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    Social Organisation of Proboscis Monkeys (Nasalis larvatus) in the Lower Kinabatangan, Sabah, Malaysia

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    Social Organisation of Proboscis Monkeys (Nasalis larvatus) in the Lower Kinabatangan, Sabah, Malaysia

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    Social Organisation of Proboscis Monkeys (Nasalis larvatus) in the Lower Kinabatangan, Sabah, Malaysia

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    128 views19 pages

    Social Organisation of Proboscis Monkeys (Nasalis Larvatus) in The Lower Kinabatangan, Sabah, Malaysia

    Uploaded by

    Ramesh 'Zimbo' Boonratana
    Social Organisation of Proboscis Monkeys (Nasalis larvatus) in the Lower Kinabatangan, Sabah, Malaysia

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    Attribution Non-Commercial (BY-NC)
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    Malayan Nature Journal 2002, S6(1),57 - 75 Social Organisation of Proboscis Monkeys (Nasalis larvatus) in the Lower Kinabatangan, Sabah, Malaysia RAMESH BOONRATANA' Abstract, Nasalis larvatus is a large, sexually dimorphic, monotypic arboreal colobine, endemic to the island of Bomeo, where itis largely restricted to riverine, peat swamp and mangrove forests of the coastal lowlands. The paper describes the social organisation of WV. larvatus observed from January 19900 December 1991, in the Lower Kinabatangan region of northem Borneo. The basic social unit of N. larvatus is a one-male group, comprising a single adult male, several adult females and their offspring. The social structure of N. larvatus in the Lower Kinabatangan is flexible, and comprised relatively stable one-male, all-male and predominantly male non-breeding groups. Different groups frequently selected river-side sleeping sites that were close to one another. Some groups associated ‘more than otherstdid, implying a secondary level of social organisation, the band. Key Words. Proboscis monkey, Nasalis larvatus, social organisation, Borneo. INTRODUCTION It was not until recently that long-term studies made available information concerning the social organisation of Nasalis larvatus. Earlier observations gave conflicting reports. Studies conducted in Brunei Bay (Kern 1964; Macdonald 1982) and in East Kalimantan (Jeffrey 1979) reported N. larvatus living in loosely organised multi- male troops that mixed and separated frequently. Another study in Sabah (Kawabe and Mano 1972), however, reported that multi-male groups were highly integrated and cohesive. More recent studies 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 and b; 1991a; 1992), and in the Lower Kinabatangan area of Sabah (Boonratana 1993), have shown that NV. larvatus has a flexible social structure. The basic social unit is a one- male group (OMG), comprising one adult male, several adult females, and their offspring (Bennett 1986; Bennett and Sebastian 1988; Yeager 1989; Rajanathan and Bennett 1990; Boonratana 1993). This corresponds to the modal pattern of social organisation in the Colobinae (Newton and Dunbar 1994). 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 199% ' P.O. Box 54, Chiang Mai University, Chiang Mai 50202, Thailand. Email: rboonratana@ hotmail.com. 57 Boonratana 1993). Furthermore, there seem to be two levels of social organisation, the OMG and the band (Yeager 1989; 19914; Boonratana 1993). This paper describes and compares the social organisation of N. larvatus at two sites in the Lower Kinabatangan region of Sabah, observed during a two-year study on its ecology and behaviour (Boonratana 1993), conducted from January 1990 to December 1991 STUDY AREA ‘The study was conducted at Sukau (118°30"E/5°30°N) and Abai (118°32°E/5°41°N), located along the Kinabatangan River in eastern Sabah, Malaysia. 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 plantations are sparsely scattered throughout (Boonratana 1993). ‘The Kinabatangan River is Sabah’s largest river, with a length of 560 km, and catchment area of 16,800 km (Scott 1989). The lower Kinabatangan River meanders through a large flat floodplain, much of which is subject to seasonal flooding, resulting in low-stature forest with little timber of commercial value. The Kinabatangan floodplain, measuring approximately 280,000ha, is the largest and possibly the most important wetland in Sabah (Scott 1989), The river finally empties into a large delta, which is brackish and tidal. Mean temperatures did not vary much between months during the study period The mean monthly minimum for 1990 and 1991 was 23.7°C, whereas the mean monthly maximum for 1990 was 32.9°C, and 33°C for 1991. The total rainfall for 1990 was 1,816mm, and for 1991 it was 2,975mm (Boonratana 1993). METHODS To obtain information on N, larvatus’ social