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Feeding Ecology Orangutans in Danum

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Feeding Ecology Orangutans in Danum American Journal of Primatology 72:820–840 (2010) RESEARCH ARTICLE Feeding Ecology of Bornean Orangutans (Pongo pygmaeus morio) in Danum Valley, Sabah, Malaysia: A 3-Year Record Including Two Mast Fruitings 1Ã 2 3 4 2 TOMOKO KANAMORI , NOKO KUZE , HENRY BERNARD , TITOL P. MALIM , AND SHIRO KOHSHIMA 1Department of Biological Sciences, Graduate School of Bioscience and Biotechnology, Tokyo Institute of Technology, Ookayama Meguro-Ku, Tokyo, Japan 2Wildlife Research Center of Kyoto University, Kyoto, Japan 3Sabah University of Malaysia, Kota Kinabalu, Sabah, Malaysia 4Sabah Wildlife Department, Kota Kinabalu, Sabah, Malaysia We observed the diet and activity of Bornean orangutans (Pongo pygmaeus morio) in the primary lowland dipterocarp forests of Danum Valley, Sabah, Malaysia, during 2005–2007, including two mast fruitings. We collected 1,785 hr of focal data on 26 orangutans. We identified 1,466 samples of their food plants and conducted a fallen fruit census to monitor fruit availability in the study area. Their activity budget was 47.2% feeding, 34.4% resting, and 16.9% traveling. Fruits accounted for the largest part (60.9%) of feeding time, especially during mast fruiting periods (64.0–100%), although the percentages of leaves (22.2%) and bark (12.3%) were higher than those reported for P. abelii and P. pygmaeus wurmbii. Although 119 genera and 160 plant species were consumed by focal animals, only 9 genera accounted for more than 3% of feeding time (total: 67.8% for 9 genera). In particular, the focal orangutans fed intensively on Ficus and Spatholobus during most of the study period, especially in periods of fruit shortage. The percentage of fruit feeding changed markedly from 11.7 to 100% across different months of the year, and was positively correlated with the amount of fallen fruit. When fruit feeding and availability decreased, orangutans fed primarily on leaves of Spatholobus and Ficus, and the bark of Spatholobus and dipterocarp. The percentage of time devoted to feeding during mast fruitings, when the orangutans foraged almost exclusively on fruits, was lower than during seasons when the orangutan diet included leaves and bark as well as fruits. Resting increased as feeding decreased in the late stage of each fruiting season, suggesting that the orangutans adopted an energy- minimizing strategy to survive the periods of fruit shortage by using energy stored during the fruit season. Am. J. Primatol. 72:820–840, 2010. r 2010 Wiley-Liss, Inc. Key words: Bornean orangutans; feeding ecology; mast fruiting; fallback foods; activity budget INTRODUCTION Asian tropical forests, experience much greater inter- and intra-annual variability in fruit produc- Southeast Asian dipterocarp forests are known tion than do frugivorous primates in African and for their supra-annual masting events approximately South American rain forests [Fleming et al., 1987]. every 2–10 years, during which a majority of trees fruit in synchrony [Ashton et al., 1988]. Masts are periods of tremendous food abundance, permitting Contract grant sponsor: JSPS-HOPE (Primate Origins of high energy intake for various frugivorous verte- Human Evolution) Project Global Environment Research Fund, 21st Century COE Programs; Contract grant number: 0A06; brates [Curran & Leighton, 2000; Leighton & Contract grant sponsor: Graduate School of Science, Kyoto Leighton, 1983; Marshall & Leighton, 2006]. How- University, 21st Century COE; Contract grant number: A14; ever, mast fruitings are often followed by periods of Contract grant sponsor: Graduate School of Science, Tokyo Institute of Technology; Contract grant numbers: R12; A10; extremely low fruit availability [Knott, 1998]. Low Contract grant sponsor: The Japanese Association of Zoos and fruit production during intermast intervals has been Aquariums (JAZA). reported to cause famine-like emaciation or death in ÃCorrespondence to: Tomoko Kanamori, Graduate School of the sun bear (Helarctos malayanus) and bearded pigs Bioscience and Biotechnology, Tokyo Institute of Technology, W3-43, 2-12-1 Ookayama Meguro-Ku, Tokyo 152-8551, Japan. (Sus barbatus) in mixed dipterocarp forests of E-mail: [email protected] Borneo [Curran & Leighton, 2000; Wong et al., Received 24 November 2009; revised 28 April 2010; revision 2005]. In addition to mast-fruiting events, smaller accepted 28 April 2010 annual and seasonal changes in fruit production also DOI 10.1002/ajp.20848 occur [Knott & Kahlenberg, 2006]. Thus, orangu- Published online 1 June 2010 in Wiley InterScience (www. tans, the frugivorous great apes living in Southeast interscience.wiley.com). r 2010 Wiley-Liss, Inc. Feeding Ecology of Bornean Orangutans / 821 However, detailed information on their feeding state government in 1996 and managed by the Sabah ecology, especially during mast fruiting and their Foundation (Yayasan Sabah Group) for conserva- response to the drastic fluctuations in fruit avail- tion, research, education, and tourism. The study ability, is limited [Knott, 1998; Morrogh-Bernard area (510101700N, 