Will Swim for Food: Resource distribution and feeding behavior in Long-tailed Macaques (Macaca fascicularis)

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Wi Swi Fd: Rc dititi ad dig havi i Lg-taid Macaq (Macaca acicai) Ji Bavid, Nic Cat, Daia D, Ryi Sachz, Nah Thapa, Cha Wad Sittd Jy 8, 7

Atact Th dig havi i- agig g-taid acaq wa xaid at a zgit pak. Th acaq w pviid with ccia dit, it ad pat daiy at pdtid cati (th) ad ti (at at twic a day). Ovati at th ti wh d wa dititd cid that th w th paat gp acaq which d typicay at pciic cati. Ctt cptiti tw adt ad jvi wa vd at a dig it, ti ad a d typ. Th at cpt ack dig cptiti tw adt a ad adt a wa attitd t th hity th acaq which ha td i a highy id ppati wh adt a ad a a atd. Th ct dig aagt th pviid acaq appa adqat t t thi titia d. Ky wd: Macaq, d-c, ctt cptiti. Itdcti Th ajity piat iv i cia gp. Whi th advatag gp ivig i t ctai, cptiti d i a ay avida cqc gp ivig piat. It i gay accptd that th a tw typ dig cptiti, ca ad ctt. Fdig cptiti cc th withi ad tw gp. Withi gp ca cptiti cc wh aia qay ha a d ppy, ad ctt cptiti cc wh diat aia piz acc t d that thy tai tha th th th gp (Ja ad va Schaik, 988). Sca cptiti i qty vd i pci wh th d c i dipd (.g. iv piat) ad ctt cptiti wh th d c i cpd (.g. i giv piat) (Saj ad Sictt, 7). I thi tdy w xaid th c dititi ad dig havi i gtaid acaq (Macaca acicai), a pci that iv i ag cpx cia gp. Adt a, adt a, iat, ad jvi iv i gp that ca 8 t 5 idivida (d Rit, 99). Ma hw a tg diac hiachy i which thy ctt high ak wha a ak i dpdt th ak thi th. W pdictd that i a pci i which diac i cay appat w wd v ctt cptiti v d. Th acaq w tdid iv i a ataitic c ad a pviid at dit cati ad ti a daiy ai. Th acaq did t appa t chiv gp (Kcza, 998), ad th th piiity xitd cptiti tw dit gp withi th c. Additiay th acaq w pviid with dit qaity d it at va it ad giv th dit dig gi, th pttia xitd ca cptiti t cc. W w a ittd a captiv aagt ppctiv t dti whth th acaq w civig icit d.

Matia & Mthd Sjct Mky Jg i Miai, Fida wa dd i 9 y Jph DMd, wh catd th haitat that h cd tdy piat havi ad cia tct i a cvit ttig. H it itdcd ix g-taid acaq th iad Java, Idia t hi c, ad a th ky ivig th tday a dcdat th igia ix. Sic 96, Mky Jg ha p t th pic ad ha c a ppa attacti i pat ca th viit a awd t d th ky, ad cqty th ky hav c w haitatd t pp. Idtiicati (Ag, Sx, Ca, Idivida, Kp, ad Gp) Sic th a ia ith-dat th ajity th acaq w cd y caiy th it ad ag/x ca: adt a, adt a ad jvi. W ditigihd th ad thi piay ad cday xa chaactitic ad iz. W pt th day at th giig th tdy at Mky Jg aig t ditigih tw th ca. W w a a t a t ditigih ctai ky idivida. With cida aitac th kp (Tacy Cak ad Tia Caqi i patica), y th d th tdy w w a t idtiy tw ad ky. Th g-taid acaq at Mky Jg a ptd t dividd it th cia gp; tw th gp (Lki ad Capa ) tak thi a thi ad (apha-a). Th thid t ack a ca ad ad i d t a th Ap gp (Cak, Pa Cicati 7). Thght tdy pid, w a kpt tack ay igiicat chag i th gp dyaic th acaq ad ay ai t th a. Haitat Th acaq c i a tpica hadwd hack, t th haitat ha cad, ad th pat th c a vgw y ivaiv xtic pci. Th ky a hd i a.6 hcta cpd that iic thi ata vit ad i dd y a th t chai ik c that i ctiid t ctai th ky. Fcd wakway aw viit t wak thgh th c. A pit at th acaq c ad th cati a kp ptd hw i th Wid Mky Swiig P, which i a h t va pci xtic ih. Fdig Th acaq ag atay it, vgtati, ad ict. I additi, th acaq a pviid y th kp ad viit. Th kp d th acaq at pciic cati ad ti. O cati i th Wid Mky Swiig P. At pdtid ti (, 5, ad 6), th a hw hd at which th