organisation, full-day observations from dawn to dusk were made, using the scan sampling method (Altmann 1974). All observations were made using a Zeiss Dialyt 10x40B binoculars. At the Sukau study area, a focal group, SUI (Table 1) was identified and observed. Whenever SUI could not be located, then observations were made on other N. larvatus groups. Observations were made from the boat in the morning before SUI moved away from the riverside and in the evening after the group returned to the riverside. During the day when SUI moved into the forest, the group was followed on foot At Abai, it was not possible to observe any one group continuously throughout the day. This was partly due to the shyness of the study animals to the observer on foot, but mainly due to the forest being flooded during high tide. The ground was very soft and muddy during low tide, making quick and noiseless follows impossible. The presence of Crocodylus porosus in the area was also a deterrent. Thus, almost all observations were made from the boat, when the animals were by the river. The author remained with a group for as long as possible and then searched for another group when the first group was no longer visible 58 ‘Table 1. Age/sex composition of identified groups at Sukau. suI® SU2* SU3® SU4® SUS® = SU6® SUT = SU8m SUS AM AF SaM 32M nF mr JIM JF m 12M 12F 12? 11M IF mm Sue Il 1 1 I 1 1 3 3 3 8 7 7 8 8 6 0 0 0 ° 0 0 0 0 0 2 2 3 1 0 1 1 1 1 0 0 0 0 0 0 0 0 0 2 4 Z 1 3 2 1 0 1 0 0 2 0 3 0 1 0 ° 0 0 0 3 0 0 0 0 0 0 0 0 2 0 0 ) 0 ' 1 0 0 0 3 a 1 2 1 0 o 0 1 0 2 0 0 ° 0 ° 0 1 ) 2 0 0 1 1 o 0 1 2 0 3 3 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0) 0 0 0 0 0 0 0 1 0 0 1 0 1 0 0 0 20 19 7 7 1s 4 9 8 10 M=male; F= female; 12=older infant; | subadult; J2 = older juvenile; J1 = young juvenile; young infant; ?= sex unknown; *= one male group; #= non breeding group, Scan samples were recorded during a 2-minute period every 15 minutes from dawn to dusk on every full day follows, and encompassed all members of the group that could be observed during that period. An “observation” refers to one animal recorded during each scan. To obtain information on social organisation, the following data were recorded once during every observation: 1 spread of group along the river, i.e. the distance from the first to the last individual in that group: observed number of individuals, i.e. the minimum number present; estimated number of individuals, i.e. probably a more accurate assessment of the number present; age and sex of observed individual; identity of observed individual, if known, age and sex of the individual nearest to the observed individual (the nearest neighbour); the distance from the observed individual to the nearest neighbour: the number of other group members within 2.5m and 5m of the observed animal (Struhsaker 1975; Struhsaker and Leland 1979). 59 RESULTS Sukau Study Area ‘Throughout the study, six OMGs, one AMG and two non-breeding groups (NBGs) were consistently recognised using individually distinctive animals as markers (Table 1). An NBG refers to a loosely bonded predominantly male group with at least one female member (Boonratana 1993). The OMGs averaged 17 individuals per group, and ranged from 14 to 20 individuals per group. The adult sex ratio of the OMGs was 1:7.3. They were relatively stable over time and none were observed with more than one adult male. The three non-OMGs (AMGs and NBGs) averaged nine individuals per group and ranged from eight to ten individuals per group. Non-OMGs frequently changed their membership. There were, however, some groups whose membership 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-I and an infant-2. Another NBG, SU9, also had two female members that remained with the group for at least seven months. Throughout the study, solitary NV, arvatus 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. Group Spread It was difficult to estimate the spatial distribution of N. farvatus groups in the forest because of limited visibility. Group spread was estimated only when groups were roosting by the river (Boonratana 1993). The average group spread of OMGs was 13.2 m (n=396, range 3-50 m). Non-OMGs had a similar average spread at 13.1 m (n=73, range 2-35 m). Although similar in average group spread, the average group size of OMG was almost double that of the non-OMG. Apparently, non-OMGs were not as cohesive as OMGs. Intra-group Associations Details of social organisation of SUI were assessed from the percentage of time an individual was nearest to the subject (Table 2). The expected percentage for which an individual would have been nearest to the subject if all members of SUI were associating randomly was determined from the average composition of SUI throughout the study. Chi-square tests were not performed because data were not statistically independent. Results