.’’ Trans. Ent. Soc. London, 1883, 
PP. 333-342, pl. xiv. 

Wystmann, P. ‘‘ Catalogue Systématique des Passalides.”” Ann. Civ 
Mus. Genova, (2a) I (X XI) 1884, pp. 326-348. 

Schaufuss, lL. W. “‘ Beitrage zur Fauna der Niederlandischen Besit- 
zungen auf den Sunda-Inseln.’’ Horae Soc. Ent. Ross. XTX 1885, 
pp. 183-209. 

Fairmaire I. ‘‘ Coleopteres de l’Intérieure de la Chine.” Ann. Soc. 
Ent. Belg. XX XI, 1887, pp. 87-136. 

Kuwert, A. ‘‘ Die Passaliden und ihr Monographie von Kaup.’”’ Soc. 
Ent. V, 1890, pp. I, 9, and 17. 

Kuwert, A. ‘‘ Systematische Uebersicht der Passaliden Arten und 
Gattungen.’’ Deutsche Ent. Zeitschr. 1891 (1), pp. 161-192. 

Blackburn, ‘I. ‘‘New Genera and species of Australian Coleoptera, 
XX.’’ ‘Trans R. Soc. S. Australia XX, pp. 233-259. 

Heller, K. M. ‘‘ Neue Kafer von Celebes.”” Abh.u. Ber d.K. Zool. u. 

Anthr.-Ethn. Mus. zu Dresden, VI, 1896-7 (3), 24 pp., I pl. 


TgI4.]| 


1896. 


1897. 
1808. 


1808. 


1900. 
1902. 


1903. 


1903. 


1904. 


1905. 


1906. 


1907. 


IQIo. 


TOLL. 


F. H. GRAVELY: An Account of the Oriental Passalidae. 343 


Kuwert, A. “ Die Passaliden dichotomisch bearbeitet’’ Novit. Zool. 
III, 1896, pp. 209-230, pl. v—vii. 
Kuwert, A. Ditto. IV, 1897, pp. 274-306. 
Kuwert, A. Ditto. V, 1898, pp. 137-205 and 259-349. 
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 pl. 
Heller K.M. Ditto. IX, 1900 (5), 46 pp. 
Fruhstorfer. ‘‘ Tagebuchblattern’’ Insekten-Borse XIX, 1902, p. 28. 
Fauvel, A. ‘‘ Faune Analytique des Coléoptéres de la Nouvelle Calé- 
donie.’’ Rev. Ent. franc. XXII, 1903, pp. 203-378. 
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. 
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. 
181-5. 
Zang. R. (a) Numerous papers in Deutsche Ent. Zeitschr. for 1905. 
(b) ‘“Anderung in der Nomenclatur der Passaliden (Coleop- 
tera).’’ Zool. Anz. XXIX, 1905, pp. 154-5. 
(c) ‘‘ Passalidarum Synonymia. Kritische Revision der von 
Kuwert und anderen Autoren aufgestellten Gattungen 
und Arten.’’ Notes Leyden Mus. XXV, 1905, pp. 
221-232. 
(d) ‘““Zwei neue Passaliden aus den Gattungen Comacupes, 
Kp., und Aceraius, Kp.’’ Notes Leyden Mus. XXV, 
1905, pp. 233-8. 
Zang, R. (a) ‘‘ Uebersicht der Basilianus-Arten.’’ Deutsche Ent. 
Zeitschr., 1906, pp. 177-183. 
*(b) ‘ Passalini’”’ in ‘‘Nova Guinea, Résultats de l’Expedition 
scientifique Néerlandaise 4 la Nouvelle-Guinée en 
1903, sous les auspices de Arthur Wichmann, Chef de 
’Expedition.”” V (1) pp. 23-26. 


Arrow, G. J. ‘‘ A contribution to the classification of the Coleopterous 
Family Passalidae.’’ Trans. Ent. Soc London, 1906, pp. 441-469. 
Heller, K.M. “‘ Finfter Beitrage zur Papuanischen Kaferfauna. ‘‘Abh. 


u. Ber. d. K. Zool. u. Anthr-Ethn. Mus. zu Dresden, XIII, IQI0 (3), 
AW) 0402, 36 Ole 

Heller, K. M. ‘‘ Kine neue Ceracupes-Art aus Formosa.’ Ann Soc. 
Ent. Belg. LV, 1g11, pp. 256-7, 1 text-fig. 


344 Memoirs of the Indian Museum. (Vou. III, 1914.] 


1913. Gravely,F.H. ‘‘ Preliminary Account of a revised classification of Indo- 
Australian Passalidae.” J.A.S5.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. 
(b) ‘The Evolution and Distribution of certain Indo- 
Australian Passalidae, J.A.S.B. (In the Press). 


1914. ] F. H. Gravety: An Account of the Oriental Passalidae. 345 


. 


The WUNDDEOX. 


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 summary 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. 


