Research articles NAT. HIST. BULL. SIAM Soc. 54(2): 177-194 ，2006
FORAGING ACTIVITY PATTERNS OF FRUGIVOROUS OR OMNIVOROUS ANIMALS ON THE FOREST FLOOR OF A TROPICAL SEASONAL FOREST IN THAILAND ， WITH REFERENCE TO SEASONAL CHANGES
Shunsuke Suzuki 1，Shumpei Ki 加 mur a2，3， Masahiro Kon 1 ，Pilai Poonswatf ， Phitaya Phitaya Chuailual ，Kamol Plongmaf ， Takakazu Yumoto 2ペNaohiko No~ ， Tamaki Maruhashf and Prawat Wohandee 6
ABSTRA Cf
明le present study quantitatively described 出e diel forag 凶 g activity pattems of 企ugivorous or or omnivorous animals on the forest floor of a 位。 'pical seasonal forest in Th ailand and showed the the seasonal changes in the activity pattems of some species. Th e activity pattems were investigated investigated using automatic camera systems baited with fruits on 出e forest floor. A to 阻lof 11 ， 133 pictures were taken. Forty -e ight animal species could be identified in 10 ，955 pictures ， including including 30 species of mammals ， 17 birds and 1 reptile. An alyses were made for the diel activity activity pattems of animals for which 30 or more visits had been recorded. Based on 白e tempo ra1 dis 佐ibution of visits ， 18 analyzed species were classified inωthe following three categories: categories: (1) diumal species ， not active at night ， (2) noctumal species ， not active during 白e day ， (3) others ， active during both day and nigh t.百 le results of 血e ciassification were as
follows: follows: 2 mammals (Macaca nemestrina ，Callosciurus finlaysonii) 飢 d3 birds (Chalcophaps
indica ，Pit ω， cyanea ， Garrulax leucolophus) were diumal wi 出叩 almost un Inl odal tempo ra1
distribution; distribution; 2 mammals (Tupaia belangeri ， Menetes berdmorei) 飢 d3 birds (Lophura diardi ，
Gallus Gallus gallus ， Arborophila chloropus) we 問 diumal with ab 泊lO d叫 tempo ra1 distribution; 5 mammals (Rattus spp. ，Niviventer fulvescens ，Leopoldamys sabanus ，Maxomys surifer ， Hystrix brachyura) brachyura) were noctumal; 3 mammals (Tragulus javanicus ， Cervus unicolor ， Muntiacus muntjak) muntjak) were 曲e others. 百lere were significant differences in 白e temporal distribution of visits visits between the rainy 加 d 企y seasons for the Malayan porcupine (Hystrix brachyura) ， Siamese Siamese fireback (Lophura diardi) and red jungle fowl (Gallus gallus).
Key words: activity pattem ， camera trapping ，crepuscular ，diumal ， feeding activity ，Kh ao Yai National National Park ， noctumal
lSchool lSchool of Environmental Science ，The University of Shiga Pr 巴fecture ，Hikone ，522-8533 ， Japan. s“l1 ss 叩uz 此 i砲@e 切c.us 叩p.aωc 仏吋 .j 匂2Cent 陶蜘t飽附e釘r 伽fj 0併rE配∞c010 句gical Research ，Kyoto University ，Otsu ，52 0- 2113 ， Japan 3旬lailand Hombill 折口j民 t，De partment of Microbiology ， of Faculty Science ，M 油 idol Universi 旬， Bangkok 10400 ，司副land 4Research 4Research In stitute of Humanity and Nature ，Kyoto ，60 2-0 878 ， Japan
50epartment 50epartment of Human and Cultu 問， Musashi University ，Nerima ，Tokyo 17ι8534 ，Jap 組
6National 6National Park ，Wildlife and Plant Conservation.Oep 訂伽lent ，B 卸 gkok 10900 ，明 lailand Received Received 13 September 2005; accepted 25 July 2006.
177 177 178 178 SHUNSUKE SUZUKI F:f AL.
問 τRODUCTION
Tr 'O pical f 'O rests pr 'O vide en 'O rm 'O usly diverse and c'O mplex habitats and harb 'O r many mammal and bird species (HARR:r SON ， 1962; M 町 RA EI' AL. ， 1997; YASUDA ET AL. ， 2005). WmTMO 阻 (1990) presented a schematic figure describ 泊g the space and 曲nep 紅白i'O凶 ng 'O f mammals in a tr 'O pical rain f 'O rest in B 'O me 'O， by which c 'O existence may be facilitated. H 'O wever ，quantitative studies 'O n activity pattems ，which 釘 e an essential c'O mp 'O nent f'O r understanding understanding time partiti 'O ning ，have been relatively insu 伍 cient f'O r animals 'O f 住opical f 'O rests (VAN SCHAIK & G 阻 FFITHS ， 1996; MIURA EI' AL. ， 1997; KAWANISHI & SUNQUIST ， 2004). 2004). Several Several studies c'O nducted in temperate z'O nes have sh 'O wn that activity pa 仕ems 'O f mammals change seas 'O nally in acc 'O rdance with envir 'O nmental fact 'O rs such as air temperature ，day length ，and precipitati 'O n (PEARSON ， 1960; OSTERBERG ， 1962; TANAKA ， 2005). 2005). Alth 'O ugh tr 'O pical seas 'O nal f 'O rests als 'O experience marked seas 'O nal changes in precipitati 'O n ，n 'O由加 g is kn 'O wn ab 'O ut seas 'O nal changes in the activity pattems 'O f animals living living in tr 'O pical seas 'O nal f'O rests.
