NAT. NAT. HIST. BUL L. SIAM S配. 55 (1): 187-192 ， 2007

Observations Observations of Sma Il -toothed Palm Civets Arctogalidia 的ivirgata in in K.h ao Yai National Park ，Thailand ， with Notes on Feeding Techniques

J. J. W. Duckworth 1 andA. R. Nettelbeckf

Small-toothed Small-toothed Palm Civet (=Three-striped Palm Civet) Arctogalidia trivirgata occurs widely widely across . Historically it has sometimes been considered as a scarce or declining declining species ， or one confined to remote 釘 eas. This note documents observations from Kh ao Yai National Park ，which is among the sites in Southeast Asia best surveyed for mam- mals. mals. In the context of other recent records ， there is no compelling reason to consider the species species as rare ，localised or declining ， but instead simply one 由at ， through its behaviour ，is easily easily overlooked by standard mammal collecting and survey techniques.

Field Field observations During 5-9 May 1995 ， Small-toothed Palm Civets were found three times (two sI!l- gles ，one duo) along the main road north of the park headqu 紅 ters and visitor cen 住'e ， in 7'/4 hours hours spot-lighting by foot (following methodology ofDuCKWORTHETAL. ， 1994). All were in 白e canopy of roadside broad-leaved evergreen forest (see SMITINAND ， 1968) at c. 760-800 maltiωde. Animals were active ，crashing noisily as they ran (as typical ofthe species: DUCK- WOR 叩， 1997) and ，twice ，making a spectacul 紅 canopy-to ・canopy leap across the main road north north of the headquarters. Such leaps were preceded by vigorous to-and- 企o sh 嘘 ing ofthe jump-off jump-off perch ， thereby decreasing substantially the distance to be covered. All were calling ， with with two call-types just as given in Laos (see DUCKWORTH ， 1997): loud disyllabic snorts ， and soft ，rather hollow-sounding ， tchuck calls ， given about once per second. On 12 October 1995 ， a silent male Small-toothed Palm Civet was found eating figs in a small Ficus hispida (Moraceae) (Moraceae) ， only 8 feet above ground ， near the headquarters at 2225 h，and later watched for for 90 minutes (see below); the animalle 負 the bush by walking along the roadside wires (re- calling 血 e agility on ropes of a shipboard captive noted by S百 RNDALE [1884] ，阻dBANKS'S [1931] [1931] general admiration for its climbing abilities). On 21 Janu 紅 y 1996 two were observed at at about 2100 h in a1 紅 ge fI owering tree by 血e park headquarters ， foraging singly ，calling (with (with ‘high-pressure' snish notes) and moving actively; at one stage ，one animaljumped onto the the roof of a building ，making a loud bang. τ'h e animals were identified by their diagnostic calls and the clear views 白rough lO x40 binoculars binoculars under illumination by a hand-held Nitech Xcell halogen spot-light (100 ，000 cp).

Standard Standard pelage features and the diagnostic e紅 pattem (naked pink inner surface with thick dark dark intrusions 仕om the rim ， with duller pink ，lightly-haired ， outer surface) of the northem rac 四 ofthe species (see VAN BEMMEL ， 1952) were readily visible ， because no animal fI ed

IWild1ife IWild1ife Conservation Society; current address ，East Redham Farm ，Pilning ，Bristo1 ， BS35 4JG ， U.K. 2Ho1steinisches 仕asse 54 ，D-10717 Berlin ，Germany. Received Received 6 November 2006; accepted 26 February 2007

187 187 188 188 1. W. DUCKWORTH AND A. R. NETTELBECK

fr 'O m i1l uminati 'O n. The species typically permits a cl 'O se appr 'O ach and prol 'O nged viewing (e.g. W ALSTON & DUCKWOR 四， 2003; BORISSENKO ET AL. ， 2004). During 由 e visit 'O fMay 1995 ， tw 'O C 'O mm 'O n Palm Civets Paradoxurus hermaphroditus were als 'O 'O bserved; alth 'O ugh n 'O ther 'O ther species can realistically be misidentified as Small-t 'O'O thed Palm Civet ，this latter ，al 'O ng with with Masked Palm Civet Paguma larvata ，is pr 'O bably 血em 'O st likely s'O urce 'O fc 'O nfusi 'O n f'O r inexperienced 'O bservers. Giant flying squirrels Petaurista may c 'O nceivably als 'O be misi- dentified dentified as arb 'O real civets. JWD was already familiar with all these species 企om extensive n 'O cturnal mammal surveys in La 'O s (DUCKWORTHETAL. ， 1994; DUCKWORTH ， 1997; EVANS ET AL. ， 2000). Th e Small-t 'O'O thed Palm Civet 'O bservati 'O ns in Kha 'O Yi 討 were spread acr 'O ss many m 'O nths and several miles 'O fr 'O ad ，precluding any p'O ssibility that 血ey might refer t 'O released released pets. The latter is ap 'O ssible s'O urce 'O f any n'O vel rec 'O rd ar 'O und the p紅 k's headquar- ters ters (LYNAM ET AL. ， 2006: 81 ，82 ，209) ， and evidently explains s'O me an 'O mal 'O us rec 'O rds 'O f