Page. 
abdominibarbatus (Leptaulax) ie ao BYOBY 
abdominisculptus (Leptaulax) Dp OF 
Aceraiinae So 0 a .. 194-197, 215 etc., 280 etc., 811-3, 314, 315, 316-8, 
329, 331 etc. 
Aceraius ae a0 6 .. 181, 195, 197, 228 etc., 286 etc., 312-4, 314, 316-8, 
320 etc., 332 etc. 
Aceraius xs e ae .. 280, 281, 293, 299- 
addendus (Aceraius) xs - Se Aisles BO 
aequidens (Aceraius) Se 56 a 240,208 a lue G23. 
albertisi (Gonatas) bf i se SOE 
aliicornis (Aulacocyclus) .. ae ac) 27s 
altidens (Gonatas) Be cra ve 302% 
alutaceosternus (Aceraius) .. 56 2 230), 201, 323: 
Analaches ae es ae we 195. 200, S04 solo. 
Analaches se ave of . LO6: 
analis (Leptaulacides) fa a ae gOS; 
anaulax (Leptaulacides) .. is See yOlsh 
andamanarum (Leptaulacides) : COS: 
andamanensis (Bastlianus, Macrolinus, Tiberius) mon 24A2 e204 0824. 
andamanensis (Bastlianus, Tiberius) .. pa Z0yHle 
andrewesi (Aulacocyclus) .. = Bo PAE 7\n Sth, 
angusticornits (Comacupes) tt y. 209: 
angustifrons (Leptaulax) .. Xe mem O5¢ 
angustior (Eriocnemts) ov oe a 209s 
antbarbis (Leptaulax) M4 Bs .. 250, 803, 305. 
anipunctus (Leptaulacides, Leptaulax).. 22 250, 300, 326. 
anna (Leptaulax) : oh -. 303, 325, 336. 
arcuatoclypeatus (Aeesenaee) a See's 
arrowi (Ceracupes) dc oe Fa 278s SLO ool. 
aruensis (Aulacocyclus) .. pe sa Bifiig SO 
assamensis (Aceraius) ce : .. 237, 292, 323. 
Aulacocyclinae . . ae a: .. 185, 188, 189, 191-3, 204 etc., 267 etc., 311, 331. 
Aulacocyclus .. ae Bin .. 191, 192, 193, 2II etc., 272 etc., 311, 318, 310, 331. 
Aulacocyclus .. *% a 5 AM), ZO, ZiT 


Aurelius 06 te 50 Je ZO2,09 04. obo: 


346 Memoirs of the Indian Museum. 


Aurthulus 

Auritulus ve 
aurivillit (Leptaulax) 
austeni (Ceracupes) 
austeni (Tiberioides) 


barbicauda (Leptaulacides) .. 
Basilianus 

basalis (Comacupes) 

batchianae (Leptaulax) 

batesi (Macrolinus) 

beccarii (Leptaulaulax) 

bicanthatus (Azlacocyclus, Passalus, 


bicolor (Leptaulacides, Leptaulax, Passalius) 


bicolor (Leptaulax) 

bicuspis (A ulacocyclus, Taeniocerus) 
bthastatus (Aulacocyclus, Passalus) 
binominis (Basilianius) 

birmanicus (Ophrygonius) .. 
borealis (Chilomazus, Tiberioides) 
borneanus (Aceraius) 

borneanus (Aceraius) 

borneensis (Comacupes) 

bornensis (Leptaulax) 
brachyphyllus (Pleurarius) 

brevis (Gonatas) 

bucerus (Cylindrocaulus) 


burmeisteri (Eviocnemis, Guaphaloenenia 


caffer (Tiberius) 
calcuttae (Leptaulax) 


cancrus (Aceraius, Basilianus, Tiberius) 
cantori (Aceraius, Basilianus , Ophrygonius, jae ae 


cantori (Aceraius, Basilianus) 
Caulifer 

cavicornis (Aulacocyclus, Gouiacdoean: 
celebensis (Aulacocyclus) 
celebensis (Leptaulax) 
Ceracupes 

Ceratocupes 

certus (Basilianus) 

Cetejus 

cetioides (Gonatas) 

ceylonicus (Aceraius) 
Chilomazus 

chinensis (Aceraius) 
ciccalrosus (Leptaulax) 
collaris (Aulacocyclus) 
Comacupes 

Comatus (Aulacocyclus) 


Page. 

279. 

191, 192, 193, 279, 311, 319, 331. 
307-8. 

212, 262, 278, 310. 

216, 281, 320. 


308. 

195, 197, 280, 281, 284, 293, 316-7. 
267, 318. 

306, 307-8. 

296. 

251, 302, 325. 

208, 270, 319. 

257, 262-295, 307, 326, 3306. 
303, 310. 

210, 271, 319. 

272), 27. 

283. 

226, 285, 320. 

281, 320. 

238, 292, 318, 323, 338. 
2gl. 

205, 268. 

303. 

213, 262, 280, 320. 

301. 

191, 279, 3109. 

249, 298, 324, 335. 


194. 

308. 

280. 

224, 262, 284, 311, 320, 331. 
282, 285. 

193: 

181, 204, 268, 318. 

297. 

303. 


180, 188, 192, 212 etc., 277 etc., 311, 319, 381. 


278. 

282, 283. 

