SERIES OF MISCELLANEOUS PUBLICATIONS UNIVERSITY OF AMSTERDAM. Atherurus F. Cuvier, D.J. van Weers. Abstract INTRODUCTION

Iformatio about this material is preseted ad the status of te specific ad subspecific ames Beaufortia SERIES OF MISCELLANEOUS PUBLICATIONS INSTITUTE OF TAXONOMIC ZOOLOGY (ZOOLOGICAL MUSEUM) UNIVERSITY OF AMSTERDAM No. 6 Volume 6 December 0 1977 Notes o Southeast Asia Porcupies (Hystricidae Rodetia) II. O the taxoomy of the geus Atherurus F. Cuvier 189 D.J. va Weers Abstract The idividual ad geographical variatio of the Brush-tailed PorcupieAtherurus F. Cuvier 189 from easter Idia Burma Thailad Laos Chia Haia Peisular Malaysia ad some adjacet islads has bee studied from material i Europea ad America museums. is discussed. Oly oe species Atherurus macrourus (Liaeus 178) is recogized ad isufficiet argumets were foud for the distictio of subspecies o the basis of the small samples at preset available. A error i Va Weers (1976: 19) cocerig the ame of the type species of Trichys Güther 1877 is corrected i a foot ote. INTRODUCTION Seba (174: 84 pi. figs. 1 ad ) described ad figured the Brush-tailed Porcupie "Porcus aculeatus sylvestris seu Hystrix Orietalis sigularis" thereby uambiguously characterizig the peculiar tail brush by comparig the swelligs i its hairs to grais of rice. Liaeus (178: 7) gave the ame Hystrix macroura to this taxo quotig Seba thus: "Porcus aculeatus s. Hystrix orietalis. Seb. mus. 1. p. 84. t.. f. 1. Habitat i Asia." F. Cuvier (189: 48) used the geeric ame Atherurus for these porcupies with origi "Des Ides orietales" ad described them as follows: "... leur queue est logue et termiee par u faisceau de laieres corees aplaties et etraglees d'espace e espace de maiere a represeter u chapelet" which correspods with Seba's descriptio. Lyo (1906: 199) stated that Hystrix macroura Liaeusis the type species for the porcupie with the beaded tail bristle as it was the oly available ame at the time of Cuvier's descriptio ad the same author (Lyo 1907: 84) restricted the type locality of macroura to Malacca. Received: May 18 1977 0

British Rijksmuseum The The Craial The 06 This species lacks a holotype ad idividual variatio withi it is poorly uderstood. Util ow there have bee oly a few Malaya specimes scattered over some Europea ad America museums. O occasio oly oe arbitrary specime from the Malay Peisula has bee available for compariso with samples from other localities i Southeast Asia ad i other cases ewly described species were based o o more tha oe sigle specime. This has led to the descriptio of a umber of doubtful species ad subspecies. Mohr (1964) brought the umber of ames amog Asiatic Atherurus to te with the publicatio of two ew oes. She distiguished three species ad quoted eight subspecific ames whereas Ellerma (1961) recogized oly oe species with two subspecies at the same time statig: "Probably if a large umber of specimes for this geus were collected it would be impossible to divide it ito subspecies". The aim of the preset paper is a critical examiatio of the status of these species ad subspecies ad to preset some iformatio about the morphology of these taxa as far as is possible with the material of Europea ad America museums that has bee available to me. MATERIAL AND METHODS Sources. list of specimes studied comprises skis ad 9 skulls of adult specimes ad 14 skis ad 16 skulls of o-adult oes available from the collectios ofthe istitutios metioed below. AMNH America Museum of Natural History New York. BMNH Museum (Natural History) Lodo. FMNH IRSNB Field Museum of Natural History Chicago. Istitut Royal des Scieces Naturelles Bruxelles. MNHN Museum Natioal d'histoire Naturelle Paris. MSNG Museo Civico di StoriaNaturale di Geova Geoa. RMNH va Natuurlijke Historie Leide. USNM Natioal Museum of Natural History Washigto. UZMC Uiversitetets Zoologiske Museum Copehage. ZMA Zoologisch Museum Amsterdam. Geographical areas. area of distributio of the specimes studied rages from Assam i Idia to Vietam ad from the Malay Peisula to Szechwa i Chia. To eable a aalysis of the data gathered a umber of geographical areas based o the distaces ad geographical barriers betwee groups of localities have bee distiguished ad are preseted i table I. Detal age. detal age groups adult subadult ad youg are defied by Va Weers (1976: 17). The umber of o-adult specimes is too small to eable comparisos so that oly data for adult specimes appear i the tables ad histograms. M easuremets. ad exteral measuremets are defied by Va Weers (1976: 17 19). These are the same as those used i the preset

07 paper except the defiitio of the tail bristles. I Atherurus these are highly modified flatteed hairs alterately expaded ad cotracted from oe to six times. Table I. Geographical areas with locality umbers (see map fig. 1) as used i the aalysis data gathered ad umber of adult specimes studied. of the Geographical area Loc. umbers Skis Skulls Malay Peisula 1 4 6 Pulau Terutau 1 1 Pulau Peag 6 1 Pulau Aur 7 7 8 Pulau Pemaggil 8 1 1 Pulau Tioma 9 4 Teasserim Burma 10 1 1 Souther Laos 11 1 Aam Vietam 1 Thailad 141 1 Cetral Laos 1617 Toki Vietam 180 6 6 Haia Chia 1 6 6 Norther Burma 7 6 6 Assam Idia 8 1 1 Wahsie Chia 9 1 1 Ukow localities 7 SPECIMENS EXAMINED The umbers before the localities i this list of specimes examied correspod with the umbers of the localities o the distributio map ad i the histograms ad tables i the preset paper. The groupig ofthe localities i geographical areas is the same as preseted i table I. The coordiates are give i the Gazetteer. Abbreviatios: HB legth of head ad body T1 legth of tail Hf legth of hidfoot E ear Wt weight. MALAY PENINSULA 1. Skeats camp G. Taha Malaya BMNH.06 skull adult coll o... Provice of Wellesley Malay Peisula BMNH 4.10..8 ski ad skull adult rf E. J. Comp. o. 880 b Dr. Cator. UZMC M-49 ski ad skull adult Aschlud 7-1X-1848 o.. UZMC M-49 ski ad skull youg Aschlud 7-IX-1848 o. 4.. Jalor Malay Peisula BMNH..6.7 ski ad skull adult H. C. Robiso ad N. Aadale o. 18. 4. Trog Malay Peisula USNM 844 ski ad skull adult 9 W. L. Abbott -1-1897 HB 17. T1 9 Wt 9.7 lbs. uterus cotaied oe fetus 1. ich i legth.

08 Figure 1. Rage of the Southeast Asiatic Atherurus forms as determied by specimes examied. The umbers of plotted localities correspod with the umbers i the list of specimes examied ad with those i the histograms of the figs. ad 4 ad the tables I ad III. USNM 49498 skull ad skeleto adult 96-11-1897. USNM 800 ski ad skull subadult 9 W. L. Abbott -XI-1896 legth to aus 19 tail from aus 8 Wt 7 lbs. Malacca RMNH 000 ski Diard( Jetik's 1888 Atherura macrura o. b).

