okyo, 27 (2): 463-469, November 13, Elyt1・a, 「okyo, 27 (2): 463-469, November 13, l 999

The D ung Beet le Fauna (Coleoptera, The Dung Beet le Fauna (Coleoptera, ) of N ortheast Northeast Thai land

Y upa Yupa HANB00NSONG Serumal CHUNRAM

Department of Depar tment of Entomol ogy, Department of Agriculture, Ministryof Faculty Faculty of Agriculture, Agricul ture and Cooperatives, Khon Khon Kaen Universi ty, Ban gko k, 190 00 Thailand, Khon Kaen, 40002 Khon Kaen, 40002 Thailan(i,

R owan Su「a p I M P A S A L E E Rowan W. EMBERSON

Department of Department of Agriculture, Ministry of Department of Entomol ogy and Agriculture and Cooperatives, Animal Ecology, Lincoln Un iversi ty, Canterbury, Bangkok, 19000 Thailand, Canterbury, New Zealand

and

Kimio Kimio MASUMOT0

Institute of Human Living Sciences, 0tsuma Women's Institute of Human Living Sciences, 0tsuma Women's University, Tokyo, 102 -8357 Tokyo, 102 -8357 Japan

A list of the scarabaeine dung collected Abstract A list of the scarabaeine dung beetles collected from bot h cul ti vated and forested areas in Northeast Thailand is presented. Al together 154 and forested areas in Northeast Thailand is presented. Al together 154 species of 15 genera in 7 in 7 tribes are recorded. The genus Onthop/1agus is the most diverse group in wh ich 103 The genera , Cassolus, Panehls, P11acosoma and species were found. The genera Caccobius, Cassolus, Panehls, P11acosoma and Sisyphus are mainl y confined to forested habi tats, while the genera Omtts and are mainl y confined to forested habi tats, while the genera Omtts and Onl tice11us occur very commonly in the dung of domestic . The ecological habitat of each dung commonly in the dung of domestic animals. The ecological habitat of each dung species found is also species found is also recorded.

Introduction beetles (Coleoptera, Scarabaeidae) are one of the most The dung beetles (Coleoptera, Scarabaeidae) are one of the most beneficial groups of ever known. With few exceptions, they feed as larvae and adults groups of insects ever known. With few exceptions, they feed as larvae and adults on the dung of the dung of vertebrates. In In spite of the important ecological and beneficial roles of dung beetles, very little is is known about the dung beetles of Thai land as compared to those o f other regions. 0n the the world basis, classification and distribution of the Scarabaeidae, above the gener ic level, level, is generally better known than those of most other comparable sized groups of W orldwide insects. Worldwide there are nearly 5,000 described species arranged in 234 genera Yupa 464 Yupa HANB0 0NSONG et al and 10-12 t ribes (HANsKI & CAMBEFoRT, 1991 ). This wel l developed understand ing and 10-12 t ribes (HANsKI & CAMBEFoRT, 1991 ). This wel l developed understand ing o f the the group is no doubt due to their economic importance and the spectacular morpho- logical modifications of many species. 0fa11 the world's regional faunas, that of logical modifications of many species. 0fa11 the world's regional faunas, that of South- east Asia is probably the poorest known (HANsKl & KRIKKEN, east Asia is probably the poorest known (HANsKl & KRIKKEN, 1991). Records of species occurring in Thailand have been included in the three Records of species occurring in Thailand have been included in the three classic regional regional faunas covering parts of Southeast Asia (ARROW, 1931; PAULIAN, 1945; BALTHAsAR, 1963 a, b,1964). More recently, MAsUMoTo (1987, 1988, 1989 BALTHAsAR, 1963 a, b,1964). More recently, MAsUMoTo (1987, 1988, 1989 a-c, l990, 1991, 1992 a, b, 1995, 1996) surveyed the fauna o f the Scarabaeidae and 1991, 1992 a, b, 1995, 1996) surveyed the fauna o f the Scarabaeidae and Aphod iidae in in Northwest Thailand and STEBN!cKA (1992) published a summary, including previous publ ished records, o f the Aphodi idae known from Thailand. However, no wor k or publ ished records, o f the Aphodi idae known from Thailand. However, no wor k or pub- lication of dung beetles from Northeast Thailand has been made lication of dung beetles from Northeast Thailand has been made before. The No r theast The No r theast Region of Thailan usually called Isan, covers approximately 170,000 square kilometers or about 1/3 of the country. Most of the area is a 170,000 square kilometers or about 1/3 of the country. Most of the area is a high plateau with a height between 150-1,000 metres above sea-level. The boundaries plateau with a height between 150-1,000 metres above sea-level. The boundaries of Northeast are connected with the Lao border to the north and the east, with Northeast are connected with the Lao border to the north and the east, with Cambodia and the Centre Region to the south, and wi th the lower region o f the North to the and the Centre Region to the south, and wi th the lower region o f the North to the west. B iodiversity Currently, we have a fund from th e Thai land Biodiversity and Training Pro- gramme gramme to conduct a one year pilot study of the fauna of the Northeast Region of Thailand. We hope to be able to extend this to a full survey o f the fauna Region of Thailand. We hope to be able to extend this to a full survey o f the fauna o f Thai land.