Small 目 t'O'O thed Palm Civet elsewhere (e.g. Su Su ， 2005). Given Given recent rec 'O rds fr 'O m evergreen 'O r semi-evergreen f 'O rest fr 'O m Huay Kh a Kh aeng Wildlife Wildlife Sanctuary (RAB 町 OWIπ ， 1991; CONFORTI ， 1996) t'O出 .e west 'O fKh a 'O Yai ，Phu Kh ie 'O Wildlife Wildlife Sanctuary (GRASSMAN ， 2004) t'O the n'O rth ，vari 'O us La 'O sites (DUCKWORTH ， 1997) t 'O the n'O rth-east ， and Camb 'O dia (W ALSTON & DUCKWOR 百， 2003) t'O恥 east ，出 e species' 'O ccu 町'ence in Kh a 'O Yai is n'O t at all unexpected. With n 'O published survey using appr 'O priate meth 'O d'O l'O gy acr 'O ss the Nati 'O nal Park ，it is unclear h'O w wid~spread 'O r abundant the species is is there. Th ec 'O ntact 企'equency 'O f appr 'O ximately 'O nce per 2' /2 h'O urs in May 1995 ，alth 'O ugh based based 'O nt 'O'O few 'O bservati 'O ns t 'O be definitive ，c 'O mpares well with th 'O se 'O f 'O nce per 4h 'O urs in in Cat Tien Nati 'O nal Park ， Vietnam (BORISSENKO ET AL. ， 2004) and 'O nce per 7-8 h'O urs f'O r vari 'O us sites in La 'O s (DUCKWORTH ，1997). 百 is may reflect 恥'O ptimal search c'O nditi 'O ns pr 'O vided by Kh a 'O Yai's r'O ad (a l'O ng viewing distance with g'O'O d sighting int 'O the can 'O py) and ，perhaps ， the l'O w levels 'O f hunting ，at least f'O r species 'O fl 'O wc 'O mmercial value ，in 出 e headquarters headquarters area 'O f the p紅 kf 'O r the preceding tw 'O decades (NE'π 'E LBECK ， 1997).

Detection Detection of tbe species and requirements for survey

Th e species was missing 合'O m lists 'O f Kha 'O Yai mammals by b'O th SRIKOSMATARA & HANSEL (1996) and L YNAM ET AL. (2006). Th is d 'O ubtless reflects tw 'O reas 'O ns advanced l'O ng ag 'O by POCOCK (1939) t 'O expl 必n 出 e paucity general 'O f field 'O bservati 'O ns 'O n 出 e species ， and by V AN BEMMEL (1 952) f'O r the l'O w numbers 'O f museum specimens. Firstly ，it is str 'O ngly n'O ctumal (STE 悶 D 此 E， 1884; BANKs ， 1931; HARRISON ， 1962; BATTEN & BATTEN ， 1966; DUCKWORTH ，1997) ， and extensive 'O bservati 'O ns by day during 1992-1996 int 'O仕 uit 泊出is part 'O f Kh a 'O Yai ，閲覧icient t 'O detect many Bintur 'O ngs (NETTELBECK ，1997) ，f 'O und n 'O Small-t 'O'O thed Palm Civet (ARN ，pers 'O nal 'O bservati 'O ns). Sec 'O ndly ，it 'O nly very rarely c'O mes t 'O the gr 'O und. Al th 'O ugh 由e genus seems never t 'O have been subject t 'O a field s旬dy ， its its general anat 'O my (VAN BEMMEL ， 1952) c'O upled with 'O bservati 'O ns 'O f captive animals (e.g. sτ 芭RNDALE ， 1884) and general field sightings (e.g. HOSE ， 1893; DAVIS ， 1962; PAYNE ET AL. ， 1985;RAB 町 OWI 'IZ， 1991) indicate str 'O ng arb 'O reality. HAR 即 SON (1962) categ 'O rised 血especies as ‘undercan 'O py' rather than ‘can 'O py' ，but 由is' is evidently n'O t appr 'O priate in at least PartS 'O f the range. Of 32 field c 'O ntacts in La 'O s during 1992-1996 ，23 were in 血e can 'O py ，8 in the midst 'O rey ， and 1 in the underst 'O rey; n'O ne descended t 'O the fl ∞Ir (DUCKWORTH ，1997) ，alth 'O ugh GRASSMAN (2004) did catch 'O ne in a gr 'O und-level b'O x 住ap baited with a live chicken. 百lese habits habits mean that tw 'O c'O mm 'O n survey techniques f'O rn 'O ctumal animals ，c 創 nera-trapping 佃 d OBSERVATIONS OF SMALL-TOOTHED PALM CIVETS ARCTOGALIDIA TRNIRGATA 189