In In recent years ，aut 'O matic c創 nera systems have been utilized in ec 'O l'O gical research 'O n animals animals (PEARSON ， 1960; CARLEY EI' AL. ， 1970; G 郎町WOOD ， 1978; GRIFFITHS & V AN SCHAIK ， 1993; PEI ， 1995; AKABAR & GORMAN ， 1996; VAN SCHAIK & GRIF 町田S， 1996; MIURA EI' AL ，. 1997; BLANCHONG & SMALE ， 2000; JA YASEKARA EI' AL. ， 2003; O'BRIEN EI' AL. ， 2003; YASUDA ， 2004). Such systems have proven useful f'O r investiga 出 g activity pattems pattems because 'O ne 'O f the advantages 'O f aut 'O matic camera systems is that they rarely disturb disturb f'O cal animals (OSTERBERG ， 1962; CARLEY EI' AL. ， 1970; VAN SCHAIK & GRIFF 町 HS ， 1996; 1996; CUTLER & SWANN ， 1999). By using aut 'O matic camera systems baited with fruit ，we have elucidated 血e relati 'O nship between fruits characteristics and frugiv 'O res 'O n 血ef 'O rest fl 'O'O r 'O fa 住opical seas 'O nal f 'O rest in 百lailand ，which has distinct rainy and dry seas 'O ns. As a result ，we accum u1 ated c 'O nsiderable data 'O n the activity pattems 'O f many animal species. In this paper ，frrstly we describe describe the diel f'O raging activity pattems 'O f frugiv 'O r'O us 'O r 'O mniv 'O r'O us animals inhabiting 出e 佐opical seas 'O nal f 'O rest 'O f Kh a 'O Yai Nati 'O nal Park ，百lailand. We 血en investigate 出e presence presence 'O r absenc 怠'O f any seas 'O nal changes in 白 e activity pattems 'O f these species.
STUDY AREA
Th e studies were c'O nducted fr 'O m July 2000 t 'O June 2002 in Kh a 'O Yai Nati 'O nal Park (hereafter (hereafter KY ， Fig. 1).官 le p訂 k lies between the latitudes 'O f 14 ・05'-15'N and the l'O ngitudes 'O f 101 ・05'-50 ・E in 血eD 'O ngruk m 'O untain range ， and c'O vers an 釘 ea 'O f2 ， 168 km2• Its elevati 'O n ranges from 250 t 'O 1， 351 m. 百 le principal study area was l'O cated near the headquarters headquarters 'O f the Nati 'O nal Park and c'O vers appr 'O ximately 70 km 2 (KπAMURA EI' 必円 2002 ，2005 ，SMITINAND ， 1977 f'O ra detailed descripti 'O n). 百 le study area ranged 合om600 t 'O 800 m asl in elevati 'O n and was mainly c'O vered with m 'O ist evergreen f 'O rest which c'O vers appr 'O ximately 64% 'O f the t 'O tal park area and extends between 400 and 1，000 m asl (SMIT 町 AND ，1977). 百le mean annual rainfall is 2，340 mm (1993-2003) with the rainy seas 'O n usually 'O ccurring 企om April t 'O Oct 'O ber and the dry seas 'O n fr 'O m N 'O vember t 'O March (KITAMURA ET AL. ， 2004). Th e mean m 'O nthly temperature ranges fr 'O m 21 0 C FORAGING ACTIVITY PAγTERNS OF FOREST AN IM ALS 179 。
20 0 N
15 0 N
10 0 N
100 0 E
Figur e 1. Location of Khao Yai Na ti ona l Park
(Decemb 巴r and January) to 32" C (April and May). A lthough rip 巴白uits (e.g. Ficus spp .) are are available ye aJ可 ound (POONSWAD ET A L.， 1998) ， fruit diversity and abundanceω'e re lat iv e ly hi. gh i.n出巴 ra i.n y season and becoming scarce at the beginning of the dry season (BARTLEγr ， 2003).
METHODS
Automatic Camera System
We used two kinds of automatic camera syst 巴ms. One consisted of a faJ '-infrared sensor sensor and motor-driven compact cam 巴ra with a built-in tim 巴pi 巴ce. The d巴ta i. ls of this system are described i.n M 1U RA ET A ム(1 997). Since April 2001 ，we have used another system consisting of a compact camera with a built-in infrared motion sensor (S 巴nsol ・
Camera “FlliLDNOTE" ，MARIF Co. Ltd ， Japan). Th 巴 detail s of this system are de scr ibed in in Y ASUDA (2004). Camera syste ms wer 巴set i.n various habitat s 'including primary forest ， seconda ry forest ，and th 巴巴dge of fores t. Five to 50 fruits that had been co l! ected 合om the ground 釦 ound par 巴nt trees were set at the base of the same tre 巴s as bait ，and c創 ηeras were placed placed approximately 2 m away from the bai t. When fruits w 巴I巴 cons um ed by an im als or 180 180 SHUNSU 阻 SUZU 阻 ETAL.
