Longitudinal patterns of reproduction in wild female siamang (Hylobates syndactylus) and white-handed gibbons (Hylobates lar)

I present the 6- year reproductive histories of three wild female siamang (Hylobates syndactylus)and four white-handed gibbons (Hylobates lar)at the Ketambe Research Station (Sumatra, Indonesia). Reproductive output varied considerably among females. Two females failed to gestate: both were nulliparous young adult H. lar,one of which remained unpaired for 4 years after dispersing from her group, while the other lost her recently acquired mate to another female. Only one- (a white-handed gibbon)- gave birth more than once, yielding interbirth intervals of 22 and 31 months. Pair bond stability or reduced interspecific feeding competition or both factors may have contributed to the brevity of these intervals. The other females- one H. lar,and three H. syndactylus-each gave birth once, suggesting minimum interbirth intervals exceeding 4–5 years (H. lar)and 3 years (H. syndactylus)in these individuals. Even given the pronounced variation observed among H. lar,these data suggest that interbirth intervals may often exceed the 2- to 3- year interval commonly attributed to these two species. Sources of reproductive failure were 1) maternal abandonment of the neonate due to impaired ability to provide maternal care (H. syndactylus,),(2) premature or stillbirth (H. syndactylus,),and (3) pregnancy termination (H. lar).These data and a review of information on longevity and age at menarche suggest that the actual lifetime reproductive output of a siamang or white-handed gibbon female may often fall far short of the 10 offspring/lifetime originally proposed for these species. Indeed, females may rear as few as five offspring to weaning in a lifetime, which is a figure reminiscent of the reproductive potential of some pongids. Finally, variance in female reproductive success is higher than expected in these monogamous species, which suggests that females (and males) are under strong selective pressure to exert mate choice, possibly through acquisition of (new) mates and extrapair copulations. Future research must clarify the availability of opportunities for paired adults to engage in these sociosexual behaviors.     

Intragroup and Intergroup Social Interactions in White-Handed Gibbons

One of the least studied aspects of the Hylobatidae is their social behavior. In the past, researchers have attributed this to a general lack of affiliative social interactions due to small group size and the hostile nature of intergroup encounters. Data collected on white-handed gibbons (Hylobates lar) in Khao Yai National Park, Thailand, indicate that gibbons engage in higher rates of social behavior than had previously been appreciated. In addition to relatively high rates of intragroup social contact, gibbons engaged in social behavior, including play and grooming, with members of neighboring groups. These findings obligate a renewed consideration of the nature and function of territoriality in primates.     

The yellow-cheeked gibbon (Hylobates gabriellae) in nam bai cat tien (southern vietnam) revisited

Data concerning the status, habitat, and vocalizations of yellow-cheeked crested gibbons (Hylobates gabriellae) were collected during a short field trip to the Nam Bai Cat Tien National Park (southern Vietnam). Nam Bai Cat Tien may be the southernmost locality where crested gibbons (i.e. theHylobates concolor group) still survive. Fewer songs were heard at Nam Bai Cat Tien National Park than at other crested gibbon sites visited by the author. At least two gibbon groups appear to have been greatly reduced in number since previous surveys in the park. There is some evidence that both the gibbon population and the gibbon habitat in Nam Bai Cat Tien are disturbed. The first case of a great call solo song in wild gibbons of theconcolor group is reported. Great calls ofH. gabriellae are described and documented with sonagrams for the first time. They differ from those previously described forH. leucogenys.     

Reassessment of age of sexual maturity in gibbons (hylobates spp.)

From studies of both wild and captive animals, gibbons are thought to reach sexual maturity at about 6 to 8 years of age, and the siamang (Hylobates syndactylus) at about 8 to 9 years. However, a review of the literature reveals that in most cases the exact age of the maturing animals was not known and had to be estimated. This study presents seven case reports on captive gibbons of known age. Captive males of the white-cheeked crested gibbon (H. leucogenys leucogenys) and of the siamang (H. syndactylus) can breed at the age of 4 and 4.3 years, respectively. Similarly, hybrid females (H. lar × H. moloch) and siamang females can breed at 5.1 and 5.2 years, respectively. This finding may help to improve the breeding success of captive gibbon populations. It is not clear whether gibbons reach sexual maturity earlier in captivity or whether sexual maturity is also reached by 5 years of age in the wild. Possible implications for the interpretation of group size regulation and of reproductive strategies of wild gibbons are discussed.     