kp ditit appxiaty.6 kg ixd it ad had id gg which th kp thw it th wat. Pat (8 g) ad cciay ppad Mky Bicit (5 g) a dititd at cati. Th it cati i adjact t Kig Iad, h th pak ivack gia; th cd cati i adjact t th kitch. (S Pat.) Th dig tak pac at 9 ad, pctivy. Dpit th act that th a hpp (dig ctai) attachd t th c, kp typicay catt th ix pat ad icit v th c. Tw th dig hpp th pit th acaq c a id vy th day with a iia ix pat ad icit. Ath iptat c d th acaq a th t a cp which hag y chai th th wakway ad ca id y viit with did it which ca pchad i th git hp. Th acaq ca p p th chai t acc th did it. Macaq a cavg it ad icit i a aa which hd va cag va adt a acaq that hav vd th c patd icidt aggi. Data Ccti Data w cctd th idtity th gp, th acaq pt, ad th acaq dig at ach cati at ctd dig ti. I additi w a cctd data xpita aipati thi dig digd t diy dig cptiti ad dti th t highy vad d it. W pt t ti vig th havi th acaq 5 t it ti a day v a pid wk. W cdd hw ay ky w at ach dig tati dig dig. Sti w aw th typ havi ch a atig, cptiti d, ad aggi twad ach th. W dtid whth a ky wa dig y vig d ig pacd i th th ig aticati. O ach ta dividd it th dit gp cpid tw pp ach. Each gp wa digatd t v th ag/x ca: adt a, adt a, ad jvi. At it itva, ach pai v cad th gp ad cdd th idivida withi a pdtid day ad whth thy w dig (itata cdig). Bad th kw idtity ctai ky idivida, w dtid which th th gp wa pt. Additiay, p a cdd th tta ky a w a ay ta havi thy dipayd ch a pcivd chag tat, aggi, atig, vigiac, ad gp dipact (ad i cdig). Mthd data ccti w Mati ad Bat (99). O th ia day th tdy w diid th dig ti t that ky had pviid pi t th it p hw. W a ptd at a v cati ad ti th th d typ typicay d t th acaq (pat, it, ad ky icit) i qa qatiti y v, patiay paatd i th paa w aig appxiaty iv t y t.

Data Aayi W cacatd chi-qad va t dti whth th dit gp (Capa, Lki ad th Ap gp) hwd a pc th dit dig it. W pd a t-tt t dti whth th wa ctt ca cptiti withi ach gp (i.. whth th a, a jvi gt t at it). W a pd a t-tt t dti whth th ky hwd a pc it, pat, icit i d chic xpit. Rt Cptiti Btw Gp Th t hw that th th gp acaq d paaty at th dit dig cati. Kig Iad wa xcivy d y Capa gp, ad th kitch it wa xcivy d y th Ap gp. Athgh t ay vd, th taiig aa, wh th cagd a w hd, wa d xcivy y Lki gp. Athgh th gp gay d paaty, th xcpti wa Wid Mky Swiig P hw dig which Lki gp wa dipacd y Capa gp. Thi i ptd i th data y a ha a pit giv t ach gp that hw. Th Chiqa tt hwd a tatitica igiicac i which gp appad t qty at dig tati. Ta : N ach gp pc at a giv dig it Wid Mky Kig Iad Kitch Swiig P Capa 5 7.5 Lki.5 Ap 5 Chi- qa:.67 DF: P-va: <. Cptiti withi Gp Th patt dig havi ach th th gp acaq wa iia. Wh w aayzd th pctag ach ag/x ca that w vd dig wh cpad t th ach ag/x ca pt th it it it at th d wa dititd, w d that th wa dic tw th dig havi th adt a (ig a, a, a) ad th adt a (ig,, ) t th th adt a ad a d tha th jvi (ig c, c,