damaged by insects or microbes ， they were replaced with new s創 nples. Photography was continued continued for at least 5 consecutive days. Th e cameras were checked at intervals of one to 伽 'ee days ，and films and batteries were replaced if necessary. In In total ， 187 individual plants of 69 species were studied (Appendix 1). We collected all all plants not identified in the field and 酪 signed a tempor. 紅 y code. Plant specimens were later later identified by experts and/or compared with those identified specimens deposited in the the Forest Herbarium in Bangkok (BKF) and 出en revised according to the In temational Plant Plant Name In dex on the web site
Classi 貨cation of ADimals sased OD Activity Patterns
Activity Activity pat 飽ms were 佃 alyzed for each species by accumulating hourly visits for the duration duration of the s旬 dy period. 百lerefore ， a variable number of individuals con 凶buted to 白e data data of a given species. 官le study period was divided into rainy (April to October) and dry (November to M 紅 ch) seasons. Differences in 出.e diel activity pattems between the rainy and dry seasons were tested using 血e Kolmogorov-Sm 註nov two-sample test. This analysis was performed for for animal species for which 40 or more visits were recorded in both ra 泊y and dry seasons because because at least 40 samples 紅 'e needed for 出is tes t. If there was no significant difference between between rainy 佃 d dry seasons ， the data were pooled for further analyses. h 血e present study 釘 'ea ，白 e 曲ne of sunrise changes between 0545 and 0643 h over the the ye 民 and 出at of sunset between 1743 and 1847 h. According to 出e stand 紅 d time ne 紅 KY ，we divided a day into the following three time zones: daytime 仕om 0700 to 1700 h; 凶ghtt 泊施合'O m 1900 to 0500 h; crepuscular time from 0500 to 0700 h and 1700 to 1900 1900 h. Based on the temporal dis 凶bution of visits ， animal species were classified into the following 白ree categories: (1) diumal species ，which had never visited 仕uit baits in the nighttime; nighttime; (2) noctumal one ，which had never visited 血.e fruit baits 泊血.e daytime; 組 d (3) (3) the others ，which had visited the fruit baits during both daytime and nighttime. 百lI s classification classification was applied to 釘山nal species for which 30 or more visits had been recorded ， as as 30 or more visits were required for statistically tes 白19 activity pattems (see below for 白.rther explanation). FORAGING ACTNITY PAITERNS OF FOREST ANIMALS 181
Th e chi-square test was performed for the activity pattems obtained for each species with with the null hypothesis that all visits occurred at random independent of the time of day. Under 出is null hypothesis ，出 e expected visit rates are 10/24 ， 10/24 and 4/24 for the da 戸ime ，nighttime and crepuscular 也 ne ，respectively. Since the determination of chi requires 出at 血e expected value for the crepuscular time (4/24) exceeds 5， a total of 30 or more visits visits were required.
RESULTS
Animal Species Photographed
We obtained a total of 11 ， 133 pictures conta 白血g 鉱山nals. Forty-eight animal species could could be identified for 10 ， 955 pic 岡田 (98 .4% of all pictures) ，and 血e time at which 血e photograph photograph was taken was also re 心orded. Th e photographed animals included 30 species of of mammals ， 17 birds ， and 1 reptile (Table 2 and Appendix 2). Mammal species accounted for for 89.6% of all 鉱山nals identified in the pictures ， which included one species of Scandentia ， 1 Insectivora ，1 Pri mate ， 14 Camivora ，1 Pr oboscidae ，4 Art iodactyla ， and 8 Rodentia. Several Several chiropteran species were also photographed in 9 picωres ， but these could not be identified identified to species. Of 出e taxa 血at were identified to species ， the yellow 吋油 rat (Maxomys surifer) was the the most f詑quently photographed (3 ， 318 picωres) ， followed by 白 e pig-tailed macaque (Macaca nemestrina; 1，665) 叩 d 血e In dochinese ground squirrel (Menetes berdmorei; 1，073). 百le pictures with these 3 species accounted for 55.6% (6 ，056 pictures) of all 白e pict 町 es. Conversely ， 12 species of mammals ， one species of reptile ， and 6 species of birds were were photographed less than 10 times. Carnivora tended to be photographed inf 同quently ， and and 9 of 14 Carnivora species were photographed less 出an 10 times. After After omitting pictures of 出e same species 血at were taken a負.er an interval of less than than 30 min ，4 ，819 of 10 ， 955 pictures (44.0%) were regarded as representing a new visit for for a given species and used for further analyses.
Seasonal Seasonal Change of Activity Patterns