09 PULAU TERUTAU. Pulo Terutau USNM 1971 holotype of Atherurus terutaus Lyo 1907 ski ad skull adult <y W. L. Abbott 10-1V-1904 o. HB aus 440 T1 from aus 110 Wt 6.76 lbs. PULAU PENANG 6. Pulo Piag UZMC M-494 skull adult Dr. Cator 16-VII-1846. UZMC M-491 ski ad skull subadult Dr. Cator 16-VII-1846 o 0. BMNH 79.11.1.69 ski ad skull subadult Dr. Cator880c o. 4. BMNH 60..4.76 ski ad skull youg 9 E. Idia House Coll. PULAU AUR 7. Pulo Aor South Chia Sea USNM 1149 holotype of Atherura zygomatica Miller 190 ski ad skull adult 9 W. L. Abbott 6-VI-1901 HB 0 mm T1 00 mm Wt 9 lbs. USNM 1141 paratype of A. zygomatica Miller W. L. Abbott 7-VI- 1901 o. 1014 HB 00 mm T1 from aus 14 mm Wt 10 lbs. USNM 114 paratype of A. zygomatica Miller ski ad skull (sawed through legthwise) adult W. L. Abbott 7-VI-1901 HB 48 mm. USNM 1144 paratype of A. zygomatica Miller ski ad skull adult W. L. 9 Abbott 7-VI-1901 o. 10 HB 00 mm. USNM 4960 paratype of A. zygomatica Miller skull ad skeleto adult W. L. Abott 8-VI-1901. BMNH 8.1..0 ski ad skull adult 9 H. C. Robiso 16-VI-1906 F.M.S. o. 6/06 HB 16 T1 09 Hf 64 E 6. BMNH.08 ski ad skull adult J 1 H. C. Robiso 1-VI-191 o. 017 F.M.S. o. 47/1 HB 47 T1 190 Hf 68 E 7. BMNH.07 ski ad skull adult H. C. Robiso o. 4886 F.M.S. o. 49/1 HB 469 Hf 68 T1 1 E 6. USNM 1140 paratype of A. zygomatica Miller ski ad skull subadult 9 W. L. Abbott 6-VI-1901 o. 1010 HB 470 mm T1 190 mm Wt 7.7 lbs. PULAU PEMANGGIL 8. Pulau Pemaggil South Chia Sea BMNH 49.4 holotype ofatherurus macrourus pemagilis Robiso 191 ski ad skull adult Robiso o. 014 F.M.S. o. 41/1 HB 46 Hf 64 E 4. 9 H. C. BMNH.1 paratype of A: m. pemagilis Robiso ski ad skull subadult H. C. Robiso F.M.S. o. 44/1 HB 487 T1 0 Hf 67 E 4.. PULAU TIOMAN 9. Juara Bay Pulau Tioma South Chia Sea BMNH 8.1..1 holotype of

10 Atherurus tiois Thomas 1908 ski oly skull assumed to be lost H. C. Robiso 1-VI-1906 F.M.S. o. /06 HB 16 T1 176 Hf 66 E. BMNH 8.1.. paratype of A. tiois Thomas skull oly ski i the collectios of the Uiversity of Sigapore adult 9 H. C. Robiso o.. BMNH 8.1..1 paratype of A. tiois Thomas adult J H. C. Robiso o. 4. BMNH.09 ski ad skull adult <f H. C. Robiso 79 F.M.S. o. 48/1 7-V1-191 HB 78 T1 18 Hf 6 E 4. BMNH.11 ski ad skull adult 9 H. C. Robiso 4-V1-191 o. 661 F.M.S. o. 440/1 HB 44 T1 148 Hf 68 E 8. BMNH.10 ski ad skull adult <j\ H. C. Robiso 4-VI-191 o. 661 F.M.S. o. 44/1 HB 4 Hf 68 E 4. TENASSERIM BURMA 10. Thougyah Teasserim BMNH 8.8.1.40 ski ad skull adult <$ J Darlig Alle O. Hume. SOUTHERN LAOS 11. Plateau Boloves Laos AMNH 878 ski ad skull adult 9 T. D. Carter o. 448 leg. Ido-Chia Exped. 1-II-19 total legth 70 T1 0 Hf 60. AMNH 8786 ski ad skull adult T. D. Carter o. 449 leg. Ido-Chia Exped. 1-II-19 total legth 840 T1 6 Hf 70. 1. Thateg Laos FMNH 799 ski ad skull adult J. Delacour 14- XII-191 o. 109 total legth 680 T1 0 Hf71. QUANG NAM VIETNAM 1.. km W. 0. km S. Mt. Sotra Prov. Quag Nam Vietam USNM 671 ski ad skull adult T. J. Mc Ityre o. 607 1-111 prep. 1- III-1966 legth 610 T1 10 Hf 6 E alt. 00 m. USNM 67 ski ad skull adult 9 T. J. Mc Ityre o. 660 -V-1966 legth 64 Hf 6 T1 18 E 8. USNM 67 ski oly T. J. Mc Ityre o. 66 1-V-1966 legth 60 Hf 66 T1 0 E. THAILAND 14. Lomlo Mt. Dasai Loei USNM 00170 ski ad skull adult R. E. Elbel 9-111-194 R. E. 1 7007040. 1. Doi Sutep West Siam UZMC 141 holotype of Atherurus agustiramus Mohr 1964 skull oly adult evergree forest 1000 m det. 1-XI-199. CENTRAL LAOS 16. Nape Laos BMNH 8.7.1.144 ski ad damaged skull adult W. Lowe o. 84 9-1-198 HB 48 T1 0 Hf 79 E 8.

11 17. Xieg-Khouag Laos BMNH 6.10.4.19 ski ad damaged skull adult 9 J. Delacour-W. Lowe o. 17 1-1-196 HB 0 T1 17 Hf78 E 7. TONKIN VIETNAM 18. Lai Chau Toki FMNH 16 ski ad skull adult R. E. Wheeler tf 1 -IV-199 Kelley-Roosevelt Exp. o. 99 69619078. 19. Chapa Toki BMNH.4.1.490 ski ad skull adult 9 J. Delacour- W. Lowe o. 149 0-XI-199 alt. 000 ft. HB 1 T1 Hf 77 E40. BMNH.4.1.491 ski ad skull adult J. Delacour-W. Lowe o. 164 11 -XII-199 alt. 000 ft. HB 0 T1 Hf 7 E41. USNM 947 ski ad skull adult 9 J. Delacour-W. Lowe o. 17 alt. 000 ft. HB 1 T1 40 Hf 74 E 8. BMNH.4.1.49 ski ad skull subadult 9 J. Delacour-W. Lowe o. 118 alt. 000 ft. HB 4 T1 00 Hf 76 E 6. BMNH.4.1.49 ski ad damaged skull adult <$ J. Delacour-W. Lowe o. 174 HB 10 T1 4 Hf 7 E 4. BMNH.4.1.494 ski ad skull youg 9 J. Delacour-W. Lowe o. 1706-XII-199 alt. 000 ft. HB 70 T1 180 Hf69 E7. 0. Ngai-Tio Toki BMNH.1.1.9 holotype of Atherurus stevesi Thomas 19 ski ad skull adult 9 H. Steves o. 16 8-VI-194 alt. 4800 ft. HB T1 8 Hf 7 E 6. HAINAN CHINA 1. Haia Chia AMNH 6641 holotype of Atherurus haiaus Alle 1906 ski ad skull adult IX-190. Nodoa AMNH 6014 ski ad skull adult 9 C. Pope o. 8. Nodoa AMNH 6014 ski ad skull adult 9 C. Pope o. 66-1-19. Nodoa AMNH 60147 ski ad skull adult 9. C. Pope o. 676 7-1-19. Nodoa AMNH 6004 skeleto youg. Nodoa AMNH 117640 ski ad skull adult C. Pope. Nodoa AMNH 6004 ski ad skull adult C. Pope o. 98. 11-1-19. NORTHERN BURMA. 0 miglie Est di Bhamo Dit di Mogioch Birmaia MSNG 4177 skull ski o. 4176 youg L. Fea 188.. Haibum N.O. Burma AMNH 1 148 ski ad skull adult 9 H. C Rave -11-19 Veray-Hopwood Exped. o. 006 169. AMNH 11486 ski ad skull adult H. C. Rave -11-19 Veray-Hopwood Exped. o. 0160 14070. AMNH 11487 ski ad skull adult 9. H. C. Rave -11-19 Veray-Hopwood Exped. o. 0 6701870. AMNH 1148 ski ad skull subadult 9 H. C. Rave 9-1-19 Veray-Hopwood Exped. o. 14 6187 71.

