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.     

Interspecific territoriality in gibbons (Hylobates lar and H. pileatus) and its effects on the dynamics of interspecies contact zones

We investigated the ecology and interspecific interactions of the two gibbon species (Hylobates lar and H. pileatus) that overlap in distribution within a narrow zone of contact in the headwaters of the Takhong River at Khao Yai National Park in central Thailand. The zone is about 10-km wide, with phenotypic hybrids comprising 6.5% of the adult population. We compared species with respect to diet, territory size, intra- and interspecific group encounters, and territory quality. The two gibbon species exploited the same types of resources within their territories despite variation in the relative abundance of food-plant species between territories. The gibbons were interspecifically territorial, and males of both species displayed aggressive behaviors at both intraspecific and interspecific territorial boundaries. There was no marked difference in the amount of overlap between territories of conspecific and heterospecific pairs of groups. Although the habitat was not homogeneous, territory quality did not vary significantly between species. The species have not diverged in habitat preference or in diet. Neither species dominated in interspecific encounters, and both were reproducing well in the contact zone. We analyzed the potential consequences of several types of interspecific interactions on individual dispersal options and on the structure of the contact zone. Interference competition through interspecific territoriality affects the dispersal of individuals into the range of the other species. In general, territorial competition coupled with limited hybridization leads to predictions of a narrow contact zone or parapatry between species; thus, behavioral and ecological interactions between species need to be considered as potential factors in explaining range borders of primate species.     

Mating system structure and population density in a polygynous lizard, Sauromalus obesus (= ater)

Increases in population density often are associated with achange in mating system structure in numerous taxa. Typically,male interactions are minimal in extremely low density populations.As density increases, males exhibit territoriality but if densitybecomes too high, the energetic cost of defending a territorywill eventually outweigh the reproductive benefits associatedwith territoriality. Consequently, males in high density populationsmay abandon territoriality and adopt dominance polygyny, lekking behavior, or scramble competition. We investigated the relationshipbetween population density and mating system structure in threepopulations of the chuckwalla, Sauromalus obesus (= ater),near Phoenix, Arizona. Densities in the Phoenix Mountains (2.7chuckwallas/ha) were lower than any population previously studied.In the Santan Mountains (10.9 chuckwallas/ha), densities weresimilar to populations studied in the Mojave Desert, and inthe South Mountains (65 chuckwallas/ha), densities were the highest yet recorded. Male mating behavior was examined by determininghome range overlap and by making direct behavioral observations.Male home range size decreased with increasing population density.There was little overlap in home ranges among males in allthree populations, whereas home ranges of males and femalesconsistently overlapped, indicating that males were strictly territorial. This conclusion was supported by behavioral observationsof interactions among individuals in a natural setting. Thenumber of females wihin male territories was correlated withfood resources (plants) in all three populations. Female homerange size appeard to be related to food resources whereasmale home ranges appeared to be related to female distribution,population density, and geology. The retention of territoriality in spite of high population densities raises new questions aboutthe relationship between density and resource defense.     

Influence of food availability and climate on behavior patterns of western black crested gibbons (Nomascus concolor) at Mt. Wuliang,Yunnan, China

Food abundance and climatic factors can significantly affect the behavior of animals and constrain their activity budgets. The population of western black crested gibbons (Nomascus concolor) in Mt. Wuliang lives in montane forest and is close to the northern extreme of the distribution for gibbons (Hylobatidae). Their habitats show remarkable seasonal variation in terms of food availability, temperature, and rainfall. To understand behavioral adaptations of western black crested gibbons to different sets of ecological conditions, we examined relationships among food availability, mean temperature, rainfall, and behavior patterns by observing two groups for 1 year each. Our results revealed that activity budget was affected by food availability and mean temperature. The gibbons spent more time eating flowers when that resource was more available and spent less time moving when fruit was more available. The gibbons spent less time feeding and more time resting, and spent less feeding time on fruit and leaves when the mean temperature was lower. These results suggest that the gibbons displayed a pronounced preference for flowers as a food resource and adopted a time minimizer strategy when high‐nutrient food items (i.e., fruit) were more available. In addition, the gibbons adopted an energy‐conserving strategy during periods of low temperature. The flexibility of behavioral patterns in responding to food availability and temperature may potentially improve the gibbons’ prospects of surviving and reproducing in a northern montane forest.     

