Tropical Biodiversity 6(3): 179-187 (1999) DISPERSAL IN PROBOSCIS MONKEYS (Nasalis larvatus) IN THE LOWER KINABATANGAN, NORTHERN BORNEO Ramesh Boonratana!? 1. Wildlife Conservation Society, 185" Street and Southern Boulevard, Bronx, NY 10460-1099, USA. 2. P.O. Box 54, Chiang Mai University, Chiang Mai 50202, Thailand, ABSTRACT Nasalis larvatus is a large, sexually dimorphic, monotypic arboreal colobine, endemic to the island of Borneo, where it is largely restricted to riverine, peat swamp and mangrove forests of the coastal lowlands. The social structure of N. larvatus is flexible, comprising relatively stable one- male, all-male and predominantly male non-breeding groups. Changes in composition were recorded in aN. farvatus one-male group, during a two-year study carried out in the Lower Kinabatangan, northern Borneo. Both male and female N. larvatus were observed to depart from the group. Male emigrations are most likely to avoid inbreeding and intra-sexual competition. Female emigrations could result from various reasons, which include inbreeding avoidance, infanticide avoidance, improved dominance status, and reduced feeding competition. Keywords; Proboscis monkey, Nasalis larvatus, Dispersal, Lower Kinabatangan. Received: 16 March 1999; Accepted: 12 November 1999 INTRODUCTION Only recently did long-term studies (Bennett, 1986; Bennett and Sebastian, 1988; Yeager, 1990a; Boonratana, 1993) report dispersal in NV. Jarvatus, These long-term stud- ies carried out at Samunsam Wildlife Sanctuary in Sarawak (Bennett, 1986; Bennett and Sebastian, 1988; Rajanathan and Bennett, 1990), at Tanjung Puting National Park in Kalimantan (Yeager, 1989, 1990a, b, 1991a, 1992), and in the Lower Kinabatangan area of Sabah (Boonratana, 1993), showed that N. farvatus has a flexible social structure with a one-male group (OMG) as the basic social unit. In addition, young juvenile males leave their natal group and join loosely bonded all-male groups (AMGs) (Bennett, 1986; Bennett and Sebastian, 1988; Yeager, 1989; Rajanathan and Bennett, 1990; Boonratana, 1993), Besides male emigrations, female N. larvatus were also observed to depart from OMGs (Bennett, 1986; Bennett and Sebastian, 1988; Rajanathan and Bennett, 1990; Boonratana, 1993), This paper describes the age/sex composition of N. larvatus groups in the Lower Kinabatangan region of northern Borneo, made during a two-year study (Boonratana, 1993) on the ecology and behavior of V. larvatus. It further describes the changes in composition of a focal group, and discusses the possible reasons for dispersal in N. larvatus180. Dispersal in Proboseis Monkey METHODS The study was conducted at two sites, Sukau (118°30"E/5°30"N) and Abai (118°32°E/ 5°41'N), located along the Kinabatangan River in eastern Sabah, Bornean Malaysia, from January 1990 to December 1991, The Lower Kinabatangan region is mostly under forest on flat land that had been subjected to different degrees of disturbance. The forest at Sukau is predominantly riverine, whereas at Abai it is predominantly mangrove. Open water, hills, villages and oi! palm plantations are scattered throughout the region (Boonratana, 1993) ‘To obtain information on the groups’ age/sex composition, monthly census surveys were conducted along the rivers, N. Jarvatus groups sleep in trees next to rivers every night (Kern, 1964; Kawabe and Mano, 1972; Jeffrey, 1979; Macdonald, 1982; Salter er al., 1985; Bennett, 1986; Bennett and Sebastian, 1988; Rajanathan and Bennett, 1990; Yeager, 1989, 1991a, b; Boonratana, 1993). ‘Thus, by travelling in a boat along the rivers before dusk and just after dawn, it was possible to observe most groups. Surveys were conducted using a small boat fitted with a 25-HP outboard engine. The surveys began at 1630 hours and usually ended at 1830 hours, between the time when animals arrived at the river and when it was too dark to count well. The surveys were repeated the next morning at 0545 hours following the same route taken the previ- ous evening and ending at 0715 hours. This allowed two attempts at group counts, and age/sex identifications to be made for any one night (Bennett, 1986; Bennett and Sebastian, 1988; Rajanathan and Bennett, 1990; Boonratana, 1993). Each month, fifteen nights (five nights by the author and ten nights by the author’s field assistant, Mr. D.S. Sharma) of river surveys were conducted at Sukau, and five nights at Abai (Boonratana, 1993). RESULTS Composition of N. larvatus Groups Based on individually distinctive animals within groups, six OMGs, one AMG and two non-breeding groups (NBGs) at Sukau, and one OMG at Abai were consistently recognized throughout the study (Table 1). An NBG refers to a loosely bonded predominantly male gtoup with at least one female member (Boonratana, 1993). The OMGs at Sukau averaged 17 individuals