Methods Nineteen Nineteen provinces in the Northeast Region were surveyed in both cultivated and forested forested areas, Natural Parks and Wildli fe Sanctuaries to sample the dung beetle fauna. Samp l ing was carr ied out throughout the year in several di fferent ways such as Samp l ing was carr ied out throughout the year in several di fferent ways such as hand searching searching directly on the dung pads, light trapping (only the dung beetles that are noc- turnal turnal and positively phototactic were caught) and pit fall traps baited with pig dung, rotten rotten chicken, etc. Forest leaf litter was also collected and extracted with Berlese fun- nels to samp le the smal l l i t ter dwel l ing nels to samp le the smal l l i t ter dwel l ing species. Specimens Specimens have been sorted into morphological species, or recognizable taxo- nomic un i ts, by eye or using binocular magn i f iers. I den ti ficat i ons have then been nomic un i ts, by eye or using binocular magn i f iers. I den ti ficat i ons have then been made using rel iably i denti f ied materials in the Museum at the Depar tment of using rel iably i denti f ied materials in the Insect Museum at the Depar tment of Agr icu l- ture ture in Bangkok, and MAsuMoTo's collection in Yokohama. Some specimens were identified by comparing with the types and other materials preserved in the identified by comparing with the types and other materials preserved in the Natural History History Museum, London, and Museum National d'Histoire Nature11e, Paris_Others were were determined from literature.

Results and Results and Discussion A A total of 154 dung beetle species of the Scarabaeidae,of 15 genera in7 tribes, were were found in the Northeast Region (Table t). This compares wi th 137 species of 22 Dung Beetle Fauna of Northeast Dung Beetle Fauna of Northeast Thailand 465