1 4. Dalu N.O. Burma AMNH 11484 ski ad skull youg 9 H.C. Rave 1-11-19 Veray-Hopwood Exped. o. 44 6101807.. Hkampti Plai Upper Burma BMNH.11.168 skull adult 9 Lord Crabrook-F. Kigdo Ward o. 9. FMNH 41048 damaged skull adult Lord Crabrook. FMNH 41047 skull subadult 9 Lord Crabrook o. 9 16-XII-191 100ft. 6. Nigma Upper Burma BMNH 0.70 ski ad icomplete skull youg 9 R. Kaulback o. 0 1-1-199 N 6. E 97.4 100 ft. 7. Nam Tamai Valley Upper Burma BMNH 0.70 ski ad skull adult <J R. Kaulback o. 17 1-IX-198 470 (aus)-1877 N 7.4 E 97.4 000 ft. BMNH 0.704 ski R. Kaulback o. 87 -XII-198 N 7.4 E 97.8 000 ft. ASSAM INDIA 8. Cherrapuji Upper Assam BMNH 0.11.1.77 holotype of Atherurus assamesis Thomas 191 ski ad skull adult H. W. Wells o. 6 HB 40 T1 0 Hf 6 E 4 Wt 4. alt. 400 ft. Mawphlag Assam FMNH 7877 damaged ski ad skull fragmets youg 9> D. Wal-N. Koelz 4-XII-19 legth 474 Tl 1 Hf 6 E. WANHSIEN CHINA 9. Szechua Wahsie AMNH 869 ski ad skull adult Walter Grager o. 449 408040. UNKNOWN LOCALITIES IRSNB 1918 holotype of Atherurus retardatus Mohr 1964 ski ad skeleto adult 9> Soc. Roy. Zool. Avers 18.1.196. IRSNB 1177 paratype of A. retardatus Mohr*) skeleto adult <j\ Soc. Roy. Zool. Avers -XI-196. RMNH 1940 paratype of A. retardatus Mohr *) ski ad skull adult zoological garde Rotterdam 1-1-1967. RMNH 19897 paratype of A. retardatus Mohr*) ski ad skull adult 9. zoological garde Rotterdam 1-11-1968. RMNH 078 paratype of A. retardatus Mohr *) ski ad skull adult zoological garde Rotterdam 8-IV-1969. ZMA 09 B 9 as Mohr 1964: (ot 107 recorded) paratype of A. retardatus Mohr zoological garde "Natura Artis Amsterdam - Magistra" IX-196. MNHN 196.1 skull adult Ido-Chie Capitaie Boifacy a vecu du 7 mars 190 au javier 190. BMNH 81.11..1 Malaya regio ski ad skull adult E. Gerrard 880c presumed Pulo Aor. *) At the momet the holotype was studied by Mohr these specimes were still alive i the Atwerp ad Rotterdam Zoo.

Malay Hystrix Differig "Habitat 1 RESULTS Geus Atherurus Atherurus F. Cuvier 189:48 Type species. Hystrix macroura Liaeus 178 type by mootypy. Rage. Peisula ad adjacet small islads Thailad Laos Vietam Haia ad Szechwa (Chia) Burma Assam (Idia) ad tropical Africa (extra-limital). Diagosis. from the more primitive geus Trichys *) Guther i the umber of dorsal vertebrae 14 i Atherurus agaist 16 i Trichys. The tail is shorter o average oe quarter to half the legth of the head ad body with about 19 to caudal vertebrae as agaist about to i Trichys. As for the skull characters there is scarcely a postorbital process ad the postorbital breadth is always larger tha i Trichys. The logest spies are always larger tha those of Trichys. From the least specialized represetatives of the geus Hystrix subgeus Thecurus it differs Lyo 1907 i the permaet molars showig visible roots whe i place ad i the quite differettail structure. This geus is fully characterized by its peculiar caudal bristles alterately expaded ad cotracted from oe up to six times. Accordig to Yog Hoi-Se et al. (197) Atherurus macrourus has a diploid umber of4 chromosomes agaist 46 chromosomes i Trichys fasciculata. Remarks. macroura Liaeus 178 is the type by mootypy as poited out by Jetik (1894: 07) ad Lyo (1906: 199) ad ot by subsequet desigatio by Lyo (1907: 84) as metioed by Delay (197: 11). Shaw's (1801) Hystrix fasciculata from Malacca has sometimes bee associated with Atherurus (Guther 1889; Thomas 1889 ad 1907; Bohote 1900). This has bee refuted thoroughly by Jetik (1894) Mohr (196) more recetly by Va Weers (1976) ad the desigatio of a Malaya Trichys specime as eotype for H. fasciculata by the latter author defiitely has take away the basis for this cofusio. The five omial taxa from the Africa part of the rage should probably be regarded as oe species A. africaus Gray 184 accordig to Delay (197) ad Kigdo (1974) but are ot listed here. List ofomial taxa: Hystrix macroura Liaeus 178: 7 Type. No holotype desigated. Type locality. i Asia". Lyo (1907: 84) restricted the type locality to Malacca obviously used as a syoym of Malay Peisula ad take as such i the preset paper. *) The type species of this geus is ad Trichys lipura Guther 1877 type by mootypy ot Trichysfasciculata (Shaw 1801)" as erroeously recorded by Va Weers (1976: 19).

