Research articles NAT. HIST. BULL. SIAM Soc. 54(2): 177-194，2 006 FORAGING ACTIVITY PATTERNS OFF RUGIVOROUS ORO MNIVOROUS ANIMALS ONT HE FOREST FLOOR OFA TROPICAL SEASONAL FOREST IN THAILAND， WITH REFERENCE TOS EASONAL CHANGES Shunsuke Suzuki1，Sh umpei Ki加mura2，3， Masahiro Kon1，Pi lai Poonswatf， Phitaya Chuailual，Ka mol Plongmaf，Ta kakazu Yumoto2ペNaohikoNo~， Tamaki Maruhashf and Prawat Wohandee6 ABSTRACf 明lepresent study quantitatively described出ediel forag凶gactivity pattems of企ugivorous or omnivorous animals on the forest floor of a位。'picalseasonal forest in Thailand and showed the seasonal changes in the activity pattems of some species. Thea ctivity pattems were investigated using automatic camera systems baited with fruits on出eforest floor. A to阻lof 11，133 pictures were taken. Forty-eight animal species could be identified in 10，955 pictures， including 30 species of mammals，17 birds and 1r eptile. Analyses were made for the diel activity pattems of animals for which 30 or more visits had been recorded. Based on白e tempora1 dis佐ibutionof visits，1 8 analyzed species were classified inωthe following three categories: (1) diumal species，no t active at night，(2 ) noctumal species，no t active during白e day，(3 ) others，ac tive during both day and night.百leresults of血eciassification were as follows: 2m ammals (Macaca nemestrina，Ca llosciurus finlaysonii) d3b irds (Chalcophaps 飢 indica，Pi tω， cyanea，Ga rrulax leucolophus) were diumal wi出叩almostunInlodal tempora1 distribution; 2m ammals (Tupaia belangeri，Me netes berdmorei) d3b irds (Lophura diardi， 飢 Gallus gallus，A rborophila chloropus) we問 diumalwith ab 泊lOd叫tempora1distribution; 5 mammals (Rattus spp.，Ni viventer fulvescens，Le opoldamys sabanus，Ma xomys surifer，Hy strix brachyura) were noctumal; 3m ammals (Tragulus javanicus，C ervus unicolor，M untiacus muntjak) were曲eothers.百lerewere significant differences in白etemporal distribution of visits between the rainy加d企yseasons for the Malayan porcupine (Hystrix brachyura)， Siamese fireback (Lophura diardi) and red jungle fowl (Gallus gallus). Key words: activity pattem，ca mera trapping，cr epuscular，d iumal，fe eding activity，K hao Yai National Park，no ctumal lSchool of Environmental Science，T he University of Shiga Pr巴fecture，Hikone，5 22-8533，J apan. s“l1ss叩uz此i砲@e切c.us叩p.aωc仏吋j. 匂2Cen陶tt飽e釘附蜘r伽fj0併r配E∞c01句0gicalResearch，K yoto University，O tsu，5 20-2113，Ja pan 3旬lailandHombill折口j民t，Department of Microbiology，F aculty of Science，M 油idolUniversi旬， Bangkok 10400，司副land 4Research Institute of Humanity and Nature，K yoto，6 02-0878，Ja pan 50epartment of Human and Cultu問，Musashi University，N erima，T okyo 17ι8534，Ja p 組 6National Park，Wi ldlife and Plant Conservation.Oep訂伽lent，B卸 gkok10900，明lailand Received 13 September 2005; accepted 25 July 2006. 177 178 SHUNSUKE SUZUKI F:f AL. 問τRODUCTION Tr'Opical f'Orests pr'Ovide en'Orm'Ously diverse and c'Omplex habitats and harb'Orm any mammal and bird species (HARR:rSON，1 962; M町RAEI' AL.，1 997; YASUDA ETA L.