can be summarised as follows: 1, The adult male spent more time than expected near the adult females but less time than expected near all the other age/sex classes; 60 2. Adult females spent more time than expected near the juvenile-Is and infants, but less time than expected near all other age/sex classes; 3. Sub-adult females spent more time than expected near the adult male and adult females, but less time than expected near all other age/sex classes; 4. Juvenile-2s spent more time than expected near the adult male, adult females and other juvenile-2s, but less time than expected near all other age/sex classes; 5. Juvenile-1s and infants spent more time than expected near the adult females, but less time than expected near all other age/sex classes. Thus, observed combinations show that associations between any two members were not at random but specific. ‘Table 2. Percent time members of SUI belonging to different age/sex categories were nearest to subject at Sukau study area (n=4966, weighted data). Values in parentheses show the percent time which the subject would spend nearest to an individual ofthat age/sex category if all individuals were associating randomly. Nearest Neighbour Subject AM AF SaF pon Rn nl a AM = 78.9(40) 0.2010) 4.3(5)15.5(25) L115) (5) 382. AF 3.715) 26.2(35) 0.410) 3.0(5) 29.5(25) 25.1(15) 12.25) 2540 Sak 11.405) 78.640) (5) (5) 9.9125) KIS) S) 33 Ja 4.715) 70.5(40) 0.9(10) 3.6(0) 13.9(25)4.3(15)1.9(5) 259 i 3.115) 76.5(40) (10) 1.5/5) 14.6(20) 2.815) 1.365) 1071 2 0.4(5) 89.8(40) 0.6(10) 1.115) 3.2(25) 4.210) 0.65) 491 n (5) 94,0(40)O(10)0.6(5) 1.825) 1.4(10) 2.35) 190 M=male; F=female; A=adult; Sa=subadult; J2=older juvenile; J1=young juvenile; 12=older infant; H=young infant Intra-group Spacing Intra-group spacing was assessed from the distance between the subject and its nearest neighbour (Fig. 1), and by recording the number of other individuals within 2.5m (Fig. 2) and 5m (Fig. 3) of the subject (Struhsaker 1975; 1980; Oates 1977; Struhsaker and Leland 1979; Bennett 1983). The results are summarised as follows: 1. The adult male was less than Im from any other individual for more than 50% of the time. Most of that time was spent close to an adult female. Each adult female spent over 90% of her time within 2m of another individual. Excluding adult females with clinging infants, each adult female 61 Figure 1. Percentage of time nearest neighbour was within cumulative distance from subject at Sukau (n = 4966, weighted data). 120 100 -| 80 - % Scans 40 | 20 -} o T T T T—T 0 — >0-1 >I-2->2-3 3-4 >4-S Distance (m) Figure 2. Percentage of time different numbers of individuals were within 2.5m of subject at Sukau (n = 4966, weighted data). 30 25 20 % Scans a4a2 4 5 6 7 8 9 10 >10 No. of individuals WH acaMac — E Alsh eonale sua Mower nvcnte Young evente i) Over Ea Young ine a 62 Figure 3. Percentage of time different numbers of individuals were within Sm of subject at Sukau (n = 4966, weighted data) % Scans 12° 3 4 5 6 7 8 9 10 >10 No. of individuals Eee SF paneer irre (2) Yemelneie fi Ota spent about half her time within Im of another individual. This was less than for the adult male. 3. Each sub-adult female and juvenile-2 spent 62.3% and 58% of their time respectively within 1m of another individual. They were more gregarious than either the adult male or the adult females. 4. Juvenile-1s and infants (infant-1s and infant-2s) spent more than 75% of the time near another individual. Each infant was less than 1m from any other individual more than 95% of the time. This was because they rarely ventured more than 2m from an adult female. Although SUI had a large membership, it formed a cohesive group, with small inter-individual distances, and the adult male was as gregarious as the adult females (Boonratana 1993). Inter-group Associations Different groups frequently come together to spend the night next to the rivers (Bennett and Sebastian 1988; Rajanathan and Bennett 1990; Yeager 1991a and b). SUI spent 60.4% and 68.8% of its nights within 50m and 100m respectively of another group (Fig. 4). SUI spent more nights within 100m of anon-OMG (57.3%) than with another OMG (54.2%). The non-OMGs probably followed SU1 or other OMGs that associated with SUI. A non-OMG followed SUI from their sleeping trees on 77 out of 93 complete and incomplete full-day follows. When neighbouring 63 groups could be identified with certainty, SUI spent most nights close to OMG SU2 (39.6%) and non-OMG SU7 (45.8%). These associations suggest a secondary level of social organisation, the band, with fission-fusion of stable OMGs within bands (Yeager 1989; 1991; 1992; Boonratana 1993). Figure 4, Percentage of nights when other N. Jarvatus group/groups were close to SUI at Sukau (n= 96). 70 5 oo so} 2 404 s eS © 304 20 4 04 o4 -

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