genera col lected by MAsuMoTo in Northwest Thai land. Most dung beet le species genera col lected by MAsuMoTo in Northwest Thai land. Most dung beet le species are coprophagous, wh i le a few species such as tr icornls WIEDEMANN, coprophagous, wh i le a few species such as Onthophagus tr icornls WIEDEMANN, etc., are are necrophagous wh ich feed on animal carr ion, and Onthophagus bonarae ZuNINo, Cassolus nudus SHARP, etc., feed on rotting vegetable Cassolus nudus SHARP, etc., feed on rotting vegetable matters. R ichness Richness o f dung beetle species in the forested areas is greater than in the lowland or or cultivated areas. This is possibly due to an abundant supp ly of food from animal biomass and vegetation in the forested areas where the areas are well preserved biomass and vegetation in the forested areas where the areas are well preserved and protected for faunal and floral habitats. In contrast with the cultivated ar ea s of the Northeast, there is a progressive decrease of cultivated farmlands and the domestic Northeast, there is a progressive decrease of cultivated farmlands and the domestic ani- mals like cows and buffalos are gradual ly reducing number and being replaced by mals like cows and buffalos are gradual ly reducing number and being replaced by ma- ch in e. O f the 18 genera reported here, the genus Onthophagus, Of the 18 genera reported here, the genus Onthophagus, mainly foun d in the forested areas, is the most diverse group, of which 43 species were identified and forested areas, is the most diverse group, of which 43 species were identified and ap- prox imately 60 species are sti l l unknown. There are, almost certainly, a prox imately 60 species are sti l l unknown. There are, almost certainly, a substant ial number o f add it iona1 0nt hop hagus species, in other forested hab i tats o f number o f add it iona1 0nt hop hagus species, in other forested hab i tats o f Thai land. Therefore, further work should be carried out for this Therefore, further work should be carried out for this group. The study has also shown that some dung beetle genera are restr icted to one The study has also shown that some dung beetle genera are restr icted to one k ind of habitat, such genera as Catharsius, 0nltis and Onltice11us are dominant in the of habitat, such genera as Catharsius, 0nltis and Onltice11us are dominant in the low- land or cultivated areas while other genera like Caccobius, Cassolus, Panelus, land or cultivated areas while other genera like Caccobius, Cassolus, Panelus, Phaco- us are confined to forested habitats. Since the genera Omtfs and soma and Szs us are confined to forested habitats. Since the genera Omtfs and Omtz- ce11u.s・occur very commonly in the dung of domestic animals like cow and buffalo, the local people in the Northeast use the dung beetles as a food protein source. local people in the Northeast use the dung beetles as a food protein source. Particu- 1arly, 0nltice11us 1arly, 0nltice11us cinctus (FABRlclUs) is the most favourite edible dung beetle species. In In conclusion, this study is the first report of dung beetles from the Northeast o f Thailand and the data were accumulated only from one year study. However, we Thailand and the data were accumulated only from one year study. However, we confi- dently expect the fauna of Northeast to be significantly larger than that dently expect the fauna of Northeast to be significantly larger than that documented here. With a comprehensive sampling programme, using various baited pitfall here. With a comprehensive sampling programme, using various baited pitfall traps and litter extraction, in different forested areas, the number of identified species and litter extraction, in different forested areas, the number of identified species will, we are sure, increase we are sure, increase significantly. In In addition to the Scarabaeidae, material of the Aphodiidae and Geotrupidae are also being accumulated for future reference. We have material of about a also being accumulated for future reference. We have material of about a halfof the19 species species of the Aphodi idae currently known from the Northeast of Thai land (STEB- N1cKA, l992) N1cKA, l992) and at least three species of the Geotrupidae have been collected.

Yupa 466 Yupa HANB00NSONG e1 al .

Dung Table t . Dung Beet les from Nor theast Thailand

Spec ies Fo o d so urces Habitats

Tribe Tribe Gymnopleurini cow, Gymnopleurus (P,a'agymnop leut'us) cow, pig, li ght trap cul tivated area '11ela na' ' ius '11ela na' ' ius H AROLD G. G. (P ) r ud is SHARP Pl9 cul tivatedarea G. G. ( P ) s lml a tus OLI VIER bar king deer, pig forested area cow, pig, light (1. (Garreta) ruficornis MoTscHULsKY cow, pig, light trap cul tivatedarea Tribe Tribe Sisyphini 99 99 Sfsyp Ms ' n a m f1 M A s U M o T 0 Ip ' Ip ' Ip forested area S. S. / 0n g l p eS ( 0 L IVI ER) forested area Sisyphus 1 sp. Sisyphus 1 sp. indef. forested area Tribe Tribe Canthonini Panehls to' lki nensls PAULIAN wild cat forested area P/1acoso,na fa11ac11aetum P/1acoso,na fa11ac11aetum MAsuMoTo wi ld cat, jackal forested area gaur, Casso lus nud us SHARP gaur, mushroom forested area Tribe Tribe Coprini Synapsfs birnlam'ctis Synapsfs birnlam'ctis GILLET Pl 9 forested area Synapsis 2 Synapsis 2 spp. indet. pig forested area cow, He11ocopt ls bucep/1alus (FABRlclUs) cow, buf falo, gaur, banteng cultivated & forested areas ao'm'IMs BATES elephant forestedarea buffalo, cow, elephant, Catharsi ils birmanens is LANsBERGE buffalo, cow, elephant, pig cultivatedarea buffalo, cow, gaur, pig, l ight C. JaVamls LANSBERGE buffalo, cow, gaur, pig, l ight trap cultivatedarea buff;ale, C m o t o s s u s ( L I N N E ) buff;ale, elephant cultivated & forested areas Cop ' ' is Cop ' is an g l ls f f co r川's ARROW elephant forested area C. C. Ca「1n lCt lS G I LLET gaur forested area elephant, C. CO「pule'1tus GILLET elephant, gaur forested area buffalo, CI「IS SHARP buffalo, elephant cultivated & forested areas cow, samber deer, elephant, pi g, C. l aev ig atu.s' GILLET cow, samber deer, elephant, pi g, l ight trap cultivated & forested areas l'inSOni C ' t o 、l'inSOni WATERHOUSE buffalo, cow, pig, light trap cultivatedarea C. sf Mc l es HOP pig cultivatedarea elephant, gaur, C. (M i c ' 'oc〔)pr is) 、レ 'italis1 GILLET elephant, gaur, banteng forested area cow, buf falo, pig, gaur, C. (M )1eflexus (FABRIclUs) cow, buf falo, pig, gaur, elephant, light trap cultivatedarea C. C. ( pa ''ac e - s ) ca r加cops FELSCHE elephant, pig forested area elephant, C. ( p ) pu11Ctu latus WIEDEMANN elephant, cow cultivated & forested areas C. ( p ) f ut c i ceps C. ( p ) f ut c i ceps FELSCHE elephant forested area Cop r is 2 Cop r is 2 spp. indet. elephant forested area Tribe Tribe Onitini hhbb fl t Pe W W eP eP a n f OMf fs 0 ''da「f C A M B E F O R T eeoc forested area r a nuu t, a u 0. 0. excavatus ARROW 9la la forested area 99 19 oc Pli o 0. 0. vil'e'Is LANSBERGE , cul ti vatedarea h p a o ffs3 spp. 0川ffs3 spp. indef. , cultivatedarea Dung Beetle Fauna of Northeast Dung Beetle Fauna of Northeast Thailand 467