194, 201, 314, 315, 334. 
301. 

288. 

195, 196, 281, 316-7. 
288, 322. 

806, 307-8. 

275. 

I8I, 192, 193, 204, 267, 318, 331. 
267. 


1914. | F. H. GRAVELY: 


compergus (Kaupiolus) 


comptoni (Aceraius, Episphenus, Peaches) 


consequens (Leptaulax) 

convexifrons (Basilianus, Ophrygonius) 
cormocerus (Comacupes) 

cornutus (Ceratocupes) 

crenatipennis (Macrolinus, Tiberius) 
crinitus (Heterochilus) 

cyclotaenius (Leptaulax) 


cylindraceus (Aulococyclus, Golnacupés, Dassen 


cylindraceus (Comacupes, Passalus) 
Cylindrocaulus .. 


darjeclingt (Leptaulax) 
darjeilingt (Lepaulax) 
dentatus (Leptaulax, Passalus) 


deyrollei (Aulacocyclus, Taeniocerus) .. 


differens (Gonatas) 

differentispina (Leptaulax) .. 
dilatus (Aulacocyclus) 

dindigalensis (Leptaulax) 

dispar (Ertocnemis) 

dissimilis (Macrolinus) 

divaricatus (Leptaulax) 

dorsalis (Eviocnemis, Trapezochilus) 
duivenbodei (Macrolinus) 
dunsiriensis (Ophrygonius) 


edentulus (Aulacocyclus, Passalus) 
emarginatus (Aceraius, Passalus) 
Embryulcus 

Epilaches 

Episphenoides 

Episphenus 


Eriocnemis 
errans (Aulacocyclus) 


eschscholtzi (Leptaulacides, ental} xe 


evidens (Leptaulax) 
exterris (Leptaulax) 


faberi (Eviocnemis) 
felderi (Comacupes, Aeistorthus) 
filius (Analaches) 


flachi (Chilomazus, Rapliends: aches) 


flachit (Laches) 

foveicollis (Comacupes) 
foveipunctatus (Aulacocyclus) 
fraternus (Macrolinus ) 

fratricornis (Aulacocyclus) 
fronticornis (Ceracupes, Passalus) 


An Account of the Oriental Passalidae. 


Page. 

334. 

218, 262-265, 281, 311-2, 314, 320. 
306, 307-8. 

225, 284, 381. 

208, 269. 

278. 

244, 295, 324, 333 

280. 

255, 305, 326, 336. 

207, 269, 318, 331. 

208, 269, 275. 

191, 192, 193, 279, 311, 319. 


393: 
303- 


I81, 185, 252, 262-265, 303, 326, 336. 


272, 381. 
300. 

307. 

276, 319. 
308. 

299: 

296. 

308. 

247, 297, 324, 335. 
294, 323: 
225, 284. 


275. 

287, 288, 290, 291, 292, 293. 
202. 

195. 

200, 314, 315, 333. 


> 


347 


189, 194, 196, 217 etc., 281 etc., 311-3, 314, 315, 


316, 317, 320, 330. 
297, 298. j 
275. 
307-8. 
308. 
303. 


2098. 

267. 

283. 

219, 281. 
281. 

267, 318, 331. 
274. 

290. 

274. 

212, 277, 3109. 


348 Memoirs of the Indian. Museum. [Vor. VIII, 


fronticornis (Ceracupes) 
fruhstorfert (Laches) SE 
fruhstorjert (Leptaulacides) .. 
furcicornis (Passalus) 


gelon (Eriocnemis, Plesthenus) 
geminatus (Leptaulax) 

geminus (Leptaulax) 

generosus (Phraortes) 

germari (Aceratus, Gonatas) 
glaber (Leptaulacides, Leptaulax) 
glabricollis (Leptaulax) 
glabriusculus (Aulacocyclus) 
glabriventris (Leptaulax) 
Gnaphalocneminae 


Gnaphalocnemis 

Gnaphalocnemis group 

Gonatas 

Gonatas group .. 

grabowskyi (Cetejus) 

gracilis (Hyperplesthenus, Labienus) 
grandis (Aceraius, Passalus) 
gravidus (Pelopides) 


hansemanni (Leptaulax) 
helferi (Aceraius) 
Heterochilus 

heynei (Protomocoelus) 
himalayae (Leptaulax) 
himalayensis (Aceraius) 
hirsulus (Aceraius) 
humerosus (Leptaulax) 
humerosus (Leptaulax) 
Hy perplesthenus 
Hyperplesthenus group 


ignolus (Eviocnemts) 
illegalis (Aceraius) 
imitator (Taeniocerus) 
impar (Hyperplesthenus) 


inaequalis (Aceraius, Basilianus, Opsieoniie: Pas- 


salus) 
inaequalis (Bassilianus) 
incidens (Aceraius) 
inctpiens (Leptaulax) 
indicus (Basilanus, Epishenus) 
indicus (Leptaulax) 


infantilis (Aceraius, < Neues Epilaches, ene 


innocuus (Passalus) 


Page. 
278. 
281. 
308. 


275. 


298. 

303, 306. 