百le animal species for which 40 or more visits were recorded 泊 bo 血 the rainy and dry dry se 部 ons 釘 e Ii sted in Table 1 also (see Table 2 for the activity pattems). 百lere were significant significant di 首erences 泊 the temporal distribution of visits between the ra 泊y and dry seasons seasons for the Malayan porcup 泊e (Hystrix brachyura) ， the Siamese frreback (Lophura diardi) diardi) and the red jungle fowl (Gallus gallus) (Kolmogorov ・.Smirnov two-sample test ， P < 0.05; Table 1， Fig. 2). In In the rainy season ， H. brachyura exhibited unimodal activity pattems with a plateau at at between 2100 and 0200 h. However ， in the dry season ，activity 泊出is species was bimodal bimodal with peaks just after dusk 叩 d just before dawn (Fig. 2a). Lophura diardi exhibited almost almost entirely timodal activity in both the rainy and dry seasons ， but the moming peak was slightly greater th 佃 the aftemoon peak in the rainy season ， but smaller 泊血e 合y season season (Fig. 2b). Gallus gallus also exhibited bimodal activity with dist 泊ct peaks 泊血e early early moming and the late aftemoon 泊 the rainy season ， whereas the morning peak was 182 182 SHUNSUKE SUZUKI ET AL
Tabl 巴1. The results of Kolmogorov-Smirnov two-sample test for the sp 巴Cl 巴s for which 40 visits or more w 巴re r巴cord 巴d in both the rainy and dry seasons . Total ， the total total number of visits; Rainy ， the number of visits in the rainy season; Dry ，th 巴 number of visits in the dry season; ns ， not significant; へpく0.05;
Species Species name English name Total Rainy Dry D P Macaca Macaca nemestrina Pig-tailed macaque 350 233 11 7 0.085
Muntiacus Muntiacus m l/ nりak Barking deel 259 135 124 0.078 ns Cervus Cervus unico/OI Sambar deer 1 39 57 82 0.182 ns Menetes Menetes berdmorei lndochine se ground squirr 巴l 527 396 131 0.049 ns I Niviventer Niviventer fit/vescens Chest nu t rat 194 60 1 34 0.139 ns Maxomys Maxomys SUI ザer Ye l1 0w rajah rat 2081 1 500 58 1 0.022 ns ystrix Hystrix brachyura MaJayan porcupine 216 1 52 64 0.291
ふ3 Loph l/ ra diardi Siamese fireback 2 19 106 113 0.198 Ga l/ us ga//us Red jungl efow l 1 12 67 45 0 .274 司
a. a. Hystrix brachyura b.Lo phura diar ιfi c. Gallus g. 且llus (D=0.291 ，p nU atA。 30 唱' 民 U 晶 20 司 守 内 A eUKU 晶 10 。 。 12 18 0 6 。 6 12 18 。 6 12 18 Ti me ofday Fig Ul巴 2. Activity patl 巴rns of th 巴animals that 巴xhibited sign ifi cant seaso nalc hang 巴 8in b巴hav ior. Ab8c issa ，time of of day; ordinate ，the rate (% ) of visits for eac h hou r. Top ，rainy 8巴a80 n，fro l11 Apr il to Octob 巴r; bo 目OIn ， dry dry seaso n，fTom トiovember 10 Marc h. Open co lumll ，c1 ayt im 巴; gray col umn ，c repu scu lar time; c1 08ecl co lumll ，Il ighttime FORAGING ACTIVITY PATIERNS OF FOREST ANIMALS 183 obscured 泊白e dry season (Fig. 2c). In In the other 5 species ，activity pattems were not significan t1 y different between rainy and and dry seasons (Kolmogorov-Smimov two-sample test ，P > 0.05). Th us ， for these species ， 出e data for the duration of the study period were combined for further analyses. Diel Diel Activity Pattern of Each Species 百le 18 species for which 30 or more visits were recorded 訂 e classified into 血ree categories categories (Table 2; see Appendix 2 the for data of the species for which less than 30 visits were were recorded). Four mammal and all of the 6 bird species are regarded as diumal. 百lese diumal diumal species may be further classified into two subcategories; one (D- l) conta 加ing species species that had almost unimodal activities with either a distinct or a vague peak around noon or aftemoon ， and the other (I ト2) containing species that were primarily bimodally active active with distinct peaks in the moming and late aftemoon. Th e former subcategory ， D- l ，contained 2 mammals (M. nemestrina and Callosciurusfinlaysonii) and 3 bird species (Chalcophaps (Chalcophaps indica ，Pitta cyanea and Garrulax /euc%phus ，Fig. 3). Approximately 90% of of 由e members 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equently around noon whereas whereas C.fin/aysonii (Fig. 3b) and two bird species (P. cyanea (Fig. 3f) and G. /euc%phus (Fig. (Fig. 3e)) were photographed more frequently in the aftemoon. 百le latter subcategory ， D ー2 ，contained 2 mammals (Tupai ・a be/angeri (Fig. 4a) and M. berdmorei (Fig. 4b)) and 3 bird species (L. diardi for both seasons (Fig. 2b) ， G. gallus for both seasons (Fig. 2c) ， and and Arborophila ch/oropus 4c)). (Fig. This bimodal activity pattem was particularly conspicuous conspicuous in the two mammal species ， and more than 50% of their visits were recorded in 出 e crepuscular time (Table 2). Five Five rodent species (Rattus spp. ，Niviventer fu/vescens ， Leopoldamys sabanus ，Maxomys surifer ， and H. brachyura for both the rainy and dry seasons) were regarded as noctumal (Figs. (Figs. 2a ， and 5) ， with more than 90% of the visits for these species being recorded in the nighttime. 百四e rat species (Rattus spp. ， N. fu/vescens and M. sur 砕r) exhibited peaks in visiting visiting frequency during the first hour of nighttime between 1900 and 2000 h. 