Table t. (Continued)

Species Foo d sou rces H ab i tat s

Tribe Tribe Oniticeli ini D' 'epanoce' '11s D' 'epanoce' '11s st' '1atulus PAULIAN COW cul tivated area On i l ice11us On i l ice11us ci ncttls (FABRICIUS) cow, buf falo, gaur cul tivated area cow, 0. tesseliatus HAROLD cow, buff;ale forested area 0. ft''eyi 0. ft''eyi BAt_THASAR cow, buff;ale cul ti vated area cow, Onitice11ils 2 spp. indet. cow, buffalo forested area 1a f ong us 1a f ong us a 川s(ARROW) elephant, gaur, deer forested area L L g a g a t i n u s ( H O P E ) elephant, gaur forested area

. /1anaeo ldes (WESTWOOD) pig forested area cow, L. ''/1adatnlstus (FABRICIUS) cow, buffalo, pig cul ti vated area elephant, L t1'identatus(BoUcoMoNT) elephant, gaur forested area L. L. venato1 (FABRl clUs) COW forested area gaur, L. vertagus (FABRICIUS) gaur, pi g forested area Tribe Tribe Onthophagini Caccobius (Caccophi11ls) monkey, bird, pig, cow forested area llMc0' '11s llMc0' '11s FABRICIUS dddddddddd et et et et ete te te te te te te te te te fffffff sssss ooo er er re re re re re aaaaaaaa er er er er er eaar er ere re re re re a a re re re re re re re re re re re a 011thophagus 011thophagus act'opict tls BOUCOMONT Pl 9 aaaaaaaaaa 0. pi iular it ls 0. pi iular it ls LANsBERGE pig elephant, wild cat, 0. ba lthasa''i VsETETKA elephant, wild cat, pig pig, rotten passion 0. bone ' ae Z UNIN0 pig, rotten passion fruit oo ooo ooo 0. b''utus 0. b''utus ARROW pig rr 0. c0'acinus 0. c0'acinus BOUCOMONT pig 0. 0. c''ass1co//fs BOUCOMONT mon key f f fff 0. dapcauensis 0. dapcauensis BoUcoMoNl pig rr ol'pll1ensfs 0. ol'pll1ensfs MAsUMoTo pig a r mon key aadd 0. 0. dupe''t1 BoUCOMONT t rlt 99

jackal, jackal, wild cat 3 11 0. 0. g,'acif!'pes BoUcoMoNT a a aa V a c uu cow, buffal o, gaur, li ght trap, 0. /1asflfer LANSBERGE cow, buffal o, gaur, li ght trap, banteng re re lt V a cow, cow, pi g c c c f f f f c c cf f f f f cf tt