The 14 Atherura zygomatica Miller 190:4 Holotype: USNM 1149 ski ad skull adult 9. Type locality: Pulo Aor. (Loc. o. 7 i map fig. 1) Atherurus haiaus Alle 1906: 470 Holotype: AMNH 6641 ski ad skull adult. Type locality: Haia. (Loc. o. 1 i fig. 1) Atherurus terutaus Lyo 1907: 87 Holotype: USNM 1971 ski ad skull adult <$. Type locality : Pulo Terutau. (Loc. o. i fig. 1) Atherurus tiois Thomas 1908: 10 Holotype: BMNH 8.1..1 $ ski. Type locality: Juara Bay PulauTioma. (Loc. o. 9 i fig. 1) Atherurus macrourus pemagilis Robiso 191: 90 Holotype: BMNH 49.4 ski ad skull adult Type locality: Pulau Pemaggil. (Loc. o. 8 i fig. 1) Atherurus assamesis Thomas 191: 98 Holotype: BMNH 0.11.1.77 ski ad skull adult Type locality: Cherrapuji Assam. (Loc. o. 8 i fig. 1) Atherurus stevesi Thomas 19: 0 Holotype: BMNH.1.1.9 ski ad skull adult 9. Type locality: Ngai-Tio Toki. (Loc. o. 0 i fig. 1) Atherurus retardatus Mohr 1964: 10 Holotype: IRSNB 1918 ski ad skull adult 9. Type locality: Ukow from Zoo Rotterdam. Atherurus agustiramus Mohr 1964: 108 Holotype: UZMC 141 skull adult ski of head ad tail with brush. Type locality: Doi Sutep Chegmai Thailad. (Loc. o. 1 i fig. 1) Craial measuremets: Measuremets of 1 craial dimesios for 16 samples from the geographic areas give i table I are preseted i table II. Some of them ad some craial characters ot icluded i this table are further discussed below. The remaiig craial dimesios preseted i table II reflect maily the differeces as already expressed by the craial legth measuremets. Craial legth. occipito-asal legth ad the bassilar legth are also preseted i two histograms figures ad. Both histograms show the cosiderable differeces i skull size. All the islad populatios are distictly smaller (occ. as. 1. 8794 mm) tha the populatios of Teasserim Laos Thailad Toki ad Wahsie (97 109 mm). However the samples of the cotietal populatios from Aam N. Burma ad Assam are equally

1 Figure. Frequecy distributio of the occipito-asal legth of skulls of Atherurus macrourus. Adult specimes oly. Locality umbers correspod with those o the distributio map (fig. 1). Specime from ukow locality is MNHN 196.1. Figure. Frequecy distributio of the basilar legth of skulls of Atherurus macrourus. Adult specimes oly. Locality umbers correspod with those o the distributio map (fig. 1).

I 16 Figure 4. distributio Frequecy of the relative size of the asals of Atherurus macrourus expressed as a percetage of the occipito-asal legth of the skull. Adult specimes oly. Locality umbers correspod with those o the distributio map (fig. 1). small ad that from the Malay Peisula is itermediate i size (9 101 mm). The basilar legth measuremets reflect these proportios i the same way (respectively 6979 mm 8189 mm ad 778 mm). Therefore it would be quite arbitrary to divide these populatios ito two or more taxoomic uits o craial legth aloe. Nasal legth. most of the origial descriptios of Asiatic Atherurus species the asal legth has bee used as a diagostic character. Therefore the relative legth of the asals expressed as a percetage of the occipitoasal legth is preseted i a histogram (fig. 4). This relative measure has bee chose because the absolute size differeces have bee stressed sufficietly by the two craial legth measuremets. The holotype of A. terutaus Lyo 1907 has absolutely as well as relatively the smallest asals (1.7 mm 44%). But sice it is the oly specime available from that locality o coclusios as to its specific or subspecific status o this character are justified. The same ca be said of the specime from Wahsie Chia represetig the upper extremity of this measuremet i table II ad fig. 4 (.0 mm 1.7%). There is a clear differece betwee the relative asal sizes of the sample from S. Laos (0. 1.0%) ad those of the

I Accordig Miller 17 Malaya specimes (6.9 8.1%) but the measuremets of all the other cotietal populatios are itermediate betwee these two. The relative size of the asals of the five remaiig islad populatios varies from 6 to 0% so it is obvious that this character is of little use for diagostic purposes. Nasal breadth. Thomas' (19) descriptio the asals of A. stevesi are called arrower eespecially their posterior breadth. Whe compared with the Malay Peisular populatio this character does ot hold good. The posterior breadth of the combied asals of the holotype from Toki is 4.% of the asal legth while i six Malaya specimes this ratio varies from 40.7 to.0% ad i five specimes from Toki from 4. to 1.%. The most importat diagostic character i Mohr's (1964) descriptio of A. agustiramus is the differece betwee the posterior ad aterior breadth of the asals i the holotype respectively 1.4 ad 16.4 mm. However these dimesios fall withi the rage of variatio of the Asiatic speciees as may appear from table II ad extremer ratios are foud i a specime from Malaya (11. ad 1.4 mm) ad i oe from Laos (11.7 ad 16. mm). Lachrymal boe. to Miller's descriptio (190) there is a costat differece betwee the form of the lachrymal boe of A. zygomaticus ad that of the mailad aimal. Miller gives fully 8 mm for the lateral dimesio of the latter's lachrymal boe but of the six Malaya specimes available three are below that value (. 10 mm m 7. mm). The same author metios measuremets of about mm for zygomaticus from Pulau Aor but the seve available measuremets rage from to 9 mm (m 6.1) so this differece is ot costat ad is smaller tha origially described. Accordig to Miller (190) agai the forward extesio of the lachrymal boe i the dorsal skull surface i zygomaticus is scarcely visible whe the skull is viewed from above. Measuremets of the zygomaticus specimes show legths from 0 to 4.4 mm ad the Malaya sample to 7. mm. So there is some overlap ad i 6 measuremets of the remaiig Asiatic samples the dorsal legth of this boe appears to vary from 0 to early 8 mm obviously correlated with craial legth. Therefore the populatio of Pulau Aor caot be distiguished from the cotietal populatios o this character. Zygoma shape. (190) stated that the zygoma of his A. zygomatica is shorter tha that of the mailad aimal. Expressed as a percetage of the basilar legth the zygoma legth (measured from the rim of the ifraorbital forame to posterior the posterior edge of the zygomatic of the process squamosum) of seve specimes from Pulau Aor rages from 8. to 40.7% (m 9.%) ad six Malaya specimes vary from 7.4 to 4.8% (m 9.6%) so these samples do ot differ at all i this respect. Miller (190) also stated that compared with the cotietal forms the zygoma of the Pulau Aor form is deeper i proportio to its legth. To test this statemet the height of the jugal boe at the aterior jugal suture of part