Demography of Agile Gibbons (<Emphasis Type="Italic">Hylobates agilis</Emphasis>) in a Lowland Tropical Rain Forest of Southern Sumatra,Indonesia: Problems in Paradise

Gibbons are among the best-studied Asian primates, but few studies address their demography and life history strategies. We used annual censuses to study the demography of agile gibbons (Hylobates agilis) between 1998 and 2009 in rain forests of Bukit Barisan Selatan National Park, Indonesia. The population declined from 22 individuals (9 groups) to 14 individuals (5 groups) over the 12 yr of study. Infant survival to the juvenile age class was 33.3%, and 16.7% of infants survived to the subadult age class. The interbirth interval was 3.83 ± 1.15 yr and birth rate was 0.22–0.28 infants female^–1 yr^–1. Two groups colonized the study area but subsequently disappeared. We documented 7 immigrations, 17 disappearances, and ≥10 transients in the population. Compared to lar gibbons (Hylobates lar) and Bornean white-bearded gibbons (Hylobates albibarbis), Way Canguk’s agile gibbon population is characterized by slow reproduction, low survival, and high group turnover. We hypothesize that, although the habitat is high in fruit resources, agile gibbons may be displaced or excluded from the best fruit resources by larger and more numerous competitors, incurring costs of decreased opportunities to forage and increased travel, and leading to higher mortality for young agile gibbons. The reproductive potential of this agile gibbon population is insufficient to compensate for high mortality, and the population is unlikely to persist without immigration from outside the area. Given the agile gibbons’ endangered status and limited capacity to respond demographically to change, it is likely that intensive management interventions will be required to conserve this species.     

Population dynamics of Arashiyama west Japanese macaques

Demographic data have been collected on the Arashiyama Japanese macaque population from 1954 until the present, through the fissioning of the original group into two parts in 1966, and through the translocation of one of the two groups to Texas in 1972. Population dynamics are reported for the Arashiyama West group in Texas during 1972 to 1979 and then compared to data from Japan. After a short period of adjustment for the translocated group, during which time natality rates were relatively low and mortality rates were relatively high, many aspects of population structure, birth rates, and survivorship showed trends similar to those observed in Japan. This suggests that both long-term homeostatic processes and shorter-term responses to environmental fluctuations are significant to the study of nonhuman primate demography.     

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.     

Fecal Glucocorticoid Measurements and Their Relation to Rearing,Behavior, and Environmental Factors in the Population of Pileated Gibbons (<Emphasis Type="Italic">Hylobates pileatus</Emphasis>) Held in European Zoos

Pileated gibbons (Hylobates pileatus) are rated as endangered according to the International Union of Conservation of Nature (IUCN) Red List. The captive population suffers from poor breeding success and is threatened to become overaged. Although several factors are likely to contribute to the poor breeding success, one in particular may be chronic stress associated with prolonged periods of high glucocorticoid (GC) output. We investigated fecal GC levels of pileated gibbons (Hylobates pileatus) and their relationship to specific life-history variables and environmental factors. After validation of an enzyme immunoassay for the measurement of 5-reduced 3α,11β-dihydroxy cortisol metabolites to assess GC output reliably in pileated gibbons, we collected fecal samples over several days from all 36 European adult pileated gibbons located in 11 institutions and compared GC levels to intrinsic individual parameters, husbandry, behavior, and breeding history. Age, sex, and origin (wild vs. captive born) had no effect on GC levels. However, unnaturally reared gibbons had higher GC levels and showed more behavioral abnormalities than parent-reared individuals. Further, nonreproducing gibbons living in a pair without infants had higher GC concentrations than gibbons living in a family, bachelor group, or as singletons. With respect to environmental factors, a large size of the inside enclosure and the existence of visual protection from visitors was associated with lower fecal GC output. The data indicate that rearing and housing conditions appear to correlate to GC levels in pileated gibbons housed under captive conditions. It is hoped this knowledge will support the future management of the species in captivity and thus lead to a more successful breeding of this endangered primate.     

Are Montane Forests Demographic Sinks for Bornean White‐bearded Gibbons Hylobates albibarbis?

