Group Composition and Monandry in Grizzled Langurs, Presbytis comata, on Java.

Grizzled langurs, Presbytis comata, a largely sexually monomorphic species, are reported to occur in populations where either the majority of groups comprise 1 adult male with 1 adult female, or where groups comprise 1 adult male with multiple females. As such, they may have a monandrous mating system. I investigated whether 1-male/1-female groups indeed form a significant part of the species' social system, and whether habitat variation (forest fragment size, distance to the forest edge, altitude) affects social organization. I found the species from sea level to 2,565 m above sea level in groups from 1 to 13 individuals. I recorded mostly 1-male/multifemale groups with offspring or, alternatively, all-male groups. Two out of 55 groups comprised 1-male/1-female groups with offspring. Group size was negatively correlated with altitude and forest fragment size, and positively correlated with increasing distance from the forest edge. Altitudinal variation in group sizes was driven mainly by fewer adult females being present in groups at higher elevations; the number of adult males (almost invariably 1), subadults, juveniles, and infants, as well as the infant/adult female ratio, showed little altitudinal variation. One-male/1-female groups have been recorded repeatedly over a 25-year period in a high-altitude population on Mt. Patuha, West Java, but even here, on average, three fifths of the groups comprise 1 adult male with multiple females. At high-altitude sites, P. comata may indeed have a monandrous mating system, but at lower elevations it seems similar to that of other Presbytis langurs.

Maternal kin bias in affiliative behavior among wild adult female blue monkeys.

Kin-biased cooperative and affiliative behavior is widespread in social mammals and is expected to increase fitness. However, despite evolutionary benefits of cooperating with relatives, demographic circumstances may influence the strength of kin bias. We studied the relationship between maternal kinship and affiliative behavior among 78 wild adult female blue monkeys (Cercopithecus mitis) from 8 groups monitored for 1-5 years. We compared behavior and kinship matrices, controlling for rank differences. Using multivariate models, we examined effects of demographic variables on the extent to which females groomed disproportionately with close adult female kin. Female blue monkeys, like other cercopithecine primates, generally preferred closer maternal kin for grooming and spatial association, although there was also substantial variation. Kin bias was weakest for association (at 7 m) while feeding, intermediate for closer (1 m) association while resting, and most intense for grooming. Grooming kin bias was stronger when a female had more very close relatives (either her mother or daughters), when her group contained more adult females, when she groomed with a lower percentage of group-mates, and when she had fewer total kin. Dominance rank did not predict variation in kin bias. Females generally groomed with all kin, but in larger groups they increased the number of unrelated grooming partners and total grooming time. The increased kin bias intensity in larger groups resulted from the addition of unrelated partners with whom grooming occurred less often than with kin, rather than from time constraints that drove females to select kin more strongly. In natural-sized groups, it may be common that females groom with all their adult female kin, which are present in limited numbers. The addition of grooming partners in larger groups may benefit female blue monkeys who rely on collective action in territorial defense; group-wide cooperation may thus influence grooming decisions in this species.

Social behavior, foraging strategies, and fecal glucocorticoids in female blue monkeys (Cercopithecus mitis): potential fitness benefits of high rank in a forest guenon.

Socioecological theory predicts that aggressive feeding competition is associated with linear dominance hierarchies and reproductive advantages for high-ranking females. Female blue monkeys contest fruits and have a linear dominance hierarchy, yet previous research has shown no evidence that high-ranking females benefit from greater feeding success or fertility. Here, we assess whether individuals differ in fecal glucocorticoid (fGC) excretion and examine proximate determinants of such differences to infer potential fitness correlates of rank, using data collected from two study groups in the Kakamega Forest, Kenya. We found that higher ranking females had preferential access to fruits in both groups, although the behavioral mechanisms leading to this effect varied between groups. Despite a consistent rank difference in feeding on fruits, an overall rank effect on fGCs emerged in only one group; females of this group spent comparatively more time feeding on fruits, fruits accounted for a greater proportion of the diet, and females engaged in more frequent food-related agonism. In addition, more females in this group were lactating during a period of low fruit availability, when rank effects on fGCs were particularly strong. Regardless of fruit availability, among lactating females of both groups higher rank was associated with lower fGC levels, indicating lower energetic stress in higher ranking females when energy demands were particularly high. Individual rates of agonism, a potential psychological stressor, were unrelated to fGCs at all times. After we accounted for rates of agonism and feeding on fruits, females of one group who groomed others more had lower fGCs, suggesting that variable social coping behavior can contribute to fGC variation in some groups. This study provides the first empirical evidence that high-ranking female blue monkeys may obtain fitness benefits from their social status, by gaining priority of access to fruits during critical times in the reproductive cycle.

Optimal foraging on the roof of the world: Himalayan langurs and the classical prey model.

Optimal foraging theory has only been sporadically applied to nonhuman primates. The classical prey model, modified for patch choice, predicts a sliding "profitability threshold" for dropping patch types from the diet, preference for profitable foods, dietary niche breadth reduction as encounter rates increase, and that exploitation of a patch type is unrelated to its own abundance. We present results from a 1-year study testing these predictions with Himalayan langurs (Semnopithecus entellus) at Langtang National Park, Nepal. Behavioral data included continuous recording of feeding bouts and between-patch travel times. Encounter rates were estimated for 55 food types, which were analyzed for crude protein, lipid, free simple sugar, and fibers. Patch types were entered into the prey model algorithm for eight seasonal time periods and differing age-sex classes and nutritional currencies. Although the model consistently underestimated diet breadth, the majority of nonpredicted patch types represented rare foods. Profitability was positively related to annual/seasonal dietary contribution by organic matter estimates, whereas time estimates provided weaker relationships. Patch types utilized did not decrease with increasing encounter rates involving profitable foods, although low-ranking foods available year-round were taken predominantly when high-ranking foods were scarce. High-ranking foods were taken in close relation to encounter rates, while low-ranking foods were not. The utilization of an energetic currency generally resulted in closest conformation to model predictions, and it performed best when assumptions were most closely approximated. These results suggest that even simple models from foraging theory can provide a useful framework for the study of primate feeding behavior.