examination examination of hunted anim a1 s in markets and villages (many of which 紅 e sn 紅 'ed) ，greatly under-record under-record this civet ，by comparison with ground-dwelling sm a1 1 carnivores. Furthermore ， searches searches for footprints would be unlikely to detect Sm a1 1-toothed Pa1 m Civet ，and it is anyway unclear unclear whether its prints 釘 'e distinct from other similarly-sized Southeast Asian carnivores. Finally ，village reports about sm a1 1 carnivores demand extreme care if identification to spe- cies cies is a恥 mpted ，al 出ough given 白is species' distinctive behaviour and ca1 1s ，habitual night hunters hunters might well know and recognise Small-toothed Palm Civets. Active Active spot-lighting is essenti a1 in surveying Sm a1 1-toothed Palm Civets. Recent spot- light light searching showed 白紙 the species is common in evergreen forest in south and centr a1 Laos (DUCKWORTH ， 1997; the north remains unsurveyed by this technique) ，and provided the first first record ， other than interpolation on generalised range maps ， for Cambodia (W ALSTON & DUCKWORTH ， 2003) and 出 e first recent record in Vi etnam (BORISSENKO ET AL. ， 2004). Both these these latter were from sites (Seima Biodiversity Conservation Ar ea and Cat Tien National Park) ，where ，部 in Kh ao Yai ，extensive large mammal surveys using a conventional mix of dium a1 observation ，camera-trapping ， sign searches and discussions with local people failed to to find the species ，despite generating impressive complements of other large mammal species (W ALSTON & DUCKWOR 四， 2003; POLET & L 別 G ， 2004). Small carnivore community assess- ments relying largely on c制 nera-trapping find the species only incidentally (e.g. HOLDEN ， 2006 ，LONG & MINH HOANG ，2006) ， or not at all (e.g. Az LAN ，2003 ，JOHNSON ET AL. ， 2006). No survey should conclude that Small-toothed Pa1 m Civet is absent or even scarce at a site unless unless 閲覧icient spot-lighting has been undertaken. For similar reasons ， past opinions that the the species is rare (e.g. DANG HUY HUYNH & PHAM TRONG ANH ， 1974) or in strong decline (e.g. (e.g. LEKAGUL & McNEELY ， 1977) should be questioned.

Feeding behaviour Because Small-toothed Palm Civet usually feeds high in the canopy ，we had not observed its its precise feeding methods previously (and have done so only once since); nor do they s田 mto have have been recorded in the literature ， even in descriptions of captive anim a1 s such as Sτ 'E RNDALE

(1 884) ，BANKS (1 931) ，HEUBEL (1 940) and Bx 甘 EN & BATTEN (1 966). The 佃 imalon 12-13 October October 1995 a鉦'o rded an excellent oppo 託unity to observe it feeding during 000 0-0 130 h at a range of 4- 8 m. lt fed on very juicy clustered cauliflorous figs Ficus hispida of c. 2.5 cm diameter. diameter. Most in the sm a1 1tree were green and u町 ipe. Ripe figs were more yellow in colour ， and the civet selected these by sniffing each fruit for a second or so at c. 2.5 cm range ， as it moved its head slowly past each clump of figs. Th e anim a1 plucked the whole fig from 白e branch branch with its mouth ，and 血en made 2-5 rapid ，inaudible ， mandibulations seemingly to get the the fruit into position and ，perhaps ， to split its skin. The civet 血.en squeezed its jaws together deliberately deliberately (with clearly audible squelching noises) a number of times 1-2 seconds ap 釘 t(1 4， 15 ，8，24 ，20 ，15 ，25 ，15 ，16 ，6 times on 10 respective fruits). Seemingly ， the tongue pressed the the fruit to the teeth and/or roof of the mouth and squeezed or rubbed out the fig juice between jaw movements. The squashed residue of rind ，flesh and the seeds (which latter were very small) ，was then allowed to drop ('spat ou t' would be ωo active a verb). During much of the observed observed feeding period ， the civet wagged its ears slowly but almost ceaselessly. A frugivore is likely to handle fruits in different ways depending on the spe- cies. cies. For example ， Paradoxurus jerdoni usually consumes seeds but with the the large ，fleshy 企uit of Palaquium eU 伊ticum () it either chews up or drops out 190 190 J. W. DUCKWORTH AND A. R. NEITELBECK