I estimated habitat-specific population densities for a population of Bornean white-bearded gibbons Hylobates albibarbis inhabiting seven distinct forest types at Gunung Palung National Park, West Kalimantan, Indonesia. Population densities in montane forests (0.44 individuals/km²) were almost ten times lower than those in the next best habitat (upland granite forest; 4.2 individuals/km²) and far lower than those in lowland forest types. Demographic data on 33 gibbon groups living across the seven forest types showed that reproduction was substantially depressed in montane forests compared to high-quality lowland habitats. A simple model suggests that montane forests are demographic sinks for gibbons at Gunung Palung. Follow-up data from observations of montane groups 5 yr after the initial observation period support this result. As high-quality lowland forests (source habitat for gibbons) are being disproportionately lost in and around Gunung Palung National Park due to illegal logging and conversion to oil palm plantations, an increasing percentage of the remaining forest in the park comprises sink habitat for gibbons. This result has disquieting implications for the long-term viability of gibbon populations at Gunung Palung. In addition, as montane forests are generally low-quality habitat for most rainforest vertebrates, and since lowland forests are being lost at alarming rates across the tropics, source-sink population dynamics similar to those I describe here may characterize populations of other tropical vertebrate species.     

Combining species distribution modeling and distance sampling to assess wildlife population size: A case study with the northern yellow-cheeked gibbon (Nomascus annamensis)

Population size and distribution data for wildlife species play an important role in conservation and management, especially for endangered species. However, scientists seriously lack data on the population status of many species. The northern yellow-cheeked gibbon (Nomascus annamensis) is found in southern Lao PDR, central Vietnam, and northeastern Cambodia. The population of the species has significantly declined due to hunting, habitat loss, and the wildlife trade. To examine the population size and distribution of N. annamensis, we conducted a field survey in Song Thanh Nature Reserve, Quang Nam Province, central Vietnam from February to April 2019 using the audio point count method. We combined Distance Sampling and Ecological Niche Modeling to estimate the population of the gibbons. Results showed that the total suitable area for the gibbons was about 302.32 km², with the two most important variables of the habitat model being the distance-to-villages and forest type. We detected 36 gibbon groups through field surveys and estimated 443 (95% CI, 278–707) gibbon groups in Song Thanh Nature Reserve. Our results indicate that the gibbon population in Song Thanh Nature Reserve is the largest known population of N. annamensis in Vietnam. In addition, our study was the first to combine species distribution modeling with distance sampling to estimate gibbon density and population size. This approach might be useful in surveying and monitoring gibbon populations because it takes imperfect detection probability into account in estimating gibbon population density while estimating the area of potential habitat using environmental variables.     

Experimental field studies of Asian ape social systems

The Asian apes, orangutans and gibbons, possess unusual social systems among anthropoid primates. Social groups of gibbons consist of mated adult pairs and their offspring; mature orangutans are primarily solitary. Recent experimental field research has begun to yield insights into the behavioral mechanisms employed by these animals to maintain their characteristic patterns of social dispersion. While spatial separation between female orangutans appears to be maintained passively, aggression, which is manifest during direct encounters and long-distance vocal interactions, mediates male asociality. Male-male aggression is the result of intense intrasexual competition occurring between animals for mating access to females. To reduce intrasexual competition, male orangutans have adopted alternative mating tactics. In contrast to female orangutans, female gibbons show marked agonistic tendencies toward conspecifics. Female territoriality contributes to preventing males from becoming polygynous. Male gibbons, restricted to monogamous relationships, attempt to ensure their paternity through intrasexual aggression. These observations suggest that the spatial dispersion of females constrains male mating options in both species. However, variations between orangutan and gibbon social systems can be understood as consequences of the temporal dispersion of sexually receptive females. The temporal clumping of females, due to relatively high operational sex ratios, limits the ability of male gibbons to acquire multiple mates. Conversely, an extremely low operational sex ratio in orangutans creates a strong selection pressure for intrasexual competition and polygamous mating. These considerations provide a novel framework for interpreting the social systems of the African apes.     