c). Th t a aizd i Ta. Th a ad tadad dviati a icdd i th appdix. Ta : T-tt t pctag adt v jvi dig. Cap Gp- Ap Gp-Kitch Cap Gp-Kig P Shw Iad Ma v. Jvi p<. p<. p<. Fa v. p<. p<. p<. Jvi Fig. a Ma a acq pt ad dig v ti at "P Shw" (Data tictd t 5 dig).5.5 a.5 # a # a dig.5 5 6 7 8 9 5 ti (it)

Fig. Ma a acq pt ad dig v ti at "P Shw" (Data tictd t 5 dig).5.5 a.5 # a # a dig.5 5 6 7 8 9 5 6 ti (it) Fig. c Ma jvi acq pt ad dig v ti at "P Shw" (Data tictd t 5 dig) 6 5 j v i # jvi # jvi' dig 5 6 7 8 9 5 6 Ti (Mit)

Fig. a Ma a acq pt ad dig v ti at "Kitch Sit" 5.5.5 a.5.5 # a # a dig.5 5 6 7 8 9 5 6 ti (it) Fig. Ma a acq pt ad dig v ti at "Kitch Sit".5 a.5.5 # a # a dig 5 6 7 8 9 5 6 ti (it)

Fig. c Ma jvi acq pt ad dig v ti at "Kitch Sit" 6 5 j v i # jvi # jvi dig 5 6 7 8 9 5 6 ti (it) Fig. a Ma a acq pt ad dig v ti at "Kig' iad".5.5 a.5 # a # a dig.5 5 6 7 8 9 5 ti (it)

Fig. Ma a acq pt ad dig v ti at "Kig' iad".5.5 a.5 # a # a dig.5 5 6 7 8 9 5 ti (it) Fig. c Ma jvi acq pt ad dig v ti at "Kig' iad" 7 6 5 j v i # jvi # jvi dig 5 6 7 8 9 5 ti (it)

Th t dtatd that th d ppy wa t qay had y a th idivida i a th gp. Th adt a ad a taid t th d ppy that th kp dititd, whi th jvi gt th pti. Thi i a xap ctt cptiti, a ppd t ca cptiti i which th d ppy i dititd ag th gp vy. Cptiti withi th gp ccd v th gg at th p hw. F th t hw i which data wa cctd, gg w thw it th wat ad 5 w at dig th vati pid. Thit gg w tivd y a acaq, wa tivd y a a, ad wa at y a ih (p<., tw-taid iia tt). Th idtity ach acaq that tivd a gg wa kw. Th a that tivd th gg w a high- akig w d ky with i tat. O th gg that w at dig th vati pid, Capa tivd ad cd % th, tha ay th idivida. Fd chic xpit Th t th d chic xpit a aizd i ig. Uig a tw-taid t-tt w d that th ky pd it v ky icit; thy a pd pat v ky icit. Hwv th wa igiicat dic tw th it ad th pat. Th p va a ptd i ta. Fig : Fd chic xpit 8 7 6 5 # ky Fit pat Bicit 5 6 7 8 9 5 6 7 ti (it)