，20 05). WmTMO阻 (1990)presented as chematic figure describ泊gthe space and曲nep紅白i'O凶ng 'Ofm ammals in at r'Opical rain f'Orest in B'Ome'O， by which c'Oexistence may be facilitated. H'Owever，q uantitative studies 'Ona ctivity pattems，w hich釘ean essential c'Omp'Onent f'Or understanding time partiti'Oning，h ave been relatively insu伍cientf'Ora nimals 'Of住opical f'Orests (VAN SCHAIK & G阻FFITHS，1996; MIURA EI' AL.，1 997; KAWANISHI & SUNQUIST， 2004). Several studies c'Onducted in temperate z'Ones have sh'Own that activity pa仕ems'Of mammals change seas'Onally in acc'Ordance with envir'Onmental fact'Ors such as air temperature，d ay length，a nd precipitati'On( PEARSON， 1960; OSTERBERG， 1962; TANAKA， 2005). Alth'Ough tr'Opical seas'Onal f'Orests als'O experience marked seas'Onal changes in precipitati'On，n 'O由加gis kn'Owna b'Out seas'Onal changes in the activity pattems 'Ofa nimals living in tr'Opical seas'Onal f'Orests. In recent years，a ut'Omatic c創nerasystems have been utilized in ec'Ol'Ogical research 'On animals (PEARSON，1 960; CARLEY EI' AL.，1 970; G郎町WOOD，1978; GRIFFITHS & VA N SCHAIK， 1993; PEI， 1995; AKABAR & GORMAN， 1996; VAN SCHAIK & GRIF町田S，1996; MIURA EI' AL .，1997; BLANCHONG & SMALE，20 00; JAY ASEKARA EI' AL.，20 03; O'BRIEN EI' AL.，2 003; YASUDA，2 004). Such systems have proven useful f'Ori nvestiga出gactivity pattems because 'One 'Oft he advantages 'Ofa ut'Omatic camera systems is that they rarely disturb f'Ocal animals (OSTERBERG，19 62; CARLEY EI' AL.，19 70; VAN SCHAIK & GRIFF町HS， 1996; CUTLER & SWANN，1 999). Byu sing aut'Omatic camera systems baited with fruit，w eh ave elucidated血erelati'Onship between fruits characteristics and frugiv'Ores 'On血ef'Orest fl'O'Or'O fa 住opicalseas'Onal f'Orest in百lailand，which has distinct rainy and dry seas'Ons. As ar esult，w e accumu1ated c'Onsiderable data 'Ont he activity pattems 'Ofm any animal species. In this paper，f rrstly we describe the diel f'Oraging activity pattems 'Off rugiv'Or'Ous 'Or'O mniv'Or'Ous animals inhabiting 出e佐opicalseas'Onal f'Orest 'OfK ha'O Yai Nati'Onal Park，百lailand.We血eninvestigate出e presence 'Ora bsenc怠'Ofany seas'Onal changes in白eactivity pattems 'Oft hese species. STUDY AREA Thes tudies were c'Onducted fr'Om July 2000 t'O June 2002 in Kha'O Yai Nati'Onal Park ・ (hereafter KY，F ig. 1).官lep訂klies between the latitudes 'Of1 405'-15'Nand the l'Ongitudes 'Of1 01・05'-50・Ein血eD'Ongruk m'Ountain range， and c'Overs an釘ea'Of2 ，168 km2• Its elevati'On ranges from 250 t'O 1，351 m.百leprincipal study area was l'Ocated near the headquarters 'Oft he Nati'Onal Park and c'Overs appr'Oximately 70 km2 (KπAMURA EI'必円 2002，2005，S MITINAND，1 977 f'Ora d etailed descripti'On).百lestudy area ranged合om600 t'O 800 m asl in elevati'Ona nd was mainly c'Overed with m'Oist evergreen f'Orest which c'Overs appr'Oximately 64% 'Oft he t'Otal park area and extends between 400 and 1，000 m asl (SMIT町AND，1977).百lemean annual rainfall is 2，340 mm( 1993-2003) with the rainy seas'On usually 'Occurring企omApril t'O Oct'Ober and the dry seas'Onf r'Om N'Ovember t'O March (KITAMURA ETA L.，2 004). Them ean m'Onthly temperature ranges fr'Om 210C FORAGING ACTIVITY PAγTERNS OF FOREST ANIMALS 179 。 200N 150N 100N 1000E Figure1. Location of Khao Yai NaitonalPa rk (Decemb巴rand January) to 32"C (April and May). Although rip巴白uits(e.g. Ficus spp.) are available yeaJ可ound(POONSWAD ET AL.