Gibbon foraging decisions and the marginal value model

We use data from an observational field study of frugivory in two sympatric gibbons, lar (Hylobates lar) and siamang (H. syndactylus), to test assumptions and predictions of the marginal value model (MVM). A key prediction of the MVM is that marginal gain rates at the time of leaving the patch are equal across patch types. We found that this is not the case for gibbons: rates of energy intake at the end of feeding sessions were significantly different for different types of fruit, and we could not attribute this to temporal variation in fruit availability. Initial and final caloric intake rates were highly correlated. This suggests that gibbons do not adjust the time spent in patches in order to maximize the average rate of energy intake. Similar results were obtained for all other currencies considered. Gibbon foraging appears to satisfy several, but not all, assumptions of the MVM. As required by the model, fruit patches occur as discrete units, patches are encountered sequentially, travel time between patches exceeds search time between items within a patch, search for and search within patches are incompatible activities, and intake rates decline over time spent in a patch. However, the declining rates we detected may be an effect of satiation instead of patch depletion, patches probably are not encountered at random, and group members may not forage independently. Thus, our results suggest that the MVM is not an adequate model of gibbon foraging behavior, but they do not invalidate the MVM per se.     

A preliminary study of the wild siamang gibbon (Hylobates syndactylus) at Fraser's Hill,Malaysia

The wild siamang gibbon was studied at Fraser's Hill, Malaysia. The study area was covered with a well developed forest and offered a suitable habitat for siamang gibbons. Other primates living in this area were the white-handed gibbon, the duskey leafmonkey, the banded leaf-monkey, and the pig-tailed monkey. The siamang gibbon groups observed have a monogamous structure consisting of one pair of adult individuals and one or more subadults which are assumed to be the offspring of the adults. The adult male showed behavior typical of a group leader. As subadults become older, they tend to become spatially separated from the mother group. Each group was observed to range freely within an exclusive area of the forest into which no other group was observed to intrude. Each group emitted loud vocalizations which seemed to maintain the spacing between the groups.     

Homosexual behaviour in wild white-handed gibbons (Hylobates lar)

Homosexual mounting in apes and prosimians is rare. Male-male mounting was observed between an adult male white-handed gibbon and an adolescent male in the same group. The behaviour is discussed in terms of the gibbon group's social structure and the development of the adolescent. It is suggested that this behaviour provides reassurance for the adolescent gibbon.     

Interspecific spacing between gibbons (Hylobates klossii) and langurs (Presbytis potenziani) on Siberut Island,Indonesia

Spatial arrangement and social interactions of two sympatric and ecologically similar primate species, Hylobates klossii and Presbytis potenzianai, are described from field observations made between July 1972 and October 1974 on Siberut Island, Indonesia. Gibbon territories and langur home ranges overlap extensively. Because gibbons have the ability to supplant langurs at shared food sources, langurs are at a competitive disadvantage. To avoid or decrease the frequency of hostile interactions with gibbons, langurs locate their core areas on boundaries between adjacent gibbon territories, which permits langurs to retreat across these barriers in response to gibbon movements. Langurs further enhance segregration by leaving their sleeping trees earlier than gibbons, gaining additional feeding time on contested food sources. This form of interspecific spatial organization between gibbons and langurs resembles certain predator-prey spacing systems, where territorial boundaries between adjacent predators serve as sanctuaries for prey populations.     

Behavioural development of twin siamangs (Hylobates syndactylus)

Multiple births are very rare among gibbons. The birth of siamang twins at the Zürich Zoo in 1992 therefore presented a valuable opportunity to observe the development of the twins and to contrast it with a survey of previous reports on the development of single offspring of siamang and gibbons of thelar group. Furthermore, the hypothesis that the presence of twins among siamang may facilitate the occurrence of helping behaviour (defined as the care of offspring by individuals other than their parents) was re-examined (Dielentheis et al., 1991). The Zürich twins (one male and one female) were observed for a total of 74 hr during their first year of life. The results show that: (1) The twins exhibited more rapid behavioural development than that reported for single offspring. (2) A clear difference between the twins was observed: the female twin developed more rapidly than the male. (3) Neither the twins' father nor their older sisterRama was ever observed carrying the twins. The hypothesis ofDielentheis et al. (1991) is hence not supported by the present study, although it is possible that the older sisterRama did not carry the twins because she was younger than the juvenile in that study. (4) Siamangs may have a longer maturation period than gibbons of thelar group.     