Ta : Fd chic xpit t-tt t. Fit v. Bicit Pat v. Bicit Fit v. Pat P va <. <. >. Scia Bhavi ad Dgaphic Chag Ov th c tdy at at iv a acaq had ai. W vd tw a with w ai i Capa gp, tw i Lki gp, ad i th Ap gp. Wh th a wa a w akig ky, at th ith w aw h tak a pit piti dig dig ti. F xap, th it ay wa t a w akig a i Capa gp ad Lippy J, 7. B th ith, Lippy ay tayd twad th ack th gp dig dig ti. At th ith, h t tayd ight xt t th c wh t th d wa thw dig th dig at Kig Iad ad at with th high akig ky at th Wid Mky Swiig P dig. O th high akig a (Gadpa) v t h iih hi gg at th Wid Mky Swiig P J 9. O Jy, 7, Ggia, th hight akig a i Lki gp (Cak, Pa Cicati 7), had a ay. Pi t thi, Capa gp ay ca t th it tw p hw th day, ad Lki gp ay ca t th hw at i th at. At Ggia had th ay, Lki gp tatd cig t th it p hw th day, ad Capa gp did t hw p ti th 5 hw. Dig th tdy pid w aw th itac gp dipact. O Jy 6 at th hw at th Wid Mky Swiig P, Lki gp wa th th it iv it vati. At iv it, Capa gp vd it th vati aa th t id th p ad Lki gp vd t th ight id ad t th aa. O Jy, at th d th p hw t which Lki gp had c, ight at w tppd iciay cdig data Capa gp ca i ad chad Lki gp. Capa acd Lki tp a wakway, ad Lki a. Tiki (high akig a i Capa gp) ad Mty ( apha a Capa gp) chad th t Lki gp th aa. At thi ctati, Capa gp t a w, ad y Hidi, a w akig a with a w ay Lki gp, tayd t at th it t i th p. Th thid icidt dipact ccd dig d chic xpit Jy. Iitiay, Lki gp hwd p t th dig aa. At iv it, Capa ad Tiki hwd p ad tw thid Lki gp t th aa. Tiki dipacd Lki, ad Ggia t cayig a pic. Lki tayd i th aa, ad Capa thatd hi. Lki giacd iivy at Capa, t ctid t at it. Aggi wa vd i a gp ad at a cati, t wa t citty qatiid.

Dici O t cid th xitc th gp g-taid acaq ivig withi th c at Mky Jg. Th th gp ay d at dit ad citt cati. At y cati, th Wid Mky Swiig P tw gp w vd dig, t t at th a ti. Cap gp wa pt at th p dig twic a t a Lki. Sigiicaty tw ccai Cap gp wa vd t dipac Lki Gp. W itpt thi a vidc that Cap gp wa th diat gp. Dig th dig xpit Capa ad Tiki dipacd at Lki ti gp (Lki wa th w hi gp that aid t h a giacd at Cap). Additia ppt thi c th d chic xpit dig which th acaq cay vad th it ad pat tha th icit. At th Wid Mky Swiig P th ky w d ixd it ad tw highy pizd had id gg. Ittigy at th tat tdy Cap gp wa tiy pt at th ig p hw. Twad th d tdy Lki gp pacd Cap a th gp vd at th ig p hw. Thi chag cicidd with th ith th ay th hight-akig a (Ggia) ad ca atiach Lki gp. Iat ica th tat thi th (va Ndwijk ad va Schaik, 987) ad i thi patica cptitiv dig itati it i vy pi that th ith Ggia iat cd high tat t Lki gp a a wh. Withi a th gp a iia patt dig cptiti gd. Ipctiv whth th d it w it, pat, icit, ctt cptiti tw adt ad jvi wa citty vd. F th th d typ ctt cptiti wa vd tw adt a ad a dpit th act that adt a acaq a qty diat v th a (Kig, ). Th vati i th tat w-akig a wig th ith h iat ad h qt aiity t d i citt with what i kw at dig cptiti i a acaq ( av). That adt hw ctt cptiti with jvi g-taid acaq i citt with th vati Saj ad Scictt (7). Th ac ctt cptiti tw th tw x i pzzig. Hwv i at a acaq av thi ata gp at xa atity ad igat t ath gp. Th pc a chai ik ctiid c dig th pit pvt th Mky Jg acaq avig thi gp (Rw, 996). Cqty, i ctat t acaq i at th a ad a acaq at Mky Jg a atd. Fth, th act that a th acaq a th dcdat th igia ix ad i 9 ak th acaq at Mky Jg highy id. Thi c gtic atihip ay acct th ac ti ctt cptiti d tw th x. A iia vati ha vd tac c-dig i atd Japa acaq (Bi ad Chapai, ). Hwv at th Wid Mky Swiig P dig a acaq w ch iky t tiv th gg tha a, hwig that thi icdiy iitd ad cpd c, th i