， 1998)，f ruit diversity and abundanceω'e relatively h.igh i.n出巴rai.nyseason and becoming scarce at the beginning of the dry season (BARTLEγr，2 003). METHODS Automatic Camera System We used two kinds of automatic camera syst巴ms.One consisted of a faJ'-infrared sensor and motor-driven compact cam巴rawith ab uilt-in tim巴pi巴ce.The d巴ta.ilsof this system are described i.n M1URA ET Aム(1997).Since April 2001，w e have used another system consisting of a compact camera with a built-in infrared motion sensor (S巴nsol・ Camera“FlliLDNOTE"，M ARIF Co. Ltd，Ja pan). Th巴detailsof this system are described in YA SUDA (2004). Camera systems wer巴seti.n various habitats'i ncluding primary forest， secondaryf orest，a nd th巴巴dgeof forest. Five to 50 fruits that had been col!ected合omthe ground釦oundpar巴nttrees were set at the base of the same tre巴sas bait，a nd c創ηeraswere placed approximately 2m away from the bai .tWhen fruits w巴I巴consumedby animalso r 180 SHUNSU阻 SUZU阻 ETAL. damaged by insects or microbes，th ey were replaced with new snples.Photography was 創 continued for at least 5c onsecutive days. Thec ameras were checked at intervals of one to伽'eedays，a nd films and batteries were replaced if necessary. In total，1 87 individual plants of 69 species were studied (Appendix 1). Wec ollected all plants not identified in the field and signedat empor.ycode. Plant specimens were 酪 紅 later identified by experts and/or compared with those identified specimens deposited in the Forest Herbarium in Bangkok (BKF) and出enrevised according to the Intemational Plant Name Index on the web site <ht甲://www.ipini.org/index.html>.These specimens are kept in our laboratory in Khao Yai National Park. Based on the pictures，w e were able to determine when and which animal species visited the fruit baits. Wef ollowed SRIKOSAMAT 成 A& HANSEL (1996) with P釘tial modifications by CORBET & 1也LL(1992) for the taxonomic nomenclature of m創nmals， and followed LE臥 GUL& ROUND (1991) for that of birds. Identiち岨gsmall mammals based solely on pictures might be difficult. However，w eh ave conducted住appingof small mammals in order to study their population dynamics加KY(SUZUKI ETA L.，un published data). Thus，w e were able to identify small mammals by comp紅泊g血osein the pictures with those血athad been trapped using their body size and color. Since the cameras were町iggeredcontinuously if an animal or ag roup of組加als stayed within the detection area of the sensor，o ne picωre did not necessarily represent a single visit by ag iven animal. To overcome this problem，w e followed O'B町EN(2003) and YA SUDA (2004) who regarded ap icωre as indica白19as ingle visit by ag iven species if the picture was taken morちthan30 minutes after the last picture of that species. Classi貨cationof ADimals sased OD Activity Patterns Activity pat飽mswere佃 alyzedfor each species by accumulating hourly visits for the duration of the s旬dyperiod.百lerefore，av ariable number of individuals con凶butedto白e data of ag iven species. 