出e seeds (D. MUDAPPA in litt. 2006). It seems possible that fruit-pressing may be a habitual feeding feeding style of Small-too 血ed Palm Cive t: the species' singular dentition ， with small peg-like premolars premolars and molars ， non-occluding molar rows even when jaws shut ， as well as an undu- lating lating palate (G 阻 GORY & HELLMAN ， 1939; POCOCK ， 1939; LEKAGUL & McNEELY ， 1977) would seem to assist 出is sort of 仕uit-processing. According to POCOCK (1 939) ，somewhat similar similar characters 釘 'e shown by Binturong Arctictis binturong ， another civet which eats much

fruit fruit (LAMBERT ， 1990; NETTELBECK ， 1997). Fruit 幽 pressing is evidently but one of several feeding techniques techniques feeding used by Small-toothed Palm Civet ，and indeed recorded stomach contents could could seem to suggest that 由e species is quite carnivorous (POCOCK ， 1939; DAVIS ， 1962; LIM BOO LIAT & BET 四 RτON ， 1977). Notably ，HARRISON (1 962) was pe 中lexed when he found only only animal material in two stomachs examined ，despite captives much prefe 凶 ng fruit over flesh. flesh. However ，fruit-pressing and ， with some plant species ，any form of fruit consumption would leave very few ，potentially no ， remains in the stomach (or in faeces) identifiable by conventional conventional methods ，and such diet 紅 y analyses will not ，therefore ，reflect the true relative 凶 portance of 仕ui t. Moreover ，fruit ，however consumed ，is often well digested and absorbed 泊to the system ，making its detection difficult for species with either few large seeds ，e.g. mango Mang iJ era indica ， or insignificant seeds e.g. cultivated bananas Musa spp. (D. Mu- DAPPA in litt. 2006). Civets are said to be significant seed dispersal agents in Asian forests ， wi 血 Small-toothed Palm Civet sometimes highlighted because of its 位 ong frugivory (e.g. PAYNE 1995 ，MEIJAARDETAL. 2005). However ，fruit-pressing will 問 sult in minimal disper- sal sal at best for the fig. Squ 註rels also shed most or all seeds in the uneaten fig residue ，and hence hence have no value for the in fig dispersal (LAMBERT ， 1990); by contrast ， the other chiefly frugivorous frugivorous sympatric civet ，Binturong ， does ingest and 也en pass out quantities large of fig seeds seeds (LAMBERT ，1990) ， as does Brown Palm Civet (MUDAPPA ， 2001). Small-toothed Palm

Civet's Civet's peculi 紅 dentition suggests that fruit-pressing is among its main feeding techniques. However ，a single observation c組 allow only a tentative suggestion ，and full studies of the species' species' feeding behaviour and role in seed dispersal would be most informative. Acknowledgments.-ARN thanks the National Research Council ofThailand for per- mission mission to conduct gibbon research in 血e course of which these sightings were made ， the 出en superintendent Mr Vallobh and other staff of Kh ao y:泊 National Park ，and the German Exchange Service (DAAD) for funding. JWD thanks the Wildlife Conservation Society Lao Pro gramme which covered the costs of his May 1995 visit and we bo 白 thank Warren Brock- elman for regular support and advice ，including 出 e identification of the fig tree and editorial work on the note. We thank Lon Grassman Jr for information about his record of the species ， Divya Mudappa and Philip Round for commenting on an earlier draft ，Ted Tizard for sourc- ing ing some nowadays obscure references ，and Harry Van Rompaey for his bibliographic work which alerted us to the latter. OBSERVATIONS OF SMALL-TOOTHED PALM CIVETS ARCTOGALIDIA TRNIRGATA 191

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