How Reliable are Density Estimates for Diurnal Primates?

Primate population assessments provide the basis for comparative studies and are necessary prerequisites in determining conservation status. The most widely used assessment method is line transect sampling, which generates systematic data quickly and comparatively inexpensively. In contrast, the presumably most reliable method is long-term monitoring of known groups, which is both slow and costly. To assess the reliability of various analytical methods, we compared group and population densities for white-handed gibbons (Hylobates lar carpenteri) and Phayre’s leaf monkeys (Trachypithecus phayrei crepusculus) derived from transect walks with those from long-term group follows at Phu Khieo Wildlife Sanctuary, Thailand. Our assistants and we regularly walked a 4-km transect over 30 mo (480 km total), resulting in 155 gibbon sightings and 125 leaf monkey sightings. We then estimated densities via 1) DISTANCE and 2) the Kelker method based on perpendicular distances (PD) or animal-to-observer distances (AOD). We compared the 3 estimates to values based on known home ranges (95% kernels), accounting for home range overlap, combined with group size data. Analyses of line transect data consistently overestimated group densities for both species, while underestimating group size for leaf monkeys. Quality of results varied according to the group size and spread of each species. However, we found, in accordance with previous studies, that values derived via AOD (or its derivations) matched most closely with population estimates based on home range data.     

The relationship between duet songs and pair bonds in siamangs, Hylobates syndactylus

One of the most commonly cited functional explanations for animal duet songs is strengthening of the pair bond. However, the evidence to support this view is, at best, limited. This study provides support by documenting a relationship between pair bonds and duet singing in siamangs. As a working hypothesis, we assume that if duetting were related to pair bonding, we might expect to see a relationship between duetting intensity and indicators of pair bond strength. Like most gibbon species, siamang pairs produce loud, long and well-coordinated duet songs. We recorded daily frequency and duration of duetting and three generally accepted indicators of pair bond strength (mutual grooming, behavioural synchronization and distance between mates) in 10 siamang groups in zoos. Duetting activity was positively correlated with grooming activity and behavioural synchronization, and negatively correlated with distance between mates. These results suggest that the production of coordinated duets by siamang pairs is related to pair bonding. Copyright 2000 The Association for the Study of Animal Behaviour.     

Sympatry between white-handed gibbons (Hylobates lar) and pileated gibbons (H. pileatus) in Southeastern Thailand

White-handed gibbons (Hylobates lar) are not known to occur to the east or southeast of Bangkok. The reliably documented localities ofH. lar nearest to this area are about 120 km northeast of Bangkok. There, in the Kao Yai National Park, is the only known zone of contact betweenH. lar and the pileated gibbon (H. pileatus), another species of the so-calledlar group. Unpublished documents dating from 1925 indicate, however, that sympatry between these two species may also have existed in the region of Sriracha, about 80 km southeast of Bangkok. Therefore, a large zone of overlap in the distribution of the two species may originally have existed. In most parts of this hypothetical zone, gibbon habitat appears to have been destroyed, with the Khao Yai Park possibly representing the last remnant of the once large contact zone.     

Interactions between lar gibbons and pig-tailed macaques at fruit sources

Interactions are reported between white-handed gibbons (Hylobates lar) and pig-tailed macaques (Macaca nemestrina) in Khao Yai National Park, Thailand, in which gibbons selected ripe fruit from sources before macaques arrived on 4 occasions during June and July of 1989. The macaques foraged near gibbons or from shared fruit sources during 13% of gibbon observation time. Gibbons made their presence known in fruit trees when macaques approached on 3 occasions and macaques did not enter fruit trees occupied by gibbons. An aggressive interaction is reported in which a single white-handed gibbon vigorously excluded 22–28 macaques from a rare and valuable fruit tree (Sandoricum koetjape). The observations suggest that exploitative and interference competition may exist between these species.     