Language evolution: the origin of meaning in primates.

Research on alarm calls has yielded rare glimpses into the minds of our closest relatives. A new study suggests that primates monitor the effect alarm calls have on others.

Revisiting play elements and self-handicapping in play: a comparative ethogram of five Old World monkey species.

Play behavior has been viewed as a mixture of elements drawn from "serious" behavior, interspersed by ritualized play signals. Two other types of play behaviors have been overlooked: patterns that are dissimilar from any serious behavior and patterns with self-handicapping character, that is, those that put the animal into unnecessary disadvantageous positions or situations. Here the authors show that these 2 types of patterns can constitute a major part of play repertoire. From our own videorecordings and observations, we constructed play ethograms of 5 monkey species (Semnopithecus entellus, Erythrocebus patas, Chlorocebus pygerythrus, Cercopithecus neglectus, and Cercopithecus diana). The authors evaluated the self-handicapping character of each pattern and in Hanuman langurs also the (dis)similarity to serious behavior. Of the 74 patterns in the 5 species, 33 (45%) were judged to have a self-handicapping character. Of 48 patterns observed in langurs, 16 (33%) were totally dissimilar to any serious langur behavior known to us. The authors discuss the possibility that the different types of play elements may have different functions in play.

Personality in free-ranging Hanuman langur (Semnopithecus entellus) males: subjective ratings and recorded behavior.

The authors obtained behavioral observations and personality ratings for 27 free-ranging Hanuman langur males. Subjects were rated using a questionnaire based on the human Five-Factor Model (FFM). Behavioral observations were taken over 5 months using an ethogram that included 50 behaviors. Principal Component Analysis (PCA) of ratings revealed Agreeableness_(R), Confidence_(R), and Extraversion_(R) components. Each personality dimension was associated with a unique set of observed behaviors. PCA of 36 behavioral indices revealed Dominance_(B), Involvement_(B), and Activity_(B) components. Bivariate correlations showed that Agreeableness_(R) was negatively correlated with Dominance_(B); Confidence_(R) was positively correlated with Dominance_(B) and Involvement_(B) but negatively correlated with Activity_(B); and Extraversion_(R) was positively correlated with Activity_(B). Dominance rank was positively correlated with Confidence_(R) and Dominance_(B) but negatively correlated with Agreeableness_(R) and Activity_(B). These results highlight the comparability of behavioral coding and personality ratings and suggest that some aspects of personality structure were present in the common ancestor of Old World monkeys.

Ranging behavior of the François' langur (Trachypithecus francoisi) in the Fusui Nature Reserve, China.

Data on activity budgets and ranging patterns were collected from March to December 2001 for one group of François' langurs (Trachypithecus francoisi) inhabiting a forested part of the Fusui Nature Reserve, Guangxi province, China. Our results indicate that the total size of the home range of the study group during the study period was 19 ha. The majority of their activities (52%) occurred within a small area, 22%, of their home range, and was concentrated in or near quadrats containing their sleeping sites, which may reduce the time and energetic cost of travel. The extent of the ranging behavior varied between months, with the smallest, 7 ha, recorded in July and the largest, 13.5 ha, in November. There was no significant difference between seasons. The monthly mean daily path lengths varied from 341 to 577 m. The daily path lengths showed significant seasonal changes: the path lengths were longer during the dry season than in the rainy season, which may be related to the scarcity of preferred food resources during the dry season.

Daytime behavior of Pteropus vampyrus in a natural habitat: the driver of viral transmission.

Flying foxes, the genus Pteropus, are considered viral reservoirs. Their colonial nature and long flight capability enhance their ability to spread viruses quickly. To understand how the viral transmission occurs between flying foxes and other animals, we investigated daytime behavior of the large flying fox (Pteropus vampyrus) in the Leuweung Sancang conservation area, Indonesia, by using instantaneous scan sampling and all-occurrence focal sampling. The data were obtained from 0700 to 1700 hr, during May 11-25, 2016. Almost half of the flying foxes (46.9 ± 10.6% of all recorded bats) were awake and showed various levels of activity during daytime. The potential behaviors driving disease transmission, such as self-grooming, mating/courtship and aggression, peaked in the early morning. Males were more active and spent more time on sexual activities than females. There was no significant difference in time spent for negative social behaviors between sexes. Positive social behaviors, especially maternal cares, were performed only by females. Sexual activities and negative/positive social behaviors enable fluid exchange between bats and thus facilitate intraspecies transmission. Conflicts for living space between the flying foxes and the ebony leaf monkey (Trachypithecus auratus) were observed, and this caused daily roosting shifts of flying foxes. The ecological interactions between bats and other wildlife increase the risk of interspecies infection. This study provides the details of the flying fox's behavior and its interaction with other wildlife in South-East Asia that may help explain how pathogen spillover occurs in the wild.

Male monkeys remember which group members have given alarm calls.

Primates give alarm calls in response to the presence of predators. In some species, such as the Thomas langur (Presbytis thomasi), males only emit alarm calls if there is an audience. An unanswered question is whether the audience's behaviour influences how long the male will continue his alarm calling. We tested three hypotheses that might explain the alarm calling duration of male Thomas langurs: the fatigue, group size and group member behaviour hypotheses. Fatigue and group size did not influence male alarm calling duration. We found that males only ceased calling shortly after all individuals in his group had given at least one alarm call. This shows that males keep track of and thus remember which group members have called.