An ecological study of troop fissions of Japanese monkeys (Macaca fuscata yakui) on Yakushima Island,Japan

Wild, habituated, Japanese monkeys were observed from 1975 to 1979 on Yakushima Island, Southern Japan. The monkey troops had a continuous distribution in a warm temperate forest. Demographic data on local populations was collected. The population density was 33 animals/km². The growth rate of the studied troop was 3.0% per year. A significant correlation between home range areas (R) and troop size (P) was found (r=0.955,p<0.005), using anR-P equation,R=1.84P. One troop split into three troops through two successive fissions. Twenty-one intertroop encounters were observed. Five types of encounters were distinguished. The encounters were apparently territorial defence. Increases in birth rate and socionomic sex ratio after the fissions were prominent. The following four factors had a direct effect upon the dispersion of the troops after fission: (1) dominance relation between the fission troops; (2) social pressure of the neighbors; (3) troop's attachment to its home range; and (4) structure of the environment. The home range of Japanese monkeys is a territory, and territoriality is a population regulating mechanism which serves to reduce competition for food.     

Ecology and behavior of wild black-crested gibbons (Hylobates concolor) in China with a reconsideration of evidence for polygyny

The black-crested gibbon,Hylobates concolor, is one of the few species of gibbons that has not yet been the subject of a long term field study. Field observations in the Ai Lao and Wu Liang Mountains of Yunnan Province, China indicate that in this area the habitat and ecology of this species differ markedly from those of other gibbons that have been studied to date. These differences are correlated with some behavioral differences. In particular, these gibbons apparently have greater day ranges than other gibbons. It has also been suggested that this species lives in polygynous groups. To demonstrate this requires observation of groups with two or more females with young. Our own observations and those from other recent studies suggest that there are alternative explanations consistent with available data.     

Density and population estimate of gibbons (Hylobates albibarbis) in the Sabangau catchment, Central Kalimantan, Indonesia

We demonstrate that although auditory sampling is a useful tool, this method alone will not provide a truly accurate indication of population size, density and distribution of gibbons in an area. If auditory sampling alone is employed, we show that data collection must take place over a sufficient period to account for variation in calling patterns across seasons. The population of Hylobates albibarbis in the Sabangau catchment, Central Kalimantan, Indonesia, was surveyed from July to December 2005 using methods established previously. In addition, auditory sampling was complemented by detailed behavioural data on six habituated groups within the study area. Here we compare results from this study to those of a 1-month study conducted in 2004. The total population of the Sabangau catchment is estimated to be about in the tens of thousands, though numbers, distribution and density for the different forest subtypes vary considerably. We propose that future density surveys of gibbons must include data from all forest subtypes where gibbons are found and that extrapolating from one forest subtype is likely to yield inaccurate density and population estimates. We also propose that auditory census be carried out by using at least three listening posts (LP) in order to increase the area sampled and the chances of hearing groups. Our results suggest that the Sabangau catchment contains one of the largest remaining contiguous populations of Bornean agile gibbon.     

Demographic analyses of the dusky shark, <Emphasis Type="Italic">Carcharhinus obscurus,</Emphasis> in the Northwest Atlantic incorporating hooking mortality estimates and revised reproductive parameters

Demographic analyses were conducted for the dusky shark, Carcharhinus obscurus, in the Northwest Atlantic. Fishery-independent, fishery-dependent, and previously published data were used to estimate age specific by-catch mortality and reproductive cycle for incorporation into the analyses. Correspondence analyses were conducted for viability (dead or alive), shark length, month, soaktime, and region. Dead sharks were associated with small size, long soak times, and the spring season, while live sharks were associated with large size and shorter soak times. Data on reproduction suggested a 3-year reproductive cycle consisting of a 2-year gestation period and a 1-year resting period. Litter sizes ranged from 3–12 embryos. Decreasing age-specific fishing mortality and a 3-year reproductive cycle were used in age structured life tables which incorporated stochasticity in life history parameters through Monte Carlo simulation to estimate annual population growth. Demographic analyses showed that the dusky shark population would decline even at low levels of fishing mortality despite low natural mortality for neonates. The demographics of this species make it among the most vulnerable to excessive fishing mortality, and suggest stringent regulatory measures may be required to recover collapsed populations.     

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.     