官官印 artiodactyl species were recorded during both day and night (Fig. 6). Tragu/us javanicus javanicus (Fig. 6a) and Muntiacus muntjak (Fig. 6c) were active mainly during day with peaks peaks around the dawn and dusk ， whereas Cervus unic%r (Fig. 6b) was active mainly at night night although it was also photographed during the day. In In all species for which 30 visits or more were recorded ， the temporal distributions of of visits were significan t1 y different from the pattem expected from the null hypothesis (Table (Table 2 ，Chi-square test ，P < 0.05). DISCUSSION Comparison with Previous Studies Th ere have been relatively few studies that have investigated the activity pattems of multiple multiple animal species coexisting in tropical forests (VAN SCHAIK & GRIFFITHS 1996; MIURA Ef AL. 1997). VAN SCHAIK & GRIFFITHS (1996) studied the activity pattems of 31 同∞ Table Table 2. Activity pattems of the species for which 30 or more visits were recorded. Pictures ，total number of pictures; total visits ，total hF number of visi 臼; Dayt 加 e，number of visits recorded during the dayt 加 e; Crepuscular time; number of visits recorded during 出ec 問:p uscul 紅白ne; Nighttime ，number of visits re 氾orded during nighttime. Percentages of the total number of visits are shown in the parentheses. Abbreviations for the categories of activity pattems 訂 'e as follows: 0- 1， diumal with unimodal distribution; 凶， diumal with b加 odal distrib 凶 on; N ， noctumal; 0 ，others. 紳げ< 0.001 for x? 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!O ry 岡田ー D81D e visi 包 time S悶 dentia Tupaudae Tupaia belangeri No 揃lem ttee shrew 190 121 54 (45) 釘 (55) 。(的 75 .7 *** 0- 2 防白 lates ce 問。 pitheci 伽E Macaca nemeslrina Pi g- 凶100 macaque 1665 350 320 (91) 27 (8) 。(的 224 .3 *** 0- 1 Arti叫 actyla Tragulidae Tragulidae 1干'agulus javanic 山 Le sser Malay mouse -d eer 177 96 51 σ3) 42 (斜) 3 (3) 44.8 。 *** *** 印国 dZ Ce rvi 也s .Ce 四郎 unicolor Sambar 321 139 7 (5) 25 (1 8) 107 (77) 54.6 *** 。 臼己目白 Muntiacus Muntiacus m醐 tjak B創魁 ng d民r 938 259 142 σ5) 63 (24) 54 (21) 26 .3 *'験* 。 Rodentia 伽 /o Fmlayson's squin 官l '3) 7 (η 。 *事* Sciuri a Cal sciurus fi nJay sonii 305 106 99 。 制) 69.8 0- 1 CNdR *ホ傘 Menetes Menetes berm 即 rei Ind ochin 凶 eground sq' 瓜rrel 1073 527 zω (39) 318 (61) 。。) 350.4 0- 2 Muri 也E Rat 簡明・ 85 63 。(的 3 (5) ω (95) 43. 6 *** N 回目 Niviventer Niviventer fulvescens α儲回 ut rat 242 194 。{め 17 (9) 177 (91) 12 1. 5 *** N .h Le opol ，伽I)' ss伽，nus Noisy rat 120 73 。(め 。(的 73 (1 00) ω.1 **事 N ・ Maxomys Maxomys surifer Yellow raj 油 rat 3318 2081 。(め 134 (。 1947 (94) 1375 .7 *** N Dゆdi 也s Hystrix brachyura (R ainy) Malay: 叩 pO lC up Is e 772 152 。仰) 8 (5) 144 (95) 103 .7 *事* N H. H. brac 付附仰y) 329 64 。{買} 7 (11) 57 (89) 38.4 *** N B凶s iformes GaJl iformes ph 悩ian 拍車 Lo phura 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 Galll ωgallus 侭細川 ROO junglefowl 108 m 50 (7 5) 17 。5) 。 (0) 35 .4 *** 0- 2 G.galll ω肋y) 101 45 27 (60) 18 (相) 。 (0) 24.6 *** 0- 2 Arborophila Arborophila chloropus Scaly-h 毘astω 抑rtridge 62 40 27 (68) 13 (33) 。(的 21. 2 *** 0- 2 Columbiformes Columbiformes 印刷bi 加 C加Icophaps indica Emerald dove 130 100 97 (97) 3 (3) 。 (0) 73. 2 *** 0- 1 Pas 町 iformes Pi 凶伽 e Pitta cyanea Blue pitta 42 31 27 (87) 4 (1 3) 。 (0) 18.0 *** 0- 1 Musci 悶pid 国e Garrulax leucolophus Wh ite-c res 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. a. MSCBCS nemestnns 、Callosciurus nnJ Sysonii 20 20 (N=350) 20 15 15 15 10 10 10 。5 。5 。 6 12 18 。 6 12 18 *- 巳. Ch sJ cophsps indics 20 20 (N =100) 16 仏1=31) 16 15 15 12 12 10 10 8 8 。5 。4 。4 。 6 12 18 。 6 12 18 。 6 12 18 'l'im e ofday Figur 巴 3. Act iv ity pattern s of the anim als c1 ass ifi 巴d as diurn al sp巴cies w ith an almo st unimoda l di strib ut ion. S ee Fig. Fig. 2 for further explanations. a. a. Tu paia belangeri b. Menetes berdmorei c. Ar borophila chloropus 40 可 (N=121) 30 可 (N=527) 20 、(N =40) 30 30 15 20 20 求 20 10 10 10 10 10 5 。 。 0 。 6 12 18 。 6 12 18 。 6 12 18 Ti meofday Figure Figure 4. Activity patt 巴rn s of th 巴 anima ls c1 assified as diurna l spec ies with a bimod aJ distribution. S巴巴 Fig.2 for for further ex pl anations 186 186 S H UNSUKE S UZU KJ ET AL a. a. Ra ttus 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. Le opoldamys sabanus d. Max om ys S Ul 鵠 r 20 可 (N =7 3) 20 可 (N=2081) 15 15 10 10 。5 。5 12 18 。 6 12 18 。 6 Time ofday FigUl 巴 5. Act ivit y pattern s of th巴 anim als class ifi ed as noct urnal sp 巴C I巴s. See F ig. 2 for further exp lanati ons a. a. Tr sl{U lusjavanicu8 b. Cervus unicolor c. Muntiacu8 munijak 20 、(N =9ω 15 15 、(N =259) 15 15 10 10 10 10 10 ~ 5 5 5 。 。 。 。 6 12 18 。 6 12 18 。 6 12 18 τ'im e of day Fi gur e 6. Act iv ity pattern s of th 巴 animal s cla ss ifi ed as be ing act i ve dur i. ng both the day and ni g ht. S巴巴 Fig .2 for for furth 巴r exp lanat i ons FORAGING ACTIVITY PATIERNS OF FOREST ANIMALS 187 animals animals in In d'O nesian rain f 'O rests using aut 'O matic camera systems. 