308. 

297. 

250, 301, 318, 325. 

307, 326. 

303. 

276. 

255; 303, 336. 

191, 194, 195, 197-203, 246 etc., 297 etc., 313-6, 
333. 


191, 202, 248 etc., 298 etc., 314, 318, 324, 329, 335. 
199, 202. 

203, 250 etc., 299 etc., 313, 314, 318, 325. 

199, 2or. 

334. 

202, 334. 

181, 231, 262-265, 288, 312-3, 314, 322, 332. 

201, 203, 208. 


300. 

238, 292, 323, 333. 

195, 286. 

334. 

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. 


209. 
230, 287, 321. 
270. 
202. 


227, 285, 320, 332. 

283. 

293- 

307-8. 

220, 262-265, 282, 311-2, 314, 320. 
303. 

283, 330. 


307. 


1914. | F. H. GravEty: An Account of the Oriental Passalidae. 


insignis (Leptaulax) 
insipiens (Leptaulax) 
interponendus (Leptaulax) .. 
interrogationts (Bastlianus). . 


javanus (Macrolinus) 


kaupi (Aulacocyclus) 
kaupt (Aceraits) 


Kaupioloides 
Kaupioloides group 
Kaupiolus 56 


Kaupiolus group 
kuwerti (Aceraius) 
kuwerti (Tiberioides, Tzbertus) 


Labienus 

Laches é ate 
laevicollis (Aceraius, Passalus) 
laevicollis (Aceraius) 


laevicornis (Aulacocyclus, Comacupes) .. 


laevimargo (Aceraius) 
langier (Aceraius) 
laterisculptus (Passalus) 
latipennis (Macrolinus) 
Lettaulacides 
Leptaulacinae 


Leptaulax 20 
luzonicus (Aceraius) 


macassariensis (Leptaulax) .. 
Macrolininae 

Macrolinus 

magnus (Aceraius) 

major (Gonatas).. 

malaccae | Leptaulax) 
malitiosus (1,eptaulax) 
manillae (Leptaulax) 
marginepunctatus (Passalus). . 
masoni (Comacupes) 

masont (Comacupes) 
Mastachilus 5 
mastersi (Aulococyclus, Taeniocerus) 
Mastochilus 

maxillonotus (leptaulax) 
medius (Leptaulax) 

meyeri (Aceraius) 

minimus (Omegarius) 

minor (Aceraius).. 


Page. 
303: 
308. 
303. 
284. 


290. 


200, 314, 334. 

199, 200. 

202, 314, 334. 

198. 

235, 290, 322. 

215, 226, 280, 320, 332. 


191, 202, 314, 315, 334. 
195, 281, 330. 

230, 287, 313, 322, 332. 
203. 

181, 205, 268. 

ABS AO), $3}. BBBT 
234, 290, 322. 

290- 

245, 296, 324. 

208, 204, 302. 


49 


191, 194, 199, 203-204, 251 etc., 302 etc., 311, 329, 


335-6. 


181, 185, 189, 191, 203, 204, 251 etc., 302 etc., 


318, 325-6, 330, 335-6. 
287. 


256, 305, 325- 


194-195, 197, 240 etc., 293 etc., 311, 329, 333. 
181, 197, 240 etc., 293 etc., 318, 323, 333. 


288. 

300. 

307-8. 

303. 

307. 

290. 

207, 208, 318. 

209. 

195. 

193, 273. 

178, 195, 200, 314, 334. 
307-8. 

307-8. 

287. 

335. 

240, 293, 317, 323, 338. 


350 Memoirs of the Indian Museum. (Vor. VIII, 
Page. ; 

minor (Comacupes) a x ee ley A 

minutifrons (Aceraius) vA a 200: 

mixtus (Leptaulax) Ar see 308. 

mniszecht (Eriocnemts, Ghaphalamiemtes se 6209: 

moeschlevi (Aceraius) Ae x ey 

molossus (Aceraius) on “t Me estes 

moluccanus (Kaupiolus) .. . 884, 

monticulosus (A ceraius, Lriocnemis, Giapkaloctenne 

Passalus) a ’ aes -- 249, 298, 325, 335. 

moorei (Episphenus) =e oe .. 277, 281, 811, 312, 315, 320. 

morator (Leptaulax) xe or -. 306. 

moschleri (Aceraius) a ie Aa Soci SLC 

nanus (Aceraius) a: ~ A bh 

naviculator (Gonatas) me : vs | SOD, S1S51925° 

neelgherriensis (Acevaius, Basilianus, Hnisnbenee, 

Passalus) ne é Hye we 222, 370, 283, SL), 624, 920,000. 

neelgherriensis (Aceratus, Basile) oe Ay EYE 

Neleus : ae si Yer 

niae aeesiaciox) A a SOS: 

nicobaricus (Aceraius) a te eeOL. 

nicobaricus (Macrolinus) .. he ree Ces Chopin By T 

nietnert (Leptaulacides) u Se eS OO: 

nikobaricus (Passalus) ty ds eZOL 

nobilis (Phraortes , Trapezochilus) aye 2472076 

novaebritanniae (Gonatas) .. ee ne 300: 

novaeguineae (Letaulax) .. oie -. 306, 326. 

obtusidens (Leptaulax) a =f ae S071 S20 

occulidens (Aceraius) & pete: .. 234, 290, 312-8, 322. 

oculitesselatus (Heterochilus) ar: Ba Aclor 

Ocythoe oe Ae xs sa oles 

Omegarius ‘a ae oe .. 203, 314, 385. 