Are <Emphasis Type="Italic">Hylobates lar</Emphasis> Extirpated from China?

The Nangunhe Nature Reserve in Southwest Yunnan (PRC) has long been presumed to be the last stronghold of lar (or white-handed) gibbons (Hylobates lar) in China and the likely last place of occurrence of Hylobates lar yunnanensis. We conducted a comprehensive survey to assess the status of lar gibbons at Nangunhe. We found no visual or auditory evidence of them still residing at the reserve and therefore tentatively conclude that lar gibbons have become extinct in China. It appears that large-scale destruction of primary forests in the 1960s and 1970s brought about an initial decline in their numbers, and subsequent uncontrolled hunting has resulted in their extirpation. The situation for the six Chinese ape taxa is nothing less than disastrous, with 1 taxon assumed to have become extinct during the last few years, 1 taxon not having been confirmed since the 1980s, and 2 species at the very brink of extinction with only tens of individuals remaining in China.     

Visual preference in a human-reared agile gibbon (<Emphasis Type="Italic">Hylobates agilis</Emphasis>)

Visual preference was evaluated in a male agile gibbon. The subject was raised by humans immediately after birth, but lived with his biological family from one year of age. Visual preference was assessed using a free-choice task in which five or six photographs of different primate species, including humans, were presented on a touch-sensitive screen. The subject touched one of them. Food rewards were delivered irrespective of the subject’s responses. We prepared two types of stimulus sets. With set 1, the subject touched photographs of humans more frequently than those of other species, recalling previous findings in human-reared chimpanzees. With set 2, photographs of nine species of gibbons were presented. Chimpanzees touched photographs of white-handed gibbons more than those of other gibbon species. The gibbon subject initially touched photographs of agile gibbons more than white-handed gibbons, but after one and two years his choice patterns resembled the chimpanzees’. The results suggest that, as in chimpanzees, visual preferences of agile gibbons are not genetically programmed but develop through social experience during infancy.     

Food Preferences and Nutrient Composition in Captive White-handed Gibbons, <Emphasis Type="Italic">Hylobates lar</Emphasis>

We aimed to assess spontaneous food preferences in captive white-handed gibbons and to analyze whether they correlate with nutrient composition. Via a 2-alternative choice test, we repeatedly presented 3 male Hylobates lar with all possible binary combinations of 10 types of food that are part of their diet in captivity and found the following rank order of preference: grape > banana = fig > apple > pear > honeydew melon > carrot > tomato > cucumber > avocado. Correlational analyses revealed a highly significant positive correlation between the food preference ranking and the total carbohydrate, fructose, and glucose contents of the foods (p < 0.01, respectively). With the exception of the trace mineral selenium (p < 0.05), there was no other significant correlation with any other macro- or micronutrient. In addition, the food preferences were stable across the day because rankings obtained from tests performed at 0900, 1200, and 1500 h, respectively, did not differ significantly (p > 0.05). Our results suggest that captive white-handed gibbons are not opportunistic, but selective feeders with regard to maximizing net gain of energy because only the content of carbohydrates, but not the contents of total energy, proteins, or lipids significantly correlate with the displayed food preferences. Further, the results suggest that captive Hylobates lar, in contrast to their free-ranging conspecifics, do not display marked changes in their food selection across the day.     

Biology and behaviour of reintroduced gibbons

We observed and recorded the behaviours of gibbons undergoing rehabilitation, before and after release, to document the behavioural and social changes of gibbons in the rehabilitation program and develop criteria for determining the suitability of a pair of gibbons for release. Hylobates albibarbis were observed at the Kalaweit Gibbon Rehabilitation Project in Kalimantan, Indonesia. Data were collected on animals both pre- and post-release and on wild gibbons for comparison. Data presented here show that reintroduced gibbons are capable of surviving without human intervention. In addition, their behaviour is similar to that of wild gibbons in terms of activity budgets, position in the canopy, body posture, pair association (PA) and diet. Prior to this study, no attempt has been made to quantify the rehabilitation process for gibbons, and rehabilitation project personnel require data reporting all aspects of a release so that improvements can be made. It is important to report these data for the benefit of future releases. Criteria, based on the behaviour of wild gibbons, are proposed to assist rehabilitation centers in assessing the suitability of gibbon pairs for release.     