per group, and ranged from 14 to 20 individuals per group. Repeated obser- vations at Abai (n=324) showed that OMGs averaged 14.6 individuals, and ranged from 8 to 2 individuals per group. The adult sex ratios at Sukau and Abai were 1:7.3 and 1:7.1 re- spectively. At Abai, however, there were three occasions when a sub-adult male was seen with an OMG. A similar observation was made at Samunsam Wildlife Sanctuary (Rajanathan and Bennett, 1990). It was likely that those sub-adult males were the OMG males’ offspring, which for unknown reasons remained with the OMGs. A more detailed study is required to confirm this.Tropical Biodiversity 6(3): 181 (1999) Table 1. Age/sex composition of identified groups in the Lower Kinabatangan SUI SUZ SU SU4 SUS’ SUG SU? SU8* SUS ABI AM oo 1 1 1 1 1 3 3 3 1 AF 8 7 7 8 8 6 0 0 0 8 SuM 0 0 0 0 0 0 2 2 3 0 Sar 0 1 1 1 Il 0 0 0 0 2M oO 0 0 0 0 0 2 3 2 0 DEO 3 2 1 0 1 0 0 2 0 2? 0 3 0 1 0 0 0 0 0 0 uM 3 0 0 0 0 0 0 0 0 2 JIF 2 0 0 0 0 1 1 0 0 0 20 3 2 1 2 L 0 0 0 0 RM 1 0 2 0 0 0 0 0 0 2 DF 1 0 2 0 0 L 1 0 0 1 12? 1 2 0 3 3 Il 0 0 4 IM 0 0 0 0 0 0 0 0 0 0 UF 0 0 0 0 0 0 0 0 0 0 n? 1 0 0 1 0 I 0 0 0 0 Sze 20 19 17 17 15 14 9 8 1018 Note: [Group size only from first complete count] ult; Sa=subadult; J2=older juvenile; ~ SU=Sukau; AB=Abai; M=male: juvenile; I2=older infant; 1 group. =young non breeding ‘At Sukau, the three non-OMGs (AMGs and NBGs) averaged nine individuals per group, and ranged from eight to ten individuals per group (Table 1). Non-OMGs fre- quently changed their membership. There were, however, some groups whose member- ship was consistent over a long period. Group SU7, for example, remained unchanged for 15 months, since it was first identified until the end of the study. Furthermore, it had two female members, a juvenile-1 and an infant-2. Another NBG, SU9, also had two female members that remained with the group for at least seven months. AMGs at Abai averaged 6.4 individuals (n=54), and ranged from four to nine indi- viduals. Occasionally, they were observed with two or more fully adult males (n= ‘Typically, AMGs at Abai comprised an adult male, several sub-adult and juvenile males. ‘At Sukau, solitary N. larvatus of both sexes were encountered on 22 occasions. This was recorded when an individual was more than 20m from the nearest conspecific (Boonratana, 1993). Adult males comprised 73%, adult females 5%, sub-adult males 18%, and sub-adult females 5% of all solitary observations. Twice, an adult female and once, a sub-adult female, were also briefly seen associating with AMGs. These associations normally did not last more than a few days.182. Dispersal in Proboscis Monkey At Abai, solitary N. larvatus were encountered only four times, A solitary adult male was seen twice, a sub-adult female once and a juvenile-2 male once. Once, an adult female with a clinging infant-2 was seen to associate with an AMG for a day. Changes in Composition of SU1, the Focal Group SUI was first identified in August 1990, and was observed every month until De- cember 1991 (Table 2). The group size of SU1 averaged 21 and the adult sex ratio was 1:8.4, varying from eight to nine adult females. Changes in group membership occurred during the study, but not all changes could be determined because not every member of SUI was recognized. Most changes in age/sex composition of SUI can be attributed to the reclassification of younger animals as they grew older, The adult male Tukkae, two adult females Martina and Choon, and their female offspring, Lucy and Pe-pex remained with the group throughout the study. All these individuals were observed within SUL during the author's re-visit to the study area in November 1992. By November 1990, the group size reduced to 19 individuals when a juvenile-1 male, Stanley, left the group. This presumably occurred when he became a juvenile-2, In February 1991, the adult female Martina, was observed with her female infant-1, Lucy. Martina most likely gave birth to Lucy either in late January or early February. This increased the group size to 20 individuals. In March 1991, there was another newborn in the group, increasing the size to 21 individuals, Table 2. Demographic changes in the focal group SUI Aug. Sep. Nov. Feb. Mar. Apr. Sep. Oct. Nov. Dec. 1990 1990 1990 1991 1991 1991 1991 1991 1991 1991 AMI 1 1 L 1 1 1 1 1 1 AR’ 8) Hh 9 ge BR 9 Se oR Bo SaF 14 9B 9 RR Be a RF ot 1 {feel gk 1 t iM o = 29 2 8 8 1 i 1 1 Ji Ss) ce 1 1 1 a i Es! Ge sh ete efle nih: SS Ge "S RM 1 Il i fo oof 8 Oo UO GF DF 1 t 2 it 1 2 1 0 0 0 2? 4 1 1 He hei |. OR ee Bg MM Oe Be tO, uF OL 1 0 1 Do oe 8 Oe eo WE OD Me OB Te Be Si 20 20 19 20 21 22 2 22 23 23 Note: M=male; adult; Sa=subadult; J2=older juvenile; J1=young juvenile; I2=older infant; T1=young infant; 2=sex unknown. [Note: Group size only from complete counts}