18 Table II. Craial measuremets i millimetres of adult specimes of Atherurus macrourus from geographic areas as distiguished i the preset study with rage mea of sample i paretheses umber of measuremets () ad the stadard deviatio (s). For the defiitios of the measuremets see Material ad Methods. P. Pulau Tu.Terutau PgPeag Pl. Pemaggil Te. Teasserim Vm. Vietam Ass. Assam Wah. Wahsie U.l. ukow locality (MNHN 196.1). Malay Peisula P.Tu. P.Pg. Pulau Aur P. PI. Pulau Tioma Te. Souther Laos Occ. as. 1 94.7101.4 (97.9) 6 s.8 88.8 9. 90.194.0 (9.) 8 s 1.8 89.6 86.79.8 (90.7) s.4 1 100. 100.0109.0(104.1) Basilar 1. 77.08. (80.6) 6 s.77 7.8 77.4 7.878.1 (76.7) 8 s 1.7 68.6 7.677.7(7.8) s. 10 8.9 8.89.0 (8.) L. asals.48.1 (7.0) 6 s1.06 1.7. 4.7.1 (.9) 8 s0.89 4.7 4.7. (.6) s 1.40 9.4 1.0.8(.0) L. frotals 4.641.6 (9.1) 6 s.48 6..0.7.0 (.) 8 s 1.1.9.66. (4.0) s 1.74 4.4 9.0040. (9.7) Palatal 1. 44..1 (48.0) 6 s.71 4.1 46.7 41.6 48.8 (46.) 7 s.1 4. 44.46.1 (4.4) s0.74. 1..0 (.7) L. diast. 6.41.0 (8.8) 6 s 1.48 8.0 7.6 6.07.7(7.1) 7 s0.6.9.88.1 (7.4) s0.94 9.8 0.04.(1.6) B. as. po. 11.14.(1.8)6 s 1.17 10. 1.8 1.01.(1.6) 7 s0. 11. 11.61.6(1.4) s0.88 14. 1.714.(14.0) B. as. at. 14.716.7(1.)6 s0.76 14.6 1.6 1.814.1 (1.6) 7 s0.67 1. 1.1.(14.6) s0.8 1.8 14.817.0(1.7) Postorb. b..99.1 (7.6) 6 sl.l7 4.6 6.4 4.46.7(.) 7 s0.86.1.87.4 (6.4) s0.6 8.0 6.8.8 (7.8) Zygom. b. 46.00. (48.0) 6 s 1.70 4. 44.0 44.-47.0 (4.6) 7 s0.8 4. 47.148.8(48.0) s0.68 0.0 47.48.8 (48.) Height sk..07.0 (6.0) 6 s0.94.9 4.0 4.6.6 (.) 8 s0.6.4.6.4 (4.) s 1.10 6.8.18.1 (6.8) L. madible 60.864.0 (6.) 6 s 1. 6.7 7.0.69.4 (7.9) 8 s 1.1..09.(7.7) s 1.68 60.0 6.66.6 (64.6) H. madible.0. (.) 4.6 4.16.0 (.) s0.7 4.6.(.) Alv. P4 M 17.019.(18.0)6 s0.8 16. 18. 1.017.4(16.) 6 s0.89 16.8 1.716.8(16.) s0.4 18. 17.118.(17.7) p Alv. p4 m 18.90.6(19.8)6 s0.74 17.8 19.8 17.18.1 (17.6) 7 s0. 17.618.(17.8) s0.9 19. 19.019.6(19.)

19 Table 11. Cotiued. Aam VM. Thailad C. Laos Toki Vietam Haia Chia N-Burma Ass. Wah. U.l. Occ. as. 1. 87.89. 97.10.7 97.4107.1 (10.1) 87.91.0(88.9) 90.094.9 (9.) 9.1 104.1 111. s4.17 6 s 1.48 6 s 1.8 Basilar 1. 7.07.7 8.86.9 8.4 81.486.6 (8.7) 69.77.(71.4) 74.078.0 (7.4) 76.8 89. 88.0 s.6 6 s 1. 1 6 s 1.6 L. asals 6.46.9.00.4 9.0-^9. 8.7.6 (0.4).7. (.1).77.8 (6.7).8.0.0 s 1.9 6 s 1.69 6 s0.77 L. frotals 1.8.9 9.41.9 4.4 6.4. (9.0).89.(.0) 4.89.(6.9).0 8. 40.0 s.0 6 s. s 1.9 Palatal 1. 44.0 4.0 49.7.6 1.1.9 0.4.9(.) 4.-4.6 (44.) 41.846.0 (4.0) 46.7 6..0 s1.9 s 1.6 6 s 1.6 1 L. diast..47.0 1.1.9 0.41.7 0.01.9 (1.1) 4.6.7(.).69.7 (7.4) 8.0 0.7.0 s0.80 s 1.08 6 s 1. B. as. po. 11.41. 1.41.7 11.71. 1.714.7(14.1) 9.1.(10.9) 11.614.(1.) 1.7 1. 1.4 s0.79 6 s0.98 6 s1.00 B. as. at. 1.1.8 16.016.4 16.17.0 14.416.7(1.) 11.81.(1.0) 1.81.(14.) 14.0 1.8 s0.8 6 s 1.9 6 s0.6 Postorb. b. 4.4. 6.87. 7.98.4 7.9.(8.4) 4.77.1 (.8).9.6 (6.8).4.9.7 s0.6 s0.88 s 1.64 Zygom. b. 4.74. 1.. 0.00. 49.01.8(49.6) 4.64.0 (4.8) 44.47.9 (46.4) 4. 48.8 48. s 1. 6 s0.84 6 s1.60 Height sk..74.1.77.4 7.4.86.9 (.7).04.0 (.7).46.9 (.).6 7. 7.0 s 1. 6 s0.8 6 s 1.46 L. madible 4.0.1 6.864.7 6.6.9 61.867.0(6.).6.8(4.7).09.9 (7.).9 66.1 6.0 4 6 s 1.7 6 s.09 H. madible 0.1. 4.46. 4.8.4 0.0.7(1.) 1..9 (.) 4.0 6 s0.98 4 Alv. P4 M 14.01. 16.619. 17.417. 16.118.7(17.7) 1.016.4(1.7) 16.18.1 (17.) 16.4 0.8 17. s 1.07 s0. 6 s0.97 Alv. p4 m 14.16.1 19.619.7 19.019.0 17.60.1 (19.) 16.17.0(16.6) 16.418.(17.6) 17.. 19.0 4 s0.7 s0.7

The The 0 of the material studied has bee measured ad expressed as a percetage of the zygoma legth. I six Malaya specimes to 8.9% (m 7.6%) ad i seve specimes this character varies from.7 from Pulau Aor from 9.9 to 1.7% (m 0.7%) so i this respect these samples differ sigificatly ideed. Nevertheless i this character too there is much variatio throughout the total Asiatic populatio. Five specimes from Pulau Tioma have relative heights of the jugal of 4.88.% three specimes from South ad Cetral Laos of 6..0% ad four specimes from N. Toki of 4.60.1%. Moreover oe specime (BMNH 81.11..1) from ukow locality with a most clear "toothlike process" at the zygoma (see below) has a relative zygoma height of 8.4% so either the latter character i zygomaticus shows more variatio or the former character is more widespread. So o the basis of the zygoma height with the idividual ad geographical variatio as represeted o satisfyig divisio of the species is possible ow. The lower cotour of the jugal was described by Miller (190) as follows:.. broke by a strogly developed cocavity beeath posterior jugal suture this cocavity termiatig ateriorly o the posterior upper surface of a well-marked tooth-like projectio." The sample from Pulau Aor is distiguished more clearly by this character ideed tha by those discussed before but it is ot developed equally strogly i the eight zygomaticus specimes studied either is it quite uique for this populatio. Although ot all samples were studied o this character a specime with a more or less clearly developed cocavity with toothlike process was foud i the Toki ad i the Aam sample (FMNH 16 ad USNM 671) ad i a larger umber of specimes this character was foud more or less slightly idicated. Exteral characters Body size. legth of head ad body of the Asiatic Atherurus species as appears from collector measuremets recorded i the list of specimes studied rages from 8 to 7 cm with a mea of 48 cm. Further aalysis of these measuremets has ot bee udertake because of the ucertaity about the way i which they have bee obtaied. Tail legth. legth of the tail i 1 collector measuremets varies from 11 to 6 cm beig to % of the legth of head ad body mea 40%. I dry skis the aterior border of the tail is ot easily defiablebecause of the differet ways the skis have bee prepared. Therefore the legth of the scaly part of the tail of dry skis has bee measured (table III). I Lyo's (1907: 87) descriptio of A. terutaus the shorter tail was metioed as diagostic character. The legth of the scaly part of the tail of the holotype (40 mm) is extremely small ideed whe compared with the Malaya specimes (10 10 mm). But the measuremets i table III show a idividual variatio i the sample from Pulau Aor of 70 to 140 mm i that