Census and survey of wild black-crested gibbons (Hylobates concolor concolor) in Yunnan Province, People's Republic of China

Black-crested gibbons (Hylobates concolor concolor) inhabit the subtropical forests of Southern China and Northern Vietnam, and have never previously been the subject of any systematic behavioral or ecological study. This report presents the findings of a three-month census and survey of black-crested gibbons in the Wuliang and Ailao Mountain Game Reserves in Yunnan province, China. The censusing methods used here were similar to those techniques used during other census studies of gibbons. The sites visited were subtropical broadleaf evergreen forests, with trees belonging to the families Fagaceae, Theaceae, Magnoliaceae, Lauraceae and Elaeocarpaceae. A total of 23-25 groups of black-crested gibbons were documented from 4 sites visited, and a group density estimate from all sites averaged 0.82 groups/km2. These gibbons were found to be polygynous with an average family group size of 7-8 animals, comprising 1 adult male, 1-4 adult females and numerous offspring of various ages. Some of the ecological and evolutionary implications of these findings are also presented.     

Social behavior adaptations of gibbon (Hylobates lar) in a controlled environment

A population of six adolescent—adult gibbons (three males and three females) were observed in a controlled environment in order to assess any social behavior adaptations related to forced aggregation and to a physical environment which differed from the native habitat in that there were no predators, no food scarcity and the geographic range was limited to 1.5 acres. Data on location, size and composition of naturally formed social groupings revealed that certain gibbons avoided social contact with certain others. This observation is discussed in relation toChance's (1967) concept of “attention structure.” It is viewed as representative of a dominance hierarchy serving gibbons as a social behavior adaptation when forced to aggregate.     

Comparisons of Suspensory Behaviors Among <Emphasis Type="Italic">Pygathrix cinerea</Emphasis>, <Emphasis Type="Italic">P. nemaeus</Emphasis>, and <Emphasis Type="Italic">Nomascus leucogenys</Emphasis> in Cuc Phuong National Park,Vietnam

In our study at the Endangered Primate Rescue Center of Cuc Phuong National Park, Vietnam, we aimed first to assemble a positional behavioral profile of captive gray-shanked (Pygathrix cinerea) and red-shanked (P. nemaeus) doucs that relates to the use of forelimb suspensory postures and arm-swinging locomotion. The profile is of interest because researchers have documented that red-shanked doucs more frequently use suspensory postures and locomotions than other colobines do. We confirmed that red-shanked doucs commonly use suspensory positional behaviors and also that gray-shanked doucs use suspensory behaviors at similar or even higher frequencies than those of red-shanked doucs. Our second goal was to assemble a preliminary kinematic profile of suspensory locomotion in Pygathrix within the context of the arm-swinging locomotion exhibited by northern white-cheeked gibbons, Nomascus leucogenys. Mean forelimb angles at initial contact and release of arm-swinging behaviors were remarkably consistent among gibbons and doucs despite the fact that gibbons typically used more continuous brachiation. Doucs also exhibit a greater range of forelimb angles than gibbons do. In addition, trunk orientation tends to be less vertical at initial contact for doucs than for gibbons, perhaps owing to the frequent use of quadrupedal sequences directly before or after forelimb suspension. Our behavioral and kinematic analyses add to the emerging realization that Pygathrix is capable of, and frequently expresses, a range of suspensory positional behaviors, including brachiation.     

Life history influences how fire affects genetic diversity in two lizard species

‘Fire mosaics’ are often maintained in landscapes to promote successional diversity in vegetation with little understanding of how this will affect ecological processes in animal populations such as dispersal, social organization and re‐establishment. To investigate these processes, we conducted a replicated, spatiotemporal landscape genetics study of two Australian woodland lizard species [Amphibolurus norrisi (Agamidae) and Ctenotus atlas (Scincidae)]. Agamids have a more complex social and territory structure than skinks, so fire might have a greater impact on their population structure and thus genetic diversity. Genetic diversity increased with time since fire in C. atlas and decreased with time since fire in A. norrisi. For C. atlas, this might reflect its increasing population size after fire, but we could not detect increased gene flow that would reduce the loss of genetic diversity through genetic drift. Using landscape resistance analyses, we found no evidence that postfire habitat succession or topography affected gene flow in either species and we were unable to distinguish between survival and immigration as modes of postfire re‐establishment. In A. norrisi, we detected female‐biased dispersal, likely reflecting its territorial social structure and polygynous mating system. The increased genetic diversity in A. norrisi in recently burnt habitat might reflect a temporary disruption of its territoriality and increased male dispersal, a hypothesis that was supported with a simulation experiment. Our results suggest that the effects of disturbance on genetic diversity will be stronger for species with territorial social organization.