Ophrygonius .. a se .. 196, 224 etc., 284 etc., 311, 312-3, 314, 316-7, 

318, 320, 381. 

orbatus (Macrolinus) a Ac 200; 

orientalis (Passalus) : «. 290. 

ovoleius (Leptaulax, Ophrygonius, Pasediies se, Bile 

palawanicus (Leptaulacides) it tao stoleh 

palawanus (Aceraius) ae uA ne 2006 

papua (Leptaulax) 36 is a) 305: 

papuanus (Letpaulacides) .. °* at a shat 

paradoxus (Tarquinius) a Sk .. 327, 835. 

parallelipennis (Macrolinus) 5c hee2g0: 

parallelogrammifrons (Laches) te Sip etsher 

Parapelopides .. ee a .. 202, 246, 297, 314, 324, 384. 

parreyt (Aulacocyclus) ae ou 5 290, 299. 

parryi (Aulacocyclus) ne 2s e276; 310: 


parryi (Aulococyclus) Sc ¥e is ©6208. 


1914. | F. H. Gravety: An Account of the Oriental Passalidae. 


parvus (Taeniocerus) ts 


patalis (Aulacocyclus, Auvikulus, Auritulus) 


Paxilloides 

pearsont (Episphenus) 
pectinigera (Episphenoides) 
Pelopides 

Pelopides 

Pelops 

perakensis (Aceraius) 
percheront (Aulacocyclus) 
percheronii (Aceraius) 
peregrinus (Gonatas) 
perlatus (Aulacocyclus) 
perplexus (Pseudepisphenus) 
perturbans (Taeniocerus) 


Pharochilus as Se 
philippinensis (Paxilloides) .. 
Phraortes 


pilifer (Aceraius) 
pulifer (Aceraius) 
pilipes (Pleurarius) 
planicollis (Leptaulax) 


planus (Leptaulacides, Leptaulax, Passalus) 


planus (Leptaulax) é 
platypus (Aulacocyclus, Taeniocerus) 
Plesthenus 

Plesthenus group 

Pleurariinae 

Pleurarius 

politus (Mastochilus) 

polyphyllus (Mastochilus) 

pontifex (Aceratus) 

Proculus = 
prosternisulcatus (Aceraius) . . 
Protomocoelus 

Protomocoelus group 
Pseudepisphenus 

puella (Laches) 

puertlis (Aceraius, Laches) .. 
pugnax (Aulacocyclus, Comacupes) 
pulchellus (Leptaulacides) 

punctifrons (Comacupes, Passalus) 
p} gmaeus (A ulacocyclus, Taeniocerus) 


quadridentatus (Passalus) 
quadrifer (Ophrygonius) 
quaestionis (Episphenoides) 


rectidens (Aceraius) : 
redtenbacteri (Semicyclus) .. 


Page. 

271. 

279, 319, 331. 

178. 

218, 281. 

315. 

197, 199, 201, 314. 
201, 208. 

201. 

22.0 9287) 321, 323,302" 
274. 

292. 

301. 

270. 

327. 

270. 

200, 314, 334. 

178. 

297. 

235, 291, 323, 333. 
292. 

279, 320. 

309. 

260, 262, 310, 326. 
309. 

2703 LO: 

202, 314. 

199. 

194, 199, 213 etc., 279 etc., 311, 329. 
194, 213 etc., 279 etc., 320. 
178. 

334. 

287. 

191. 

288. 

191, 199, 201, 314, 334. 
199, 200. 

327, 329, 380. 

281. 

283, 330. 

277. 

307. 

209. 

209, 271, 319. 

303- 

286. 

333. 


233, 288, 322, 332. 
178. 


352 Memoirs of the Indian Museum. 


respectabilis (Phraortes, Trapezochilus) 
reticulaticollis (Aceraius) 
Rhipsaspis 

ribbei (Leptaulax) 

roepstorfi (Leptaulax) 

roepstorfi (Leptaulax) 

rosenbergii (Aulacocyclus) 
rotundatoclypeatus (Aulacocyclus) 
rotundifrons (Macrolinus, Tiberius) 
rugulosus (Leptaulacides) 


saigonicus (Leptaulax) 
schellongi (Gonatas) 
schenklingi (Analaches) 
schmidtii (Paxilloides) 
schraderi (Pelopides) 
Semicyclus 

separandus (Leptaulax) 
sequens (Leptaulax) Se 


sikkimensis (Basilianus, Macrolinus Tiberius) 


simplex (Gnaphalocnemis) .. 
singapurae (Ophrygonius) 
singhalensts (Macrolinus, Tiberius) 
sinkepicus (Basilianus) 

socius (Laches) 

stoliczkae (Basilianus) 
stoliczkae (Comacupes) 
submedius (L,eptaulax) 
subsequens (Leptaulax) 
sulcatipons (Aulacocyclus) 
sulciperfectus (Macrolinus) 
sumatrae (Leptaulax) 
symmetricus (Parapelopides) 