11714405000E, 231–384 m a.s.l.) is a et al., 2009; Wich et al., 2006]. primary forest of approximately 2 km2 surrounding a Pongo pygmaeus morio is the most eastern tourist lodge, Borneo Rainforest Lodge (BRL). BRL subspecies of Bornean orangutan, and inhabits was established in 2004 on the banks of the Danum forests in which mast-fruiting events occur in more River, a tributary of the Segama River. The forest is tree species and individual trees than reported for composed predominantly of lowland dipterocarp habitats of other Bornean orangutan subspecies forest, interspersed with some lower-hill dipterocarp (P. pygmaeus wurmbii and P. pygmaeus pygmaeus forest. Canopy height is approximately 50 m, with in the central and western Borneo), and the emergent trees more than 70 m. The mean tree Sumatran orangutan (P. abelii) [Curran et al., 1999; density in this area was reported to be 470 trees per van Schaik et al., 2009a; Walsh, 1996]. Masting hectare (430 cm dbh), and the trees in the forests events in Sumatra are less prominent than in Borneo (430 cm dbh) included Euphorbiaceae (21%) and because fewer mast-fruiting tree species exist in Dipterocarpaceae (16%), with basal areas of 7 and those forests [van Schaik, 1996]. Moreover, the 49%, respectively [Newbery et al., 1992]. Approxi- influence of the El Nin˜o/Southern Oscillation on mately 88% of the total volume of the large trees in mast fruiting is reportedly strongest in the eastern the conservation area was reported to be diptero- part of Borneo [Ashton et al., 1988; Wich & van carps [Marsh & Greer, 1992]. Schaik, 2000]. Thus, P. pygmaeus morio living in the dipterocarp forests of East Kalimantan and Sabah, Study Period experience dramatic fluctuation in fruit availability [Leighton & Leighton, 1983; Rijksen & Meijaard, We conducted field research from March 2005 to 1999]. P. pygmaeus morio exhibits the most robust December 2007, 27 months in total, although we only jaw and smallest skull of any orangutan taxa, obtained limited data in April–May 2005 and suggesting that this species has adapted to ecological November 2005–March 2006. We obtained no data conditions different from those of other Bornean in February 2006 owing to serious flooding in the subspecies and the Sumatran orangutan [Taylor, area. The study period included two mast fruitings 2006, 2009; Taylor & van Schaik, 2007]. Thus, the (July–August 2005 and May–September 2007). We study of P. pygmaeus morio is important for under- defined a mast year as one with months in which standing orangutan feeding ecology and their ability fruit scores were higher than 1.96 standard devia- to adapt to fluctuations in fruit availability. Earlier tions above the mean for the study period, following studies on P. pygmaeus morio have been conducted the definition for mast fruiting proposed by Wich and by Mackinnon in Ulu Segama and by Rodman and van Schaik [2000]. Leighton in Mentoko [Leighton, 1993; Mackinnon, 1974; Rodman, 1988]. Since these studies, however, Climate detailed studies of the ecology of this subspecies in The mean, maximum, and minimum tempera- primary dipterocarp forests has not been undertaken, tures in the study period, recorded at Danum Valley although studies have been conducted in logged Field Research Centre of DVCA next to the study area, forests in Kinabatangan, Sabah [Ancrenaz & were 27.11C, 31.71C, and 22.61C, respectively (The Lackman-Ancrenaz, 1999; Ancrenaz et al., 2004, 2005]. Royal Society Southeast Asia Rain Forest Research In this study, we conducted field research on the Program, unpublished data). Mean monthly tempera- feeding behavior and activity budget of P. pygmaeus turestendedtobehigherinAprilandMayandlower morio living in the primary lowland dipterocarp in November and December. Annual rainfall in the forests of Danum Valley, Sabah, Malaysia, from 2005 area varied between 3,083.1 and 2,349.5 mm during to 2007, including two mast fruiting periods. We 2005–2007. Mean monthly rainfall was 234.8 mm compared seasonal changes in the feeding time (SE 5 22.1), ranging from 60.3 mm in April 2005 to allocated to individual food items and plant species, 798.4 mm in February 2006. Although the seasonality and examined the dietary response of orangutans to of rainfall was not clear during the study period, major fluctuations in fruit availability. 10-year mean values of monthly rainfall from 1988 to 2007 show that rainfall in this area tended METHODS to be higher in May–June and October–January and The Study Area lower in March–April and July–September. The study area was located in Danum Valley Conservation Area (DVCA, 438 km2), East Borneo, Observation of Animals Sabah, Malaysia (Fig. 1A). DVCA is a Class I We observed 26 identified individuals, divided Protection Forest Reserve established by the Sabah into the following six age–sex classes based on Am. J. Primatol. 822 / Kanamori et al. Fig. 1. Map of the study area. (A) The location of the study area, based at Borneo Rainforest Lodge in the state of Sabah, East Borneo. (B) Transect around Borneo Rainforest Lodge. morphological characteristics [Kuze et al., 2005]: 3 females, 2 juveniles, and 5 infants (Table I).
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