官lestudy period was divided into rainy (April to October) and dry (November to M紅ch)seasons. Differences in出.ediel activity pattems between the rainy and dry seasons were tested using血eKolmogorov-Sm註novtwo-sample test. This analysis was performed for animal species for which 40 or more visits were recorded in both ra泊yand dry seasons because at least 40 samples紅'eneeded for出istest. If there was no significant difference between rainy ddry seasons，th e data were pooled for further analyses. 佃 h血epresent study釘'ea，白e曲neof sunrise changes between 0545 and 0643 ho ver the ye民 and出atof sunset between 1743 and 1847 h. According to出estand紅dtime ne紅 KY，w e divided ad ay into the following three time zones: daytime仕om0700 to 1700 h; 凶ghtt泊施合'Om 1900 to 0500 h; crepuscular time from 0500 to 0700 h and 1700 to 1900 h. Based on the temporal dis凶butionof visits，a nimal species were classified into the following白reecategories: (1) diumal species，wh ich had never visited仕uitbaits in the nighttime; (2) noctumal one，wh ich had never visited血.efruit baits泊血.edaytime;組 d (3) the others，w hich had visited the fruit baits during both daytime and nighttime.百lIs classification was applied to釘山nalspecies for which 30 or more visits had been recorded， as 30 or more visits were required for statistically tes白19activity pattems (see below for 白.rtherexplanation). FORAGING ACTNITY PAITERNS OFF OREST ANIMALS 181 Thec hi-square test was performed for the activity pattems obtained for each species with the null hypothesis that all visits occurred at random independent of the time of day. Under出isnull hypothesis，出eexpected visit rates are 10/24，1 0/24 and 4/24 for the da戸ime，nighttime and crepuscular也ne，respectively. Since the determination of chi requires 出at血eexpected value for the crepuscular time (4/24) exceeds 5，a t otal of 30 or more visits were required. RESULTS Animal Species Photographed Weo btained at otal of 11，133 pictures conta白血g鉱山nals.Forty-eight animal species could be identified for 10，955 pic岡田 (98.4%of all pictures)，a nd血etime at which血e photograph was taken was also reorded.Thep hotographed animals included 30 species 心 of mammals，17 birds，an d 1r eptile (Table 2a nd Appendix 2). Mammal species accounted for 89.6% of all鉱山nalsidentified in the pictures，wh ich included one species of Scandentia， 1I nsectivora，1 P rimate，1 4 Camivora，1 P roboscidae，4 A rtiodactyla，a nd 8R odentia. Several chiropteran species were also photographed in 9p icωres，bu t these could not be identified to species. Of出etaxa血atwere identified to species，th e yellow 吋油rat(Maxomys surifer) was the most f詑quentlyphotographed (3，318 picωres)，f ollowed by白epig-tailed macaque (Macaca nemestrina; 1，665)叩 d血eIndochinese ground squirrel (Menetes berdmorei; 1，073).百lepictures with these 3s pecies accounted for 55.6% (6，056 pictures) of all白e pictes.Conversely，1 2 species of mammals，o ne species of reptile，a nd 6s pecies of birds 町 were photographed less than 10 times. Carnivora tended to be photographed inf同quently， and 9o f 14 Carnivora species were photographed less出an10 times. After omitting pictures of出esame species血atwere taken a負.eran interval of less than 30 min，4 ，819 of 10，955 pictures (44.0%) were regarded as representing an ew visit for ag iven species and used for further analyses. Seasonal Change of Activity Patterns 百leanimal species for which 40 or more visits were recorded泊 bo血 therainy and dry se部ons釘eIisted in Table 1( see also Table 2f or the activity pattems).