Taeniocerus 

Taentocerus 

Tarquiniinae 

Tarquinius 

Tatius i 

tavoyanus (Aceraius) 

tavoyanus (Macrolinus) 
tenassertmensis (I,eptaulax) 
tereoides (Aulacocyclus) 

teres (Aulacocyclus, Passalus) 
thoreyi (Leptaulax, Trichostigmus) 
Tiberioides 

Tiberius 

tibialis (Gonatas) Be 
timoriensis (Leptaulax, Passalus) 
timortensts (Leptaulax , Passalus) 


Page. 

248, 297. 
2g1. 

286. 

303, 3086. 
260, 309, 326. 
308. 

274, 319. 
274. 

244, 295, 324. 
308. 


303. 
300. 
200. 


307-8. 

308. 

243, 2904, 323. 
248, 298, 324. 


226, 285, 317, 320. 


205. 
286. 
281. 
282. 
206, 268, 318. 
303. 
308. 
273. 
296, 324, 333. 
308. 


246, 297, 324, 334. 


192, 193, 208 etc., 270 etc., 319. 


272. 
329, 335. 


178, 327, 328, 329, 330, 335. 


197, 203, 314. 
237) 292- 
243, 294 

308. 

273. 

273. 

261, 310, 326. 


[Vor VIII, 


189, 196, 215 etc., 280 etc., 311-3, 314, 317, 320. 


194, 195, 197, 280, 293. 


300. 
303, 326, 336. 
303. 


Ig14. | F. H. GRAVELY: An Account of the Oriental Passalidae. 353 


tonkinensis (Leptaulax) 
Trapezochilus 
Trichostigmus 
tricornis (Aceraius) 
tricuspis (Tristorthus) 


tridens (Eviocnemts, Guaplialocneniié, Ocythoé, Pas 


salus) 
tridens (Eviocnemts) 
tridentatus (Gonatas) : 
trigonophorus (Kaupioloides, ice pie Das) 
Tristorthus 


ursulus (Leptaulax, Trichostigmus) 
urus (Macrolinus) 


variolosus (Aulacocyclus) 
Vellejus : ; 
vicinus (Passalus, Remains eoteutans 


wallacei (Aceraius, Heterochilus) 
waterhousei (Macrolinus, Tiberius) 
weberi (Macrolinus) 

weber! (Macrolinus) 

westermannt (Comacupes) 


yorkensis (Gonatas) 


Page. 


202, 247 etc., 207 etc., 314, 324, 335. 
203, 261 etc., 310, 318, 326. 
235, 290, 322. 


250, 299, 325, 335. 


202, 334. 
192, 193, 268, 331. 


310, 326. 
204, 323- 


259, 307, 386. 


228, 286, 317, 321 
245, 205, 324. 
245, 296, 324. 


a ay 


‘ 
, 


Last 


P 


wiry be ; i* 


+s 7 
Aan ad Pp, ie vs 


“pied : 
eater 

wees ye 
Padme! *} ‘a 


cain 


ae 

7 ore 
erik: a . ees b hh Fat ANG 
sey ewe) PA\ iv agant ete 


4 


EXPLANATION OF PLATE XI. 


1.—Comacupes cavicornis var. borneensis (type). Head from above, x 2. 


Ia. Ditto Head from the side, x 2. 

2.—Comacupes cavicornis var. laevicornis (reed. no. *\’). Head from above, 
x2: 

2a. Ditto Head from the side. x 2. 

2b. Ditto Mentum, etc., x 4. 

3.—Comacupes stoliczkae (type). "Head from above, x 2. 

3a. Ditto Head from the side, x 2 

4.—Comacupes masont (type). Head from above, 2), 

4a. Ditto Head from the side, x 2 


—Comacupes cylindraceus (regd. no. or) Head from above, x 2. 


Si 

5a. Ditto Head from the side, x 2. 

6.—Another form of the same species (from a specimen in the British 
Museum). Head from above, x 2. 


6a. Ditto Head from the side, x 2 
7.—Taeniocerus bicanthatus (regd. no. “Y). Head from above, x 2. 
7a. Ditto Head from the side, x 2. 

7b. Ditto Mentum, etc., x 4. 

8.—Taeniocerus pygmaeus (regd. no. “°). Head from above, x 2 
8a. Ditto Head from the side, x 2. 
9.—Taeniocerus bicuspis (regd. no. >), Head from above, x 2. 
ga. Ditto Head from the side, x 2 


10.—Aulacocyclus andrewesi (from the type in Mr. Andrewes: collection). 
Head from above, x 2. 


roa. Ditto Head from the side, x 2 
I1.—Ceracupes austeni (type). Head from above, x2 
Ila. Ditto. Head from the side, x 2. 