官ley c1 assified the ph 'O t'O graphed animals int 'O the f'O ll 'O wing categ 'O ries: diurnal ，n 'O cturnal and cathemeral (出 e last last categ 'O ry had been defined by T A'πERSALL (1 987) f'O r primates as being active during b'O th day and night). In additi 'O n，出ey sh 'O wed that arb 'O real species were either strictly diurnal diurnal 'O rn 'O cturnal ，whereas te 町'es 凶al species were primarily cathemera l. H 'O wever ，their results results d 'O n'O t appear s 'O c'O nvincing because their sample sizes were relatively l'O w (1， 308 pictures 加 t'O tal) and m 'O st species (20 'O f 31 species) were ph 'O t'O gr 叩 hed less 血an 20 times. Furtherm 'O re ， the time at which the pictures were actually taken did n'O t appear t 'O be rec 'O rded 'O n their pictures by their camera systems. C 'O nsequently ， the activity patterns described described by VAN SCHAIK & G 即開口百S (1 996) d 'O n'O t appe 紅白 be s 'O accurate. By using aut 'O matic camera sy 蜘 ms ，M 即 RA EJ AL. (1997) sh 'O wed the activity pa 悦 rns 'O f animals in a tr 'O pical rain f 'O rest 'O f the Malay Peninsula by taking 2，738 picωres 'O n which the time 'O f the animal visiting was printed. They described the activity patterns 'O f representative 8 representative mammal species ，3 'O f which were n'O cturnal ，3 were diurnal 佃 d2 were active active irrespective 'O f day and nigh t. In血 e present sωdy ，we t'O'O ka c 'O nsiderable number 'O f pictures (in excess 'O f 10 ，000). In In additi 'O n ，we c'O nsidered 'O nly th 'O se pictures taken 30 minutes 'O rm 'O re after 出 e previ 'O us picture picture as c'O nstitut 泊 g 佃 independent visi t. In d'O ing S'O， we believe 'O ur results t 'O be m 'O re reliable 出 an th 'O se 'O f previ 'O us studies. Based 'O n the standard time ar 'O und KY ， we divided a day 泊 t'O the f'O ll 'O wing three p制 s; dayt 泊施， nightt 泊 le ，and crepuscul 釘 time. Whi le the defmiti 'O n 'O f crepuscular time may n'O t be precise given 出 at the timing 'O f sunset and sunrise changed seas 'O nally (alth 'O ugh the the change was remained less 血an 'O ne h'O ur) ， the activity patterns 'O f the 仕equently ph 'O t'O graphed animal species c'O uld be classified with 'O ut difficulty (Figs. 2 -6; Table 2). Bel 'O w we discuss the activity patterns 'O f species bel 'O ng 加gt 'O each categ 'O ry. Diurnal Diurnal species 百le 2 mammal species and the 3 bird species were c1 assified int 'O the subcateg 'O ry 'O f 0- 1 (diurnal with a peak 'O r peaks ar 'O und n'O'O n 'O r aftern 'O'O n ， Fig. 3). Of these ， the 2 mammals share a rather arb 'O real tendency. Furtherm 'O re ， in the 3 birds ，all 'O f which bel 'O ng t 'O C 'O lumbif 'O rmes 'O r Passerif 'O rmes ， the tendency t 'O stay 'O n the gr 'O und appears t 'O be weak when c'O mpared t 'O the birds in subcateg 'O ry D ー2 ，all 'O f which bel 'O ng t 'O Phasianidae (see bel 'O w). 官le 2 mammals and the 3 phasianids c1 assified as D ー2 (diurnal with peaks in the early m 'O rning and late aftern 'O'O n) share a tendency 'O f living mainly 'O n the gr 'O und. In a 仕opical rain rain f'O rest 'O f Singap 'O re ，many invertebrates were 'O bserved t 'O m 'O ve fr 'O m the shelter 'O f 白e f 'O rest f1'O'O rt 'O出 e can 'O py t 'O feed at 訂'O unddusk 叩 dt 'O reωm 釘 ound dawn ，when predat 'O rs such 副総O中 i'O ns 合ogs and spiders ambushed and preyed 'O n these invertebrates at the base 'O f 仕ees (W HπMORE ， 1990). Tree shrews and gr 'O und squirrels are als 'O kn 'O wn t 'O 合'equently prey prey up 'O n invertebrates as well as fruits (HARRISON ， 1961; LEKAGUL & McNE 乱 Y 1977; MACK 悶 NON ， 1978; CORBET & HILL ， 1992; MACK 町 NON EJ AL. ， 1996; EMMONS ， 2000). 官le birds 'O f the Phasianidae als 'O appe ぽ t'O eat inve 巾 brates. C 'O nsequently ，it is hyp 'O thesized that that the bim 'O dal activity pattem 'O f 白e D-2 species in KY may be related t 'O a habit 'O f preying preying 'O n invertebrates 'O n the f 'O rest f1'O'O r. 188 188 SHUNSUKE SUZUKI Ef AL. Nocturnal Nocturnal species In In the present study ，all 4 of 白.e Muridae species and the M a1 ayan porcupine ， Hystrix brachyura ， were noctum a1 (Figs. 2a ，b， and 5; Table 2). In the 3 murld species ，Maxomys sur 砕r ，Niviventer fulvescens and Rattus spp. ， and the M a1 ayan porcupine ，activity started in in the late dusk after 1800 h and ceased in the early dawn before 0600 h，Leopoldamys sabanus ，however ，became active after 1900 h in the early at nighttime and ceased before 0500 h. PEI PEI (1995) studied the activity pattem of the spinous coun 町 rat ，Niviventer coxingi ， using using an automatic camera system in Taiwan and suggested 白紙血e onset and cessation of of its activities were controlled by rapid changes in light intensity accompany 泊g sunset and sunrise. Simil 釘 regul 瓜ion of activity was reported by CARLEY ET AL. (1 970) for the deer deer mouse ， Peromyscus maniculatus ，加 Kansas ，US A.百 le onset and cessation of activities of of the noctumal species observed in the present study may a1 so be controlled by light intensity. intensity. If so ， the present results suggest that 血 e light intensities under which L. sabanus becomes active may be lower than those for the other 4 noctum a1 species. Other species 百le 3 species of Arti odactyla in KY ， Tragulus javanicus ， Cervus unicolor ，and Muntiacus Muntiacus muntjak ，were active during both day and night (Fig. 6); these species may be regarded regarded as cathemer a1 as defined by T AT 四 RSALL (1 987). 