百lerewere significant di首erences泊 thetemporal distribution of visits between the ra泊yand dry seasons for the Malayan porcup泊e(Hystrix brachyura)，th e Siamese frreback (Lophura diardi) and the red jungle fowl (Gallus gallus) (Kolmogorov.Smirnovtwo-sample test， ・ P< 0.05; Table 1，Fi g. 2). In the rainy season，H. brachyura exhibited unimodal activity pattems with ap lateau at between 2100 and 0200 h. However，in the dry season，ac tivity泊出isspecies was bimodal with peaks just after dusk djust before dawn (Fig. 2a). Lophura diardi exhibited 叩 almost entirely timodal activity in both the rainy and dry seasons，bu t the moming peak was slightly greater th佃 theaftemoon peak in the rainy season，b ut smaller泊血e合y season (Fig. 2b). Gallus gallus also exhibited bimodal activity with dist泊ctpeaks泊血e early moming and the late aftemoon泊 therainy season，w hereas the morning peak was 182 SHUNSUKE SUZUKI ET AL Tabl巴1. The results of Kolmogorov-Smirnov two-sample test for the sp巴Cl巴sfor which 40 visits or more wrercorddin both the rainy and dry seasons.T otal，t he 巴 巴 巴 total number of visits; Rainy，th e number of visits in the rainy season; Dry，th 巴 へ number of visits in the dry season; ns，n ot significant; p0.05; く Species name English name Total Rainy Dry D P Macaca nemestrina Pig-tailed macaque 350 233 117 0.085 Muntiacus mnりak Barking deel 259 135 124 0.078 ns l/ Cervus unico/OI Sambar deer 139 57 82 0.182 ns Menetes berdmorei lndochinese ground squirr巴l 527 396 131 0.049 ns I Niviventer fit/vescens Chestnutra t 194 60 134 0.139 ns Maxomys SUIザer Ye1l0w rajah rat 2081 1500 581 0.022 ns Hystrix brachyura MaJayan porcupine 216 152 64 0.291 Lophl/ra diardi Siamese fireback 219 106 113 0.198 ふ3 Gal/us ga//us Red junglefowl 112 67 45 0.274 司 a. Hystrix brachyura b.L ophura diarιfi c. Gallusg.且llus (D=0.291，p< O.Ol) ω=0.198，p< 0.05) (D=0.274，p <0.05) 25 Rainy (N=67) 20 90 30 “ 11 FOAURυ。 守喝ムA505 2100。 0。 。 12 18 0 6 6 12 18 6 12 18 言主 。A5 QL0 。anU 30 唱'晶50 tAU民 20 司A50 e守晶UKU。内 。 10。 。 12 18 0 6 6 12 18 6 12 18 Time ofday FigUl巴2.Activity patl巴rnsof th巴animalsthat巴xhibitedsignificant seasonalch ang巴8in b巴havior.Ab8cissa，ti me of day; ordinate，th e rate (%)of visits for eachho ur. Top，ra iny 8巴a80n，frol11 Apr lito Octob巴;rbo目OIn， dry season，f Tomトiovember10 March.Op en columll，1c aytim巴;gray column，c repuscular time; c108ecl columll，Il ighttime FORAGING ACTIVITY PATIERNS OFF OREST ANIMALS 183 obscured泊白edry season (Fig. 2c). In the other 5s pecies，ac tivity pattems were not significant1yd ifferent between rainy and dry seasons (Kolmogorov-Smimov two-sample test，P > 0 .05). Thus，fo r these species， 出edata for the duration of the study period were combined for further analyses. Diel Activity Pattern of Each Species 百le18 species for which 30 or more visits were recorded訂eclassified into血ree categories (Table 2; see Appendix 2f or the data of the species for which less than 30 visits were recorded). Four mammal and all of the 6b ird species are regarded as diumal.