12.—Ceracupes fronticornis (regd. no. °" a). "Head from above, x 2. 
13.—Pleurarius brachyphyllus (regd. no. $5). Head from above, x 2 


13a. Ditto Mentum, etc., x 4. 
14.—Tiberioides kuwerti (regd. no. Head from above, xX 2. 


15.—Tberioides austeni (type). Head os above, x 2. 


15a. Ditto Mentum, etc., 
16.—E pisphenus moorei (regd. no. sir), Sete from above, x 2. 
16a. Ditto Mentum, etc., x 4. 


17 Cee Pero (type). Head from above, xX 2. 
18. (regd. no. *). Head from above, X 2. 


18a. Ditto Mentum, etc., x 4. 
19.—E pisphenus comptoni var. flachi (regd. no. * and **'). Frontal area, 


etc., x 3, showing the variability of the chee of this area. 
Iga. Abnormal mentum of another specimen (regd. no. oss) X 4. 


20.—E pisphenus indicus large specimen (regd. no. *8). Head from above, x 2. 


20a. Ditto Mentum. eter act: 

20b. Same species, small specimen (from a specimen in the Bombay Natural 
History Society’s collection). Head from above, x 2. 

21.—Episphenus neelgherriensis (regd. no. 5). Head from above, xX 2. 


21a Ditto Mentum, etc., x 4. 
22,—Ophrygonius cantort (regd. no. ™""), Head from above, x 2. 


22a. Ditto Mentum, etc., x 4. 
23.—Ophrygonius cantort var. dunsiriensis (type). Mentum, etc., x 4. 


' E. moorei, see above, p. 28r. 


Mem. Ind. Mus., Vol. 11] += Plate XI 


Bemrose, Collo., Derby. 


D. Bagchi, del. 


ORIENTAL PASSALIDAE. 


EXPLANATION OF PLATE XII. 


24.—Ophrygonius inaequalis (regd. no. “). Head from above, x 2. 


24a. Ditto Mentum, etc., x 4. 


25.—Aceraius borneanus (regd. no. *"). Head from above, x 2. 


25a. Ditto Left mandible from the side, x 4. 

25. Ditto Mentum, etc., x 4. 

26.—Aceraius wallacei (regd. no. **). Head from above, x 2. 
26a. Ditto Left mandible from the side, x 4. 

26b. Ditto Mentum, ete.; x 

27.—Aceraius laevicollis (regd. no. ™8). Head from above, x 2. 
27a. Ditto Left mandible from the side, x 4. 


28.—Aceraius grandis subsp. hirsutus, a specimen of moderate size (regd. 
no. **'), Head from above, x 2. 

28a. A large specimen of the same species and subspecies (regd. no. a). 
Head from above, x 2. 

28). 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. 758) 
from above, x 4. 


30.—A ceraius grandis var. rectidens (from a specimen in the Sarawak Museum 
collection). Head from above, x 2. 

31.—Aceraius kuwerti (from a specimen in Mr. Andrewes’ collection). Head 
from above, x 2. 

32.—A ceraius illegalis (regd. no. **). Head from above, x 2. 

32a. Ditto Mentum, etc., x 4. 

33-—A ceraius méschleri (regd. no. **). Head from above, x 2. 

34.—A ceraius alutaceosternus (regd. no. =). Head from above, x 2. 

34a. Ditto Mentum, etc., x 4. 

35.—Aceraius pilifer (from a specimen in Mr. Andrewes’ collection). Head 
from above, x 2. 


36.—Aceraius himalayensis (type). Head from above, x 2. 


36a. Ditto Anterior part of head from the right side, x 4. 
37.—Aceraius assamensis (regd. no. ™*). Head from above. 

37a Ditto Anterior part of head from the right side, x 4. 
38.—A ceraius tavoyanus ‘ (regd. no. ®), Head from above, x 2. 

38a. Ditto Anterior part of head from the right side, x 4. 
39.—A ceraius helferi (regd. no. *“). Head from above, x 2 


394. Ditto Anterior part of head from the right side, x 4. 


' A. helferi, see above, p. 292. 


Mem. Ind. Mus., Vol. ll], 6sa= Plate Xl. 


D. Bagchi, del. Bemrose, Collo, Derby. 


ORIENTAL PASSALIDAE. 


<< 


iY 


a Ae 

: "i" 

j 7 ae - 

y gt les 

ip i i ae 
ne! 


- «7 
& > 
— 
; 
a ~ 
as 


EXPLANATION OF PLATE XIII. 


40.—Macrolinus nicobaricus (type). Head from above, x2. 

40a. Ditto Anterior part of head from the side, x 4. 
41.—Macrolinus andamanensis (regd. no. “*). Head from above, x 2. 
41a. Ditto Anterior part of head from the side, x 4. 
42.—Macryolinus sikkimensis (type). Head from above, x2. 

42a. Ditto. Mentum, etc., x4. 

43.—Macrolinus rotundifrons (regd. no. “°). Head from above, x2. 


44.—Macrolinus waterhousei (regd. no. **'), Head from above, x2. 


45.—Macrolinus latipennis (regd. no, *"!). Head from above, x2. 