The The The 1 from Haia of 60 to 10 mm ad i the N. Burmese sample of 60 to 110 mm. Agaist the backgroud of the idividual ad geographical variatio of the tail legth as preseted i table III oe sigle specime is isufficiet proof for a differet species or race o Pulau Terutau. Hidfoot. legth of a hidfoot i 9 collector measuremets varies from6 to 80 mm with a mea of 71 mm. Ear. legth of the ears i 9 collector measuremets varies from to 4 mm with a mea of 7 mm. Colour. idividual variatio i the blackish-brow colour of back ad sides is maily caused by a differet frequecy of visible white parts of the spies ad tactile bristles. This exposure of white parts of spies however may be iflueced by shrikage or stretchig of a ski durig preparatio. Both idividual variatio ad this pheomeo iterfere with the judgemet of geographical variatio. Mohr's (1964) A. retardatus is maily based o the presece of light ad dark logitudial stripes alog the back ad sides of adult aimals. Accordig to this author all other Asiatic Brush-tailed Porcupies oly show logitudial stripes at birth which disappear after three or four weeks. However little is kow about the developmet of the colour patter durig life ad there is othig that affirms Mohr's statemet. These dark ad light stripes are foud from rather clearly to at least slightly perceptible i all the Asiatic Atherurus specimes studied ad the type respect. series does ot differ i this Thomas (191) metiosof his assamesis: "colour above rather darker" but cosiderig the idividual variatio as observed i all the other Asiatic samples this sigle specime does ot prove that the populatio i Assam is differet i this respect. I a very light-coloured ski from Toki (BMNH.4.1.490) early 0% of the large spies ofthe back have white tips of up to mm i legth. This is probably ot commo i this geographic area. The holotype of A. stevesi Thomas 19 from Toki does ot possess whitetipped spies but ufortuately this ski is damaged ad early large spies are lackig. Other skis from that area however half of its show these white spies i a much less cospicuous degree. Miller (190) ad Thomas (191 ad 19) used a lighter or darker colour of the uderside as a diagostic character for respectively zygomaticus of Pulau Aor assamesis from Assam ad stevesi from Toki. Chi throat ad chest betwee the forelegs are whitish i all the specimes of the whole Asiatic material studied but there is ideed some variatio i the colour of the belly. I some specimes from the Malay Peisula the belly is light brow with the spies with a pale brow middle part ad a whitish tip whereas others have more white o the belly. Some specimes from Pulau Aor have the same light brow belly as the darkest specimes from Malaya whereas others from that islad have the belly more darkly mottled. Thomas' (19) type of stevesi has a white belly but other specimes from Toki have light brow bellies dirty white or light brow with a arrow white strip

The The i the middle. Apparetly there is so much variatio i the colour of the uderside that its use as a diagostic character i the small samples available is ope to questio. Spies ad tactile bristles. measuremet of the logest spie ad tactile bristle is somewhat subjective because it is ot always possible to kow whether i fact the logest spie bristle of or a ski has bee measured. Therefore although these measuremets provide a good method of comparig the developmet of the spiy coverig o mea or stadard deviatio is recorded ad oly the rages of the samples are preseted i table III. The relatio betwee skull size ad developmet of spiy coverig shows the followig tedecy. The samples with the largest skulls from Teasserim Laos Thailad ad Toki (see figs. ad ) have the highest developed spiy coverig with spies of 80 to 117 mm ad tactile bristles of 10 to 160 mm agaist spies of 70 to 7 mm ad tactile bristles of 80 to 110 mm (see table III) i the samples from Aam ad Haia with smaller skulls. This tedecy is far from costat. From the figs. ad ad table III it appears that the specime from Wahsie Chia with early the largest skull has relatively short spies ad bristles (respectively 7 mm ad 110 mm). Further the specimes from Pulau Aor with smaller skulls o a average tha those from the Malay Peisula have spie legths of 90 to 108 mm agaist 7 to 100 mm i the Malaya aimals. Moreover the sample from N. Burma with a relatively small skull legth (figs. ad ) has rather large tactile bristles (10 to 1 mm). The largest spie i a ski varies i thickess from 1. to. mm i the total material studied ad its breadth rages from 1.8 to.0 mm. These log spies o the posterior upper part of the back differ i form from the majority of the smaller spies i a ski. The log spies are thicker ad the shorter oes are broader ad thier as be demostrated with the may measuremetsof a specime chose arbitrarily (BMNH..6.7) i mm: legth thickess breadth 100.1. 97 1.8. 0 1.6. 4 1..6 0.9.6 The largest tactile bristle i the skis of all the specimes studied has a thickess ear the basis of 1.0 to 1.8 mm. Tail brush. legth of the largest tail bristle i the total material studied varies from 0 to 10 mm the greatest umber of expasios i oe bristle varies from two to six the size of the expasios is i most cases about 10 mm ad rages from 8 mm i some smaller beads to mm i termial oes which are geerally somewhat larger.

Table III. Legth of the logest spie legth ofthe logest tactile bristle ad legth of the scaly part of the tail i the skis of Atherurus macrourus studied. Adult specimes oly. The locality umbers correspod to those i the map of fig. 1 measuremets i millimetres umber ofskis available. Loc. umber L. spies Tact. br. Scaly p. tail Malay Peisula 14 7--100 11010 1010 P. Terutau 90 10 40 1 P. Peag 6 80--8 110140 110 P. Aur 7 90--108 11010 70140 7 P. Pemaggil 8 87--89 10 P. Tioma 9 76--87 11010 10010 4 Teasserim 10 87 10 1 1 Laos 11 1 16 17 97--110 1140 90140 Aam 1 70--7 10010 1010 Thailad 14 1 10 160 160 1 Toki 180 80--117 10160 10170 7 Haia 1 70--7 80110 6010 4 N. Burma 7 76--8 101 60110 4 Assam 8 8 8 140 1 Wahsie 9 7 110 10 1 Lyo (1907: 88) metioed the caudal bristles with oly oe expasio i the tail brush of the holotype ad oly specime of his ew species terutaus ad supposed that the wor appearace of the bristles might accout for this feature. This implies that each bristle should have lost oe or more beads but the form ad legth of the expasios suggest that these are termial oes. This adds to the probability of Mohr's (1964: 104) suppositio that this specime had totally shed its bristles ad was formig ew oes. Nevertheless idividual or geographical variatio as a possible explaatio may ot be left out of cosideratio. A re-examiatio of this specime yielded oe bristle with two expasios legth 8 ad 1 mm. A tail brush with bristles havig oly two expasios is foud i oe specime i the sample from Pulau Aor remaiig specimes havig four or five expasios ad also oe i the Haia sample remaiig specimes havig three expasios. The available data suggest that there is a correlatio betwee tail legth ad umber of expasios i the bristles. As discussed before this specime from Pulau Terutau has the shortest tail of all the specimes studied so it is ot ecessarily a aberrat idividual. But cosiderig the idividual variatio of this character i the Asiatic material this sigle specime caot be accepted as sufficiet proof for a differet species or subspecies o that islad. Wool-hairs. The origial descriptio of A. stevesi Thomas 19 reads: "Upper surface with much greater developmet of the close coat of woolhairs beeath the spies." Each porcupie ski possesses somethig of a udercoat of wool-hairs i most cases oly perceptible whe the spies are separated ad sometimes more clearly visible. The type ski however has