Calling in wild silvery gibbons (Hylobates moloch) in Java (Indonesia): behavior, phylogeny, and conservation

Hardly any behavioral data are available for the silvery gibbon (Hylobates moloch), an endangered primate that is endemic to the island of Java, Indonesia. We studied the singing behavior of the easternmost population of this species in the Dieng mountains, central Java, in 1998-1999. We aimed to document the timing of singing, quantify the amount of singing by the respective sexes, and explore the role of bioacoustics in density estimation. A total of 122 song bouts in at least 12 groups were monitored. No duet songs were heard. Most of the song bouts (91.5%) were female solo song bouts or female scream bouts. In contrast to an earlier study on the westernmost population of silvery gibbons, during which few if any male songs were heard, at least 8.5% of the song bouts in our study were male solo song bouts. They were significantly longer in duration than the female songs. All male song bouts uttered before dawn (0520 hr) were produced in a chorus fashion, with at least three individuals participating. Choruses occurred about once every 8.5 days, and lasted longer and occurred earlier than female solo song bouts. Most male songs (60%) started between 0355-0440 hr, when it was still dark. All female songs, in contrast, started after 0500 hr, and female singing activity peaked around 0600. Regular male singing, male chorusing, and regular predawn singing have not previously been reported for silvery gibbons. Similarly separated periods of male and female solo songs and the absence of duetting have been observed in Kloss's gibbons (H. klossii) on the Mentawai Islands, and may represent synapomorphies shared by both species. The pronounced individual-specific song characteristics of silvery gibbons allow accurate mapping of groups. The density of gibbons at our study site was established to be 1.9-3.7 groups/km2, corresponding to 6.7-13.1 individuals/km2. We reassess the suitability of gibbon songs as a means of estimating the density and size of gibbon populations, and discuss the proximate causes for the absence of duetting in silvery gibbons.     

Preliminary observations of wild black-crested gibbons (Hylobates concolor concolor) in Yunnan province,people's republic of China

Black-crested gibbons (Hylobates concolor concolor), which inhabit the subtropical forests of southern China and northern Vietnam, have until now never been the subject of any systematic behavioral or ecological field study. This report presents the findings of a three-month field study of wild black-crested gibbons in the Wuliang and Ailao Mountain Game Reserves in Yunnan Province, People's Republic of China. The sites visited consisted of temperate, subtropical, broadleaf evergreen forests, with trees belonging to the families Elaeocarpaceae, Fagaceae, Lauraceae, Magnoliaceae, and Theaceae. Black-crested gibbons were documented from four sites visited and were found to be polygynous, with an average group size of seven to eight animals. The family groups were observed to comprise one adult male, from one to four adult females, and numerous offspring of various ages. Both sexes of this species emit interactive songs, and various features of this loud vocal behavior were analyzed and are discussed here. Since all other gibbon species were found to inhabit only tropical rain forests and live in strictly monogamous family groups, some of the ecological and evolutionary implications of these findings are also presented.     

Predation on giant flying squirrels (Petaurista philippensis) by black crested gibbons (Nomascus concolor jingdongensis) at Mt. Wuliang, Yunnan, China

Predation on vertebrates is infrequent in gibbons. In a 14-month field study of the central Yunnan black crested gibbon (Nomascus concolor jingdongensis) at Mt. Wuliang, Yunnan, China, we observed gibbons attacking, killing and eating giant flying squirrels (Petaurista philippensis). During 845 h of observation on one study group, the gibbons attacked giant flying squirrels 11 times, and succeeded in 4 cases. Although all members of the group attempted to attack the squirrels, all four successful attacks were made by the same adult female. The victims were infants in three cases and a juvenile or sub-adult in one case. Black crested gibbons also attacked adult giant flying squirrels by grabbing their long tails and throwing them from the canopy, but they failed to catch or kill the prey in three cases observed. Passive meat sharing occurred in three out of four successful cases. Besides hunting giant flying squirrels, the black crested gibbons also ate eggs or chicks in two birds’ nests and one lizard.