百lese results agree with those of of previous studies conducted in Southeast Asia (VAN SCHAIK & G 悶 WπHS 1996; M 町 RA ET AL. ， 1997; MCCULLOUGH ET AL. ， 2000). MATSUBAYAS 田町AL. ， (2003) reported that 白e moused 田 r， T. javanicus ，was observed to wander or forage on the forest floor only during the the daylight periods (0500 ー1900 h) ，and 出at it moved between foraging and resting sites at at dawn and dusk 加出e Kabili ・Sepilok Forest Reserve in Sab 油， Bomeo. Mru RA ET AL. (1997) (1997) a1 so reported that T. javanicus was active irrespective of day and night in Pasoh ， 血eM a1 ay Peninsula. Conversely ，SRIKOSAMATARA & HANSEL (1996) regarded T.javanicus in in KY as being prim 紅 ily noctumal (a1 though it w 出a1 so active during the daytime ， particularly particularly in the dry season). In血 e present study ，however ， T. javanicus was recorded mainly mainly during the day (0600 ー1800 h) ， with peaks in the moming and evening (Fig. 6c). 百lU S，出 e present results are in agreement with those for other reserves of Pasoh (M 町 RA ET AL. ， 1997) and Sabah (MATSUBAYAS 田町AL. ， 2003). In In the present sωdy ， as well as 白at ofM 即 RA ET AL. (1 997) ，合uit was used as bai t. Therefore ，most species observed in 出e present study were considered to be either frugivorous frugivorous or omnivorous anim a1 s (although several carnivores were a1 so inftequently recorded). recorded). Furthermore ， the activities recorded 泊出e present study were considered to be directed directed prim 釘 ily at foraging. is It 泊lpOrtant to note ぬat 血e present results may be limited by the that fact activities other than foraging may have been Further missed. studies using other other methods will therefore be required in order to assess the activity pattems of each species species more comprehensively. Seasonal Seasonal Changes in Activity Patterns Th ere were significant differences in the activity pattems between the rainy and dry seasons seasons for the 3 species (the porcupine ，Hystrix brachyura ， and the 2 birds Lophura diardi and Gallus gallus) of the 9 species investigated conceming seasonal change (Table 1. and FORAGING ACTIVITY PATTERNS OF FOREST ANIMALS 189 Fig. Fig. 2). In all 3 species ， the activity pattems of bo 出 the rainy and dry seasons were classified classified into the same category (Table 2). However ，泊 H. brachyura the temporal distribution distribution of visits was markedly different between the rainy and dry seasons; a plateau was observed around midnight in the ra 回y season whereas peaks just after dusk 佃 d just prior prior to dawn were seen in 血.e dry season (Fig. 2a). At present ，we have no explanation for for these changes in activity pattem between the rainy and dry seasons. In In both of the bird species (L. diardi and G. gallus) ，moming activity appe 紅 'ed to be more vigorous in the rainy season (Figs. 2b and 2c). However ，it is also unclear whether 白is seasonal change in behavior is an adaptation to seasonal changes in some environmental factors. factors. We have described the diel foraging activity pattems of frugivorous and omnivorous animals animals on the forest floor and described seasonal changes in the foraging behavior of several several species for the frrst time in a tropical seasonal fores t.百 le adaptive significance of the the seasonal changes remains unclear; 出ey may be a丘町 ted by physiological cons 釘'aints 釦d/ or environmental factors such as the availability of food items. Moreover ，al 由ough fruit fruit was used as bait in the present study ，some carnivores such as the leopard cat (Prionailurus (Prionailurus bengalensis) 叩 d dhole (Cuon alpinus) were infrequen t1 y recorded (Appendix 2). 2). It remains unclear whether they came to take 仕uits or to hunt the frugivorous or omnivorous omnivorous animals that visited the baited sites. However ，their activity may influence 血e time time when the frugivores or omnivores 訂 'e active. Further studies on 血e ecology and behavior behavior of each species ，including the camivores ， are required in order to understand the variety variety of factors that determine the changes in diel and seasonal activity p副 ems of forest animals. animals. ACKNOWLEDGMENTS We are grateful to the National Research Counc i1 of Th ailand and the National Park Park Di vision of the Royal Forest Department of Th ailand ， for granting us permission to conduct conduct studies on activity in KY. We 白ank B. Saengthong ， S. Chua i1 ua ， S. Nakkuntod ， S. S. Sanguanchat ，N. Jirawatkavi and all the staff of Thailand Hombill Pr oject for supporting our our field work as well as their kind encouragement and hospitality. We extend our hearty 出釘lks to the staff at KY. 百lI s research was supported 泊 part by a Research Fund of the Japan Japan Society for the Pr omotion of Science (#1357006) and JSPS Research Fellowships for for Young Scientists for S. Kitamura. REFERENCES AKABAR ，Z. ，AND M. L. 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Fallen fruits 釦 d terrestri a1 vertebrate 合ugivores: a case study in lowland 位。 