46.—Another specimen of the same species (regd. no.**). Head from 
above, x2. 

47.—Gonatas germart (regd. no. *). Head from above, x2. 

A7a. Ditto Mentum , ete.; x 4: 

48—Trapezochilus dorsalis (regd. no. "). Head from above, x2. 

49.—Gnaphalocnemis burmeistert (regd. no. **). Head from above, x2. 

49a. Gnaphalocnemis monticulosus (tegd. no. *). Head from above, x 2. 

50.—Gnaphalocnemis tridens (from a specimen in Mr. Andrewes’ collection). 
Head from above, x2. 

50a. Ditto Mentum, etc., x 4. 

51.—Leptaulax humerosus (regd. no. **). Head from above, x 2. 

52.—Leptaulax dentatus, large specimen (regd. no. ™”). Head from above, 

Kg. 
52a. Ditto Mentum etc., x 4. 
52). Frontal areas of some specimens of the same species (regd. nos, “*, S| 


“°), all taken from one colony, x 2. 


19 
96 924 957 


52c. Frontal areas of specimens of the same species (regd. nos. “*, |) from 


other colonies, x 2. 

52d. Small specimen of the same species {regd. no. “*). Head from 
above, x2. 

53.—Leptaulax cyclotaenius, s. str. (regd. no. “'). Head from above, x 2. 

54.—Leptaulax macassariensis subsp. anibarbis (regd. no. °"). Head from 
above, x 2. 

55-—Leptaulax anipunctus (regd. no. “*). Head from above, x 2. 

56.—Leptaulax bicolor, s. str., large specimen (regd. no. “°). Head from 
above, x 2 

56a. The same species (var. vicinus), small specimen (regd. no. “”). Head 
from above, x 2. 

57.—Leptaulax roepstorfi (regd. no. **). Head from above, x 2. 


58.—Leptaulax planus (regd. no. **). Head from above, x 2. 


1914. 
Mem. Ind. Mus., Vol. 11], +e Plate X11 


D. Bagchi, del Bemrose, Collo, Derby 


ORIENTAL PASSALIDAE. 


i Ria Vol. VIII, 1912. 

‘y : fe ee 4 ine - Zoological Results of the Abor Expedition, IQII~12. 
Part I.—Introduction, Batrachia. Reptilia. Lepidoptera. Porifera. Scolopendridae. Hymenoptera. Anthophila. 
Part IJ,—Mammals. Coleoptera, I-III. Arachnida, I.° Dermaptera. Diptera. 


Part III.—Coleoptera, Iv-VI. Thysanoptera. Arachnida, II. Mollusca, I. Temmocephalidae. Hymenoptera, II, 
III. Fish. Birds. Crustacea Decapoda. Orthoptera, I. Freshwater Planaria. 


Part IV.—Hymenoptera, IV. Orthoptera, II. Odonata, Neuropteroid Insects. Reptilia (Supplement). Mollusca, 
II, Oligochaeta. hs 


' Part V.—Coleoptera, VII. Orthoptera, III. Termitidae. Lepidoptera, II. Hymenoptera, V, Crustacea Amphipoda, 


Wa Preit. 


Vol. IX, 1913. 


Part I.—Notes on the collection of Coccidae in the [ndian Museum, II. New Indian Empidae. 


Part II.—A new variety of freshwater crab from Travancore. On some Thysanura in the Indian Museum. The 
Tortoises of Chota Nagpur. Notes from the Bengal Fisheries Laboratory, Indian Museum, No. I. Cryptostomes 
from the Indian Museum. 5 


Part III.—Nouvelle étude sur les Chironomides de I’Indian Museum de Calcutta. Description de quelques nouvelles 
Cecidomyies des Indes. Description d’un nouveau Mymaride des Indes Orientales. The Anthribidae in the Indian 
Museum. Description of a new Sand-boa from the Persian Gulf. 


Payt IV.—A note on certain Ophiuroids in the Indian Museum. The Indian Barmacles of the subgenus Scalpellum. 
Notes on Freshwater Sponges, XV. Preliminary Note on a new genus of Onychophora from the N.-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 Limnocnida indica. 


Pavt V.—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 
Parnassius 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 
Assam 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. 


Payt II.—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. 


Part IJI,—On a new species of Blepharocerid fly from Kashmir, with a description of some larvae from the same locality. 
Studies in Indian Helminthology, No. I. Studies in Indian Helininthology, No. IZ. On some ney terrestrial Isopods 
from the Andaman Islands and Southern India. Miscellanea :—Note on the genus A nactinia. Metamorphosis of 
Zoanthella, Change of name ig an Indian genus.of Echinoids. Note on some Amphipods collected on the Pamirs 
at an altitude of 15,000 ft. NXenapsylla nesiotes : a correction. 


ya 


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INDIAN “MUSEUM 


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collection in the Indian Museum. By x H. GRAVELY. -Rs. 5. 


Vol, IV. Rie 


No. 1.—An Account of the Crustacea Stomatopoda of the Indo-Pacific Region based Regen 
on the collection in the Indian Museum. | By STANL EY Kemp. Rs. 15." | 7.4 bis 


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