百lese diumal species may be further classified into two subcategories; one (D-l) conta加ing species that had almost unimodal activities with either ad istinct or av ague peak around noon or aftemoon，a nd the other (Iト2)containing species that were primarily bimodally active with distinct peaks in the moming and late aftemoon. Thef ormer subcategory， D-l，co ntained 2m ammals (M. nemestrina and Callosciurusfinlaysonii) and 3b ird species (Chalcophaps indica，Pi tta cyanea and Garrulax /euc%phus，Fi g. 3). Approximately 90% of 由emembers of both of these groups were recorded in the daytime (Table 2). Of出ese， M. nemestrina (Fig. 3a) and C. indica (Fig. 3c) were recorded most仕equentlyaround noon whereas C.fin/aysonii (Fig. 3b) and two bird species (P. cyanea (Fig. 3f) and G. /euc%phus (Fig. 3e)) were photographed more frequently in the aftemoon.百lelatter subcategory， Dー2，contained 2m ammals (Tupai・abe/angeri (Fig. 4a) and M. berdmorei (Fig. 4b)) and 3b ird species (L. diardi for both seasons (Fig. 2b)，G. gallus for both seasons (Fig. 2c)， and Arborophila ch/oropus (Fig. 4c)). This bimodal activity pattem was particularly conspicuous in the two mammal species，an d more than 50% of their visits were recorded in出ecrepuscular time (Table 2). Five rodent species (Rattus spp.，Ni viventer fu/vescens，Le opoldamys sabanus，M axomys surifer，a nd H. brachyura for both the rainy and dry seasons) were regarded as noctumal (Figs. 2a，an d 5)，wi th more than 90% of the visits for these species being recorded in the nighttime.百四erat species (Rattus spp.，N. fu/vescens and M. sur砕r)exhibited peaks in visiting frequency during the first hour of nighttime between 1900 and 2000 h. artiodactylspecies were recorded during both day and night (Fig. 6). Tragu/us 官官印 javanicus (Fig. 6a) and Muntiacus muntjak (Fig. 6c) were active mainly during day with peaks around the dawn and dusk，wh ereas Cervus unic%r (Fig. 6b) was active mainly at night although it was also photographed during the day. In all species for which 30 visits or more were recorded，th e temporal distributions of visits were significant1yd ifferent from the pattem expected from the null hypothesis (Table 2，C hi-square test，P < 0.05). DISCUSSION Comparison with Previous Studies There have been relatively few studies that have investigated the activity pattems of multiple animal species coexisting in tropical forests (VAN SCHAIK & GRIFFITHS 1996; MIURA EfA L. 1997). VAN SCHAIK & GRIFFITHS (1996) studied the activity pattems of 31 Table 2. Activity pattems of the species for which 30 or more visits were recorded. Pictures，to tal number of pictures; total visits，to tal 同∞ number of visi臼;Dayt加e，number of visits recorded during the dayt加e;Crepuscular time; number of visits recorded during hF 出ec問:puscul紅白ne;Nighttime，n umber of visits re氾ordedduring nighttime. Percentages of the total number of visits are shown in the parentheses. Abbreviations for the categories of activity pattems 'eas follows: 0-1，d iumal with unimodal 訂 x? distribution;凶， diumal with b加odaldistrib凶on;N，n octumal; 0，o thers.