No Subjective Ellerma The Origial 4 lost the greater part of its spies o the middle ad posterior part of the back thus exhibitig these wool-hairs i quite a uusual ad way makig it difficult to the compare frequecy of these hairs with other specimes. Two specimes from orther Toki do ot show this character as described by Thomas whereas i four others from that geographical the area wool-hairs are more abudat. The specime from souther Chia does ot show this character very cospicuously ad southwards the specimes from Laos do ot differ i this respect from the Malaya skis. But with the material available it is ot possible to decide whether the Toki sample ot o a average from others with regard to the developmet differs or of woolhairs. CONCLUSIONS All the Asiatic Brush-tailed Porcupies belog to oe species Atherurus macrourus (Liaeus 178). With the material curretly available a satisfactory divisio ito subspecies is ot possible ad all the subspecies distiguished up to this momet are poorly characterized. The results of this study support the probability of Ellerma's suppositio (1961) that it would be impossible to divide this species ito subspecies if a larger umber of specimes was collected. Atherurus macrourus (Liaeus 178) Hystrix macroura Lieaus 178: 7 Type. type desigated. Type locality. type locality "Habitat i Asia" restricted to Malacca by Lyo (1907: 84) cosidered a syoym of Malay Peisula i the preset paper. Distributio. Asiatic rage of the geus i the list of omial taxa. Syoyms. syoyms as eumerated i the list of omial taxa. Characters. (1961: 99) gives a detailed descriptio of exteral ad craial characters. A umber of characters with their idividual ad geographical variatio is discussed i the preset paper (see Results). This species differs from the Africa form i the total absece of quills (log ad thick spies circular i cross-sectio ad with the largest diameter about mid-legth). As far as I kow some of these quills are always preset i the skis of Africa brush-tailed porcupies ad are missig oly whe they have bee shed. I A. macrourus there is o iflatio of the frotals ad the heights of the skulls expressed as a percetage of the basilar legth rage from 9. to.6% (1 m.%). East Africa populatios are macrourus-like i this respect whereas West Africa specimes show some iflatio of the frotal regio. The holotype of A. africaus Gray 184 exceeds the rage of the Asiatic series with a relative height of the skull of

6.1% ad i a specime from Camerou (BMNH 48.197) this is eve more clearly with 8.8%. Status of the amedforms A. zygomaticus of Pulau Aor was compared by Miller (190) with Malaya specimes ad was characterized as follows: lachrymal boe smaller zygoma shorter jugal deeper jugal with posterior cocavity ad a toothlike process whitish areas less developed. As discussed before (see Results) the differece i zygoma legth is ot bore out by the preset study the other differeces appeared to be smaller tha origially described ad oe is absolute whe compared with the total Asiatic material. Cosiderig the idividual variatio i this sample the small sample size ad the eormous variatio i all the craial ad exteral characters i this species a omeclatorial recogitio of this populatio i my opiio is ot justified ow. A. haiaus was diagosed by Alle (1906) with the followig differeces from the mailad aimal: smaller size darker colour shorter spies ad tail. The occipito-asal legth of the Haia sample is very small ideed but it does ot differ i this respect from the samples of the five small Malay islads Aam ad orther Burma (figs. ad ). The spies are very small too but ot differetfrom the spie legths of the Aam specimes (table III) the tail legth is ot smaller tha i the remaiig samples with smaller craial size (table III) ad the colour is probably ot outside the rage of variatio i the species. Therefore it is apparet that A. haiaus is isufficietly characterized for subspecific recogitio. A. terutaus was characterized (Lyo 1907) amog others by shorter asals smaller lachrymal boe more proouced orbital costrictio ad a shorter tail with shorter bristles. The developmet of lachrymal boe ad orbital costrictio falls withi the rage of variatioof the species but the legth of the asals ad the legth of the tail are the shortest measured i this species (fig. 4 ad table III). However with the large amout of variatio i this species i mid this sigle specime is cosidered isufficiet for the proof existece of a differet subspecies o Pulau Terutau. A. tiois was said (Thomas 1908) to have a shorter tail a lower skull ad rather large asals. However whe compared with cotietal samples the tail is ot short (table III) the asals are ot large (fig. 4) ad the relative heights of the skulls i five specimes available (0..1% of the basilar legth) fall withi the rage of the total Asiatic material (9..6% 1). A isufficiet umber of differeces could be foud i this sample to cosider the populatios of Pulau Tioma a differet subspecies. A. macrourus pemagilis was compared i Robiso's (191) diagosis with a umber of rather variable characters of zygomaticus tiois ad Malaya specimes. The oly remark that ca be made about the holotype ad at the same time oly adult specime that has bee available to me is that its

6 craial size approaches the lower limit of the rage of variatio of the species (figs. ad ). This basis is isufficiet for maitaiig subspecific rak for the populatio of Pulau Pemaggil. The mai characters ofa. assamesis from Cherrapuji Assam as give i the origial publicatio (Thomas 191) are highly variable i the Asiatic Atherurus species such as the breadth of the skull form of the asals form of the zygoma colour etc.... Compared with the geographically earest populatio that of orther Burma it has the same craial size (figs. ad ) ad scarcely loger asals (fig. 4). The relative zygomatic breadth of the skull expressed as a percetage of the basilar legth is smaller (6.4% agaist 9.6.9% i the Burmese sample) but is ot outside the rage of the species (4.8 6.9%). So this the oly available adult specime does ot show eough differeces to justify subspecific recogitio. Craial size legth ad breadth of the asals colour of the back ad belly ad developmet of the wool-hairs of A. stevesi were discussed before. Other merely trivial skull characters have bee metioed i the origial descriptio (Thomas 19) such as the upper profile of the skull form of palatal foramia ad posterior palate etc. O the characters studied the Toki sample caot be distiguished from the others except o larger size but this character aloe as stated before produces a rather artifical classificatio of the Asiatic Brush-tailed Porcupies. A. retardatus Mohr 1964 is based o the presece of light ad dark logitudial stripes alog the back ad sides of six livig adult specimes i the zoological garde "Blijdorp" i Rotterdam i 199. The origial descriptio is based o oe of these specimes which came ito the collectios of the Istitut Royal des Scieces Naturelles i Brussels after it had died ad o a specime i the collectios of the Zoologisch Amsterdam (ZMA 09 B) obtaied from the zoological garde Museum "Artis" i Amsterdam (ot ZMA 9 as Mohr erroeously recorded). The specimes from Rotterdam came "via Sigapore" ad the geographical origi of the latter specime is quite ukow. Three of the remaiig specimes i Rotterdam came ito the collectios of the Rijksmuseum va Natuurlijke Historie i Leide (RMNH 1940 19897 ad 078). Accordig to Mohr (1964) the holotype (IRSNB 1918) had lost its stripes before the preparatio ad cocerig the paratype ZMA 09 B othig is kow about the presece of stripes durig life. Little or othig is kow about alteratios i the colour patter of the Asiatic Brush-tailed Porcupies ad the type series of retardatus does ot differ at all from other samples (see also Results: colour) so there is o reaso at all for maitaiig this specific or subspecific ame. The form of the asals of the holotype of A. agustiramus Mohr 1964 from Thailad has bee discussed before (see Results). The breadth of the orbital process of the maxilla the most importat diagostic character is really very small.9 mm agaist.6.8 mm i twety ie other Asiatic specimes measured. The geographical earest specime (USNM 00170) i Thailad