cptiti tw th x. Bca w akig a tivd gg, w ccdd that ctt cptiti did xit v tw th adt a. A xpctd i ctt cptiti, Capa, th apha a, gt th t gg ad th jvi did t gt ay gg. Bca th gg w i gaty iitd ppy cpad t th ixd it, th high akig a w a t piz th highy cpd c. Thi cad th w akd th gp t xcdd gttig th gg. O a pactica v th pviiig chd ad cati th g-taid acaq at Mky Jg adqat t t thi titia d. Dipig (thwig) th d appad t giv th acaq, icdig th jvi, adqat pptity t tai d at th dig it. N gp wa cpty xcdd th dig, ad i act t th gp acaq w vd at a dig it at a daiy ai. Lki gp wh akd cd at Cap appad t hav adptd cavgig th cagd acaq withi th cpd a a ga c titi. Hwv, thy w v cpty xcdd th p hw a thy typicay d th dig th at dig ad dig th ig wig th ith Ggia ay. Th ctt cptiti that xit tw th Mky Jg acaq d t appa t it with thy titia w ig ad thi aiity t ag atay i thi c i a addd it. Pdct Iati Mky Bicit Mazi High Pti Piat 5668. Ackwdgt W wd ik t thak Tacy Cak wh aitd with ay apct th acaq, Tia Caq wh aitd with th dig th acaq, Mky Jg ttig it ky ad c a w a ttig a thi gd, D. Eva adig ach, ad th ky cpatig ad ttig tak th;

Fig ad Ta Kig Iad Wid Mky Swiig P Kitch Sit Pat O Aia Phtgaph Macaq Cpd at Mky Jg (Pit Fc Otid) idicatig th th dig cati.

Appdix: Ma ad tadad dviati pctag ag/x ca atig at ach th it. P a a jvi Ma.55667.558696. SD.779.9855.59 Kitch a a jvi Ma.857.8768.59695 SD.685.7898.5956 Kig Iad a a jvi Ma.778.6869.6785 SD.6677.675.7 Litat Citd Bi P. ad Chapai B.. Tatd c-dig i ati t dg kihip i Japa acaq. Bhavi, 8, 87-59. d Rit J.99. Bhavi ad g i ata ppati g-taid acaq (Macaca acicai) Pihd Dctia thi Uivity Utcht. Utcht, Had Ja CH. ad Schaik CP Va. 988. Rcgizig th May Fac Piat Fd Cptiti: Mthd. Bhavi, 5, 65-86. Kcza EM.998. Ipact iat taity ad kwd x ati th captiv aagt th i- agig ppati g-taid acaq (Macaca acicai) at ky jg, Miai, Fida. Upihd at Scic thi Fida Itatia Uivity, Miai Fida. Mati P ad Bat P. 99. Maig Bhavi: A Itdcty Gid- Scd Editi. Caidg Uivity P. Rw N. 996. Th pictia gid t th ivig piat. Eat Hapt, Nw Yk: Dgia P. Saj TL, 7. Fdig cptiti i Macaca acicai: A at th ay aiva tactic. Itatia Ja Piatgy. 8() Spig 58-56. va Ndwijk MA ad va Schaik CP. 987. Cptiti ag a g-taid acaq, Macaca acicai. Aia Bhavi, 5, 577-589

va Schaik CP ad Ndwijk MA Va 987. Sca ad ctt i dig cptiti ag a g-taid acaq (Macaca acicai) Bhavi, 5, 77-98.