0. 0. Jaccobells BoUcoMoNT d ad aaad aa oo o oo oo rr lr rl rr rr aaaa aa elephant, 0. /aevls HAROLD elephant, pig wild 0. lindaae MAsUMoTo wild cat, monkey, sliver pheasant t

cow, deer, porcupine, 0. luridipennis BoHEMAN cow, deer, porcupine, pig tt 0. 0. nlamfi MAsUMOT0 pig 0. 0. naa1'con MASUMOTo pig ・ 9 re V a u a cow, 0. olientalis HAROLD cow, buffalo, monkey, pig, light trap - t a o re

pig ・ 11f re

0. 0. pacjf tcus LANSBERGE arerere V uu cow, cow, buf falo, pig, l ight trap - 0. papulatus 0. papulatus BoUCOMONT ・ & V u S

cow, cow, buffalo, pig, l ight trap d e e

0. pr()fetal f us 0. pr()fetal f us HAROLD a a a ar o 「 ttt tt oo oo aaa 0. 0. ph,・utsap/1akho'mls MAsUMOT0 rot ten banana, pi g rr 0. ''ud is 0. ''ud is SHARP pig 0. 0. sa''awakus HAROLD pig rr111 addd a eaaa a a re o elephant, wi ld cat, 0. taur inus WHITE elephant, wi ld cat, pig

・ r rr VVV uuu cow, buf falo, pi g, li ght cow, buf falo, pi g, li ght trap u 0. 0. t''ag iodes BoUcoMONT ・ 0. 0. t''1tuber (WIEDEMANN) - monkey, bu f fal o, monkey, bu f fal o, pi g ・- buffalo, banteng, gaur, pig, light 0. (Co1obont /1op/1agus) buffalo, banteng, gaur, pig, light trap t''agus (FABRICIUS) Yupa HANB00NSONG et 468 Yupa HANB00NSONG et al

t. Table t. (Continued)

Species Food sources Habi tats 0. (0iglfoMf/1ophagl's) elephant, cow, buffalo, gaur, banteng cu lti vated area bonasus (FABRI cIUs ) 0. 0. (Indac11o,・lus) dolsut/1epensls Pl 9 forested area MASUMOT0 0. (Mafas/1l'a) 0. (Mafas/1l'a) angll/fceps elephant, pig forested area BOUCOMONT 0. (M ) avocetta 0. (M ) avocetta ARROW Pl9 forested area (M ) 0. (M ) 'nam'pill'eMsfs ARROW pig forested area 0. 0. (Mic,-ont/1ophagus) mon key, dog forested area falslvlgilans falslvlgilans MASUMoT0 0. 0. (.Pataphanaeo'110'p;1us) Pl9 forested area eane/ ffamls eane/ ffamls PAULIAN 0. (pa''ascatono'mls) topali 0. (pa''ascatono'mls) topali ENDRODI carrion trap forestedarea cow, pig, dead snake, li ght 0. (p ) triCOrnls (WIEDEMANN) cow, pig, dead snake, li ght trap cul tivated & forested areas 0. 0. (Proagoalerus) 'n011hot1' HAROLD cow, buf fal o cul tivated area op h o ,-us ) 0. (So,,・op h o ,-us ) 1・ecteco,・nutils bu f falo, l i ght t rap cul tivated & LANSBERGE forested areas 0. 0. (S) sagitta''1us(FABRICIUS) cow, buf fal o, banteng, gaur, li ght trap cul tivated area cow, 0. (S) senlculus(FABRICIUS) cow, buf fal o, e lephant, banteng, pi g, cul tivated area light light trap Onthop;tagus 60 spp. Onthop;tagus 60 spp. indef pig, cow, buffalo, jackal, l ight trap forestedarea

*Though the species on the list is mainly arranged according to the *Though the species on the list is mainly arranged according to the BALTHAsARsystem (1963), some changes are given as regards subgeneric names and species descri bed after changes are given as regards subgeneric names and species descri bed after l t.

Acknowledgments We We would like to acknowledge the grant received from the Thailand Biodiversity Research and Training Programme. We also thank Dr. Angoon LEwvANlcH in the In- sect Museum at the Department of Agriculture, Bangkok, for arranging access to the col lect ion of dung beet les and using facil i t ies. Deep indebtedness should be col lect ion of dung beet les and using facil i t ies. Deep indebtedness should be expressed to to Dr. Yves CAMBEFoRT, Museum National d'Histoire Nature11e, Paris, and Mr. Ma1- colm KERLEY, the Natural History Museum, London, for permi tt ing us to colm KERLEY, the Natural History Museum, London, for permi tt ing us to examine types types and other materials for determination.