紳げ<0.001 for 制. F8I凶Iy S脚cies English 防御悶 Toω Daytime Crep眠叫釘 Nigb悩me t P 臼t司!Ory 岡田ー D81De visi包 time 。 。 S悶dentiaTupaudae Tupaia belangeri No揃lemttee shrew 190 121 54 (45) 釘 (55) (的 75.7 *** 0-2 防白lates ce。問pitheci伽E Macaca nemeslrina Pig-凶100macaque 1665 350 320 (91) 27 (8) (的 224.3 *** 0-1 Arti叫actyla 。 Tragulidae 1干'agulusjavanic山 Lesser Malay mouse-deer 177 96 51 σ3) 42 (斜) 3 (3) 44.8 *** 。 印 Cervi也s .Ce四郎unicolor Sambar 321 139 7 (5) 25 (18) 107。 (77) 54.6 *** 。 dZ臼国 SMcuiruiri也伽Ea MMRNCiaeauvtlnniet簡/votiesean明cscti ueu・rsbr uefsmru mlfi醐v即entsJjracaeyekinsson ii αBFInm創dl儲魁oacy回hnsigountn凶d'rs民aet rsgqruoiunn官dl sq'瓜rrel 192037340835825 52112590673469 z1ω 499。2。。 。σ(3め的{(59'3))) 3161837。37 (( 62η(((1495)))) 1ω757。4 ((()299。制151)))) 31254260963.1....38 465 *****ホ**'傘事**験 ** 00NN--21 Rodentia 回目CNdR己目白 Leopol，伽I)'ss伽，nus Noisy rat 120 73 。め( 。的(( 73 (100) ω 1. **事 N .h 。 ・ Maxomys surifer Yellow raj油rat 3318 2081 め( 134 1947 (94) 1375.7 *** N Dゆdi也s Hystrix brachyura (Rainy) Malay:叩 pOlCupIse 772 152 。仰) 8 (5) 144 (95) 103.7 *事* N 。 H. brac付附仰y) 329 64 {買} 7 (11) 57 (89) 38.4 *** N B凶s 。 GaJliformes ph悩ian拍車 Lophura diardi侭細川 Siam附 fireback 219 106 88 (83) 18 ()1。7) 。(0) 58.7 事** 0-2 L.仰rdi肋y) 208 113 105 (93) 8 。 (0) 73.5 *** 0-2 m 。 Galllωgallus侭細川 ROO junglefowl 108 50 (75) 17 5) (0) 35.4 *** 0-2 。 G.galllω肋y) 101 45 27 (60) 18 (相) (0) 24.6 *** 0-2 Arborophila chloropus Scaly-h毘astω抑rtridge 62 40 27 (68) 13 (33) 。的( 21.2 *** 0-2 Columbiformes 。 印刷bi加 C加Icophapsindica Emerald dove 130 100 97 (97) 3 (3) (0) 73.2 *** 0-1 。 Pas町iformesPi凶伽e Pitta cyanea Blue pitta 42 31 27 (87) 4 (13) (0) 18.0 *** 0-1 Musci悶pid国e Garrulax leucolophus White-cres凶 lau凶i刷m由 65 46 43 (93) 3 (7) (0) 30.3 *** 0-1 FORAGING ACTIVITY PATTERNS OF FOREST ANIMALS 185 、 a. MSCBCS nemestnns CallosciurusnnJSysonii 20 (N=350) 20 15 15 10 10 。5 。5 。 。 6 12 18 6 12 18 *- 巳.ChsJcophsps indics 20 (N=100) 16 仏1=31) 16 15 12 12 10 8 8 。5 。4 。4 。 。 。 6 12 18 6 12 18 6 12 18 'l'imeo fday Figur巴3.Activity patternso f the animalsc1 assifi巴das diurnalsp 巴cieswith an almostu nimodaldi stribution. See Fig. 2f or further explanations. a. Tupaia belangeri b. Menetes berdmorei c. Arborophila chloropus 40可 (N=121) 30可 (N=527) 20、(N=40) 30 15 20 求 20 10 10 10 5 。 。 。 。 0 。 6 12 18 6 12 18 6 12 18 Timeofday Figure 4. Activity patt巴rnsof th巴animalsc1assified as diurnals pecies with ab imodaJ distribution. S巴巴 Fig.2 for further explanations 186 SHUNSUKE SUZUKJ ET AL a. Rat tus remotus 、• Niviventerおlvescens 25 (N=63) 20可 (N=194) 20 15 15 10 10 5 。5 。 。 。 12 18 6 12 18 6 ヌc.Leopoldamys sabanus d. Maxomys SUl鵠 r 20 (N=73) 20 (N=2081) 可 可 15 15 10 10 5 5 。 。 。 。 12 18 6 12 18 6 Time ofday FigUl巴5.Activityp atternso f th巴animalsclassified as nocturnal sp巴CI巴s.See Fig. 2f or further explanaitons a. Trsl{Ulusjavanicu8 b. Cervus unicolor c. Muntiacu8 munijak 、 20、(N=9ω 15 15 (N=259) 15 10 10 10 ~ 5 5 。5 。 。 。 。 。 6 12 18 6 12 18 6 12 18 τ 'imeo f day Figure6 . Activity patternso f th巴animalsclassified as being active dur.ing both the day and night. S巴巴 Fig.2 for furth巴rexplanations