pical rainforest in Peninsular Malaysia ， Pages 151-174 in P. M. Forget ， P. M. Lambert ，J. E. Hulme ，and S. B. Vander Wall (eds.) ，Seed Fate: Predation ，Dispersal and Seedling Seedling Establis 伽， ent. CABI Publishing ，W a1 lingfo 吋， UK. 192 192 SHUNSUKE SUZU 悶 EIAL. Appendix 1. Li st of plant species used as bait and the oumber of days wheo camera- 回 ps operated operated 00 each plant species. Family Family Species Name Trap Family Species Name Tr ap days days days An ac 温rdiace 悦 Choerospondias axillaris 295 Le guminosae Acacia sp.S K3 98 127 Ann onaceae Alphonsea sp.SK095 35 Acrocarpus Jr， 似 'i nifo /i us 23 Dasymaschalon Dasymaschalon s，ω'tepense 26 Entada rheedii 26 Desmo 哩，chinensis 4 Magnoliaceae Michelia baillonii 18 M i/i， 凶 'a cunea ω 27 Meliaceae Aglaia lawii 44 P同'm; 抑制'croca 抑 136 Aglaia spectabilis 186 Polyalthia Polyalthia sp.SK115 19 Aphanamixis polystachya 12 Polyalthia Polyalthia sp.SK195 37 Dysoxylum cyrtobotryum 37 Polyalthia Polyalthia sp. 24 Me /i a azedarach 103 Uvaria Uvaria lurida 12 Sandoricum koetjape 91 Burseraceae Burseraceae Canarium euphyU 聞 189 Moraceae Antiaris toxicaria 50 Cel ぉ岡郎総 Bhesa robusta 32 Artocarpus lacucha 31 Comaceae Comaceae Mastixia pentandra 103 Ficus altissimo 136 Cucurbitaceae Cucurbitaceae Trichosanthes tricuspidata 68 Ficus annulata 39 Dipterocarpaceae Dipterocarpaceae D伊terocarpus graci /i s 53 Ficus kurzii 27 Ebenaceae Ebenaceae Diospyros glandulosa 21 Ficus subcordata 119 Elaea 伊aceae 1: laeagnus ωゆrta 90 Ficus sp.2 2 ElaωC 釘paαae Elaeocarpus robustus 104 Ficus sp .3 12 Eupho 巾lia αae， Balakata baccata 65 Myricac 回 E Knema elegans 18 Macaranga Macaranga siamensis 36 Syzygium al 岬町醐 52 Euphorbiaceae Euphorbiaceae sp.S K25 6 14 Cleistocalyx nervos ，醐 26 Fagaceae Fagaceae Lithocarpus sp. 41 Palmae Areca triandra 69 Li thocarp ωthomsonii 23 Li vistona jenkinsiana 25 Quercus Quercus myrsina φ，/i a 125 Pro teaceae He /i cia formosana 20 Fl aco 町 tiaceae Casearia grl 仰 iifo /i a 58 Rubiaceae Anthocephal ω，chinensis 133 Casearia Casearia sp. 10 Nauclea sp.SK033 85 Gn e旬ceae Gnetum sp.SK024 39 Canthium coffeoides 47 Guttiferae Guttiferae Garcinia sp.SK442 17 Rutaceae Acronychia pedunculata 14 Icacinaceae Icacinaceae Gonocaryum lobbian 附 1 34 Citrus /i mon 7 Platea Platea latifolia 60 Citrus sp. 62 La urace 唱E Beilschmiedia sp.SK437 23 Clausena harmandiana 43 Beilschmiedia 脚 ingayi 5 Sapindaceae Mischocarpus pentapetalus 31 Cinnamomum Cinnamomum subavenium 106 Sapotaceae Pouteria stellibacca 73 Cryptoca η 'a sp.SK424 54 U1 maceae Aphananthe cuspidata 59 Phoebe Phoebe cathia 24 Appendix 2. Species for which less than 30 visits were recorded. See Table 2 for further explanation. Family Family Species English Pi c佃res To 旬l Day time Crepuscular Night time 目ame name visits time Insectivora Insectivora Crocidurinae Crocidura horsfieldi Horsfield' s shrew 。 (0) 。 ρ) (100) HOm同 αliroptera αliroptera Bat 9 9 2 (22) 。 仰) 7 (78) 〉。同 C紅nivora Canidae Cuon alpinus Dhole 32 6 5 (83) (1 7) 。 (0) 12 12 (14) 。 (0) 6 (86) Z Ursidae Ursidae Ursus thibetanus Asiatic black bear 7 。〉の叶同〈『 U. U. malayanus Sun bear 2 2 (50) (50) 。 (0) Mustelidae Mustelidae Martes j1 avigula Yellow-throated m 釘ten 2 2 2 (100) 。 (0) 。 (0) Arctonyx Arctonyx collaris Hog-badger 2 2 (50) 。 (0) (50) HJ 「同 Viverra Viverra zibetha Large Indian civet 24 17 。 (0) 4 (24) 13 (76) 〉寸吋開M V. V. megaspila L紅 ge spotted civet 。 (0) 。 (0) (100) Viverricula Viverricula indica Small indian civet 9 5 。 (0) (20) 4 (80) mZ (0) (0) (82) 臼 Paradoxurus Paradoxurus hermaphroditus Common paI m civet 53 28 。 5 (1 8) 23 O Herpestidae Herpestidae H erpestes javanicus Small asian mongoose 6 4 4 (100) 。 (0) 。 (0) 明司 Felidae Felidae Prionailurus bengalensis Le opard cat 35 19 2 (11) 6 (32) 11 (58) om 明白↓〉 Catopuma Catopuma temminckii Golden cat 2 2 。 (0) (50) (50) Pard ゆ， lis nebulosa Clouded leopard 2 。 (0) 。 (0) (1 ∞) Z P. P. marmorata Marbled cat 5 2 2 (1 00) 。 (0) 。 (0) 同室〉 断。 boscidea Elephantid 島E Elephas maximus Asian elephant 41 9 3 (33) 5 (56) (11) F 臼 iodactyla Art iodactyla Suidae Sus scrofa Wild boar 36 24 5 (21) 6 (25) 13 σ4) Rodentia Rodentia Di podidae Atherurus marourus Asiatic brush-tailed porcupine 2 。 (め 。 (0) (1 00) Reptile Reptile V鉱加idae Varanus salvator Monitor lizard (1 ∞) 。 (0) 。 (0) B凶 Phasianidae Lo phura nyc 幼emera Silver phe 悩組t 21 10 10 (1 00) 。 (0) 。 (0) liformes GaI liformes 目。凶 Cuculiformes Cuculiformes Cuculidae Carpococcyx renauldi Co raI -billed ground-cuckoo 29 24 21 (88) 3 (1 3) 。 (0) 戸市 Species Species for which less 出an 30 visits were recorded. See Table 2 for further expl 組 ation. (Cont.) wh Appendix 2. 骨 Family Family Species English Pi cturω To 旬I Day time Crepuscular Night time name name name 吋sits time Coraciifonnes Coraciifonnes Bucerotidae Anthracoceros albirostris Oriental pied hornbill 100 22 22 (100) 。 (め 。 (め Passerifonnes Passerifonnes Pittidae Pitta phayrei Eared pit 飽 (1 ∞) 。 (0) 。 (め 。 (0) 。 (0) cnonotidae Py cnonotidae Criniger pallidl ω Puff~即日ated bulbul 29 19 19 (1 ω) Muscicapidae Muscicapidae Garrulax chinensis Black-throated laugh 泊凶 uush 2 2 2 (1 00) 。 何) 。 切) cinia Lω cinia cyane Siberian blue robin 6 3 3 (100) 。 切) 。 。) Copvchus Copvchus malabaricus Whi te-rumped shama (100) 。 。 (0) 切) 回国 Monticola Monticola solitarius Blue rock~世uush (100) 。 切) 。 切) CZ 師巴阿国的 othera Zo othera citrina or 加1ge-b 伺 ded thrush 17 II II (100) 。 。) 。 何) Cyomis Cyomis sp. Blue flycatcher (1 00) 。 切) 。 仰) CNC 間同同『』』 ・F