要 約 Y. HANB00NSONG ' S. CHUNRAM ' S. PIMPAsALEE ・ R. W. EMBERsoN ・益本仁雄 :東北タ イの食 糞性コガネムシ相. - 東北 タ イ に分布す る食糞性コ ガネム シ (Scarabaeidae) について , 耕 . 作された地域と自然林の両方で採集をおこなった. その結果, 合計 7 族, 1 5 属, 154種がみと められた. そのう ち, エンマコ ガネ属 (0nthophagus) がもっ とも種数が多く 103種である. コ Cassolus, Panelus, P/1acosoma, エンマコガネ属 (Caccobius) およびCassolus, Panelus, P/1acosoma, Sisyphus などの属はおもに自 Dung Beetle Fauna of Northeast Dung Beetle Fauna of Northeast Thai land 469

, 然林内に生息 し , O n i t is 属, O m t i c e 1 1 u s 属などは, 家畜の英にきわめて普通にみられた . 添付 し , た種名一覧表には, 採集 し た 糞虫が来集し た糞の種類 およ び生息地域 を併記 し た.

References

C oleoptera, ARROW, G. J. , 193 1 . Coleoptera, Lamellicornia, part III, Coprinae. In tile: Fauna of British India, In- ,na 428 pp. Taylor and Francis, cluding Cey1o,1 and But・,na 428 pp. Taylor and Francis, London. BALTHAsAR, V., 1963. Monographic der Scarabaeidae unci Aphodi idae der Palaearkt ischen und Orientali- Region (Coleoptera; sc he n Region (Coleoptera; Lame11icornia). 1. Scarabaeidae 392 pp., 24 pls. Tschechos1ovakia Akademie Akademie Wissenschaft, Prag. - l963. Ditto, 11.628 pp.,16 -l963. Ditto, 11.628 pp.,16 pls. FINcHER, G. T., 1973. Dung beetles as biol ogical cont rol agents for gast rointestinal parasi tes of livestock. J. J. Parasito1., 59: 396 -399. HANsKI, 1., & J. KRIKKEN, 1991. Dung beetles in tropical forests in South-East Asia./n HANsK1, 1. , & Y. CAMBEFoRT (eds ), Dung Beetle Ecology, 179 -197. Princeton Uni versity CAMBEFoRT (eds ), Dung Beetle Ecology, 179 -197. Princeton Uni versity Press, New Jersey. - & Y. CAMBEFORT,1991.Species richness.In HANSKI, 1., & Y. - & Y. CAMBEFORT,1991.Species richness.In HANSKI, 1., & Y. CAMBEFORT(eds), Dutlg Beetle Ecology, 350 -365. Ecology, 350 -365. Princeton Universi ty Press, New Jersey. MAsUMoTo, K., l987. Coprophagid-beetles from Northwest Thailand (I). Ent. Rev . Japan. , Osaka, 42: 125- 131 125- 131 . 1988 . D it to (I I ). 一 1988 . D it to (I I ). Ib id., 43: 135- 143. - 1989a. -1989a. Ditto(III). Ibid.,44:125-131. Coprophagous scarabaeids related to 0,1thophagus -1989b.Coprophagous scarabaeids related to 0,1thophagus or ienta lis (Coleoptera, Sca rabaeidae ) Elytra, from Sout heast Asia. Elytra, Tto 0, 17: 175-180. om -1989c. Coprophagid-beetlesfi・om Northwest Thailand (IV). E lf. Rev. Japat1, Osaka, 44 : 87 - 96. 1990. Ditto (V). 一 1990. Ditto (V). Ibid., 45: 43-51. - 1991. -1991. Ditto(VI). Ibid., 46:27-37. - 1992 -1992 a. Ditto(VII). 1,bid.,47:35-41. - 1992 b. Ditto(Vm). -1992 b. Ditto(Vm). Ibid.,47:115-117. - 1995. Ditto(IX). -1995. Ditto(IX). Ibid.,50:27-37. - 1996. Ditto(X). -1996. Ditto(X). 11bid.,50:87-94. PAULIAN, R., 1945. Coleopteres Scarabeides de l ' Indochine. F◆taune de 1'E,1lp i・ ' eFta'1?ats,3.228pp. &1 map. La rose, map. La rose, Pa ri s. STEBN1cKA, Z., 1992. Aphodi inae from Thai land (Coleoptera; Scarabaeidae). Stuttg. Be l t r. Natut'k., (A), (481) : 1 -16.