7 has a more ormal orbital process of mm so it is very probable that the arrow orbital process of the holotype is a accidetal idividual variatio. Apparetly this ame must be cosidered a syoym too. ACKNOWLEDGEMENTS I wat to thak most sicerely Dr. P. J. H. va Bree Curator of the Mammal Departmet Zoologisch Museum Amsterdam for his advice ad criticism throughout this study. For allowig me to study specimes uder their care for ledig me material ad for cooperatio I am most grateful to the followig persos: Dr. S. Aderso (America Museum of Natural History New York) Dr. I. R. Bishop ad Daphe M. Hills (British Museum (Natural History) Lodo) Dr. F. W. Braestrup ad Dr. P. Valeti- Jese (Uiversitetets Zoologiske Museum Copehage) Dr. A. M. Husso ad Mr. J. Schoute (Rijksmuseum va Natuurlijke Historie Leide) Dr. X. Misoe (Istitut Royal des Scieces Naturelles Brussels) Dr. Fr. Petter ad Mr. J. Roche (Museum Natioal d'histoire Naturelle Paris) Dr. H. W. Setzer (Natioal Museum of Natural History Washigto) Prof. Dr. E. Tortoese (Museo Civico di Storia Naturale Geoa) Dr. L. de la Torre (Field Museum of Natural History Chicago). I am also idebted to Mr. C. M. Sep (Alkmaar) for his revisio of the Eglish ad to Dr. J. E. Hill (British Museum (Natural History) Lodo) for a fial correctio. A subvetio of the Netherlads Orgaizatio for the Advacemet of Pure Research (Z.W.O.) eabled me to visit the Museums of New York ad Washigto i 1974 to study Southeast Asiatic Hystricidae. GAZETTEER The followig geographic ames are those from the list of specimes studied. Differet spelligs of syoyms are eclosed i paretheses. Mostof the coordiates were take from the "Official Stadaid Names Gazetteer" of the Uited States Board o geographic Names Washigto some of them were derived from "The Times Atlas of the World" ed. 197 Lodo or from published iformatioof the collectors. Malay Pe isula (with adjacet islads): Guug Tahag Jalor Pulau Aor (Pulo Aur) Pulau Pemaggil Pulau Piag (Pulo Peag) 4. N 10.4 E 6.0N 101.10 E.7 N 104.1 E. N 104.0 E.0 N 100.8 E Pulau Terutao (Pulo Terutau; Ko Tarutao) 6. N 99.40 E Pulau Tioma.48 N 104.11 E Trag (Trog) 7. N 99.6 E Wellesley. N 100. E

Proc. Bull. 8 Thailad (mailad): Doi Suthep (Doi Sutep) Lomlo Mt. (Lom lo Phu) 18.48 N 98. E 17.01 N 101.0 E Laos: Nape Plateau Boloves (Phouphieg Boloves) Thateg Xieg-Khouag 18.18 N 10.07 E 1.0N 106.0 E 1.6 N 106. E 19.1 N 10. E Vietam: Chapa (Cha Pa) LaiChau Ngai-Tio (Osgood 19: pi. 9) Qag Nam Sotra (So Tra).1 N 10.0 E.04 N 10.10 E.44 N 10.6 E 1.40 N 108.00 E 1. N 108.46 E Burma: Dalu (Dalu Ga; Taro) Hai Bum (Carter 194: map) Hkampti Plai Nam Tamai Valley Thaugyah (Thougyah) 6.1 N 96.11 E 6.04 N 9. E.7.N 97.00 E 7.4 N 97.4 E 11. N 98.44 E Assam Idia: Cherrapuji Mawphlag.16 N 91.48 E.8 N 91.46 E Chia: Nodoa (Na-ta) Wahsie (Wa-Hsie) 19.1 N 109. E 0.49 N 108.4 E REFERENCES ALLEN J. A. 1906 Mammals from the islad of Haia Chia. Amer. Mus. at. Hist. : 46490pi. 69. BONHOTE J. L. 1900 O the mammals collected durig the "Skeat Expeditio" to the Malay Peisula 18991900. zool. Soc. Lodo 1900 : 86988 pi. LVI. CARTER T. D. 194 The mammals of the Veray-Hopwood Chidwi expeditio orther Burma. Bull. Amer. Mus. at. Hist. 8 : 9114 pis. 6 1 map. CUVIER F. 189 Dictioaire des scieces aturelles 9 : 10 (LevraultStrasbourg ad Paris).

Z. Notes Smithso. 191 O ew mammals from the islads of the Johore Archipelago South Chisse Sea. Proc. Proc. Proc. Field 9 DELANY M. J. 197 The rodets of Ugada: IVIII 116 89 figs. (British Museum (Natural History) Lodo.) ELLERMAN J. R. 1961 The faua of Idia icludir Pakista Burma ad Ceylo. Mammalia ed. (1): IXXX 1 48 (Zoological Survey of Idia Calcutta). GRAY J. E. 184 Descriptio of some ew geera ad fifty urecorded species of mammalia. A. Mag. at. Hist. (1) 10 : 67. GCINTHER A 1877 Report o some of the additios to the collectios of mammalia i the British Museum. Proc. zool. Soc. Lodo 1876 : 771 pis. LXIXLXXIV. 1889 Note o a Borea porcupie Trichys lipura. zool. Soc. Lodo 1889 : 777. JENTINK F. A. 1888 Museum d'histoire aturelle des Pays-Bas. Cataloguesystematique des Mammiferes (Rogeurs Isectivores Cheiropteres Edetes et Marsupiaux) 1: 180 (Brill Leide). 1894 O Trichys fasciculata (Shaw). Leyde Mus. 16 : 009 KINGDON J. 1974 East Africa mammals. A atlas of evolutio i Africa (B): IIX 4704. BIBLVII (Academic Press Lodo ad New York). LINNAEUS C. 178 Systema aturae per rega tria aturae secudum classes ordies geera species cum characteribus differetiis syoymis locis 1 (ed. 10): 184 IIII (Salvii Holmiae). LYON M. W. 1906 Type of the geus Atherurus brushtailed porcupies. biol. Soc. Washig- U.S. a- to 19. 199. 1907 Notes o the porcupies of the Malay Peisula ad Archipelago. tio. Mus. : 794 pis. LIVLVII. MILLER G. S 190 Sevety ew Malaya mammals. miscell. Coll. 4 (140): 174 pi IXIX fig. 1. MOHR E 1964 Zur Nomeklatur ud Systematik der Quastestachler Atherurus F. Cu- Gattug vier 189. Saugetierk. 9 (): 9116 10 figs. OSGOOD W. H 19 Mammals of the Kelley-Roosevelts ad Delacour Asiatic expeditios. at. Hist. zool. ser. 18 (10): 1919 figs. 90 pis. IXXI. Mus. ROBINSON H. C. A. Mag. at. Hist. (8) 10 : 899. SEBA A. 174 Locupletissimi rerum aturalium thesauri... 1 : 1178 pis. pis. ICXI. SHAW G. 1801 Geeral zoology or systematic atural history (1): IV 16 pis. 116. (Lodo).