Variation in monthly and seasonal elevation use impacts behavioral and dietary flexibility in Rhinopithecus bieti.

Black-and-white snub-nosed monkeys (Rhinopithecus bieti) rely on behavioral and dietary flexibility to survive in temperate latitudes at high-elevation habitats characterized by climate and resource seasonality. However, little is known about how elevation influences their behavioral and dietary flexibility at monthly or seasonal scales. We studied an isolated R. bieti population at Mt. Lasha in the Yunling Provincial Nature Reserve, Yunnan, China, between May 2008 and August 2016 to assess the impacts of elevation on feeding behavior and diet. Across our sample, R. bieti occupied elevations between 3031 and 3637 m above mean sea level (amsl), with a 315.1 m amsl range across months and a 247.3 m amsl range across seasons. Contrary to expectations, individuals spent less time feeding when ranging across higher elevations. Lichen consumption correlated with elevation use across months and seasons, with individuals spending more time feeding on this important resource at higher elevations. Leaf consumption only correlated with elevation use during the spring. Our results suggest that R. bieti do not maximize their food intake at higher elevations and that monthly and seasonal changes in lichen and leaf consumption largely explain variation in elevation use. These findings shed light on the responses of R. bieti to environmental change and offer insight into strategies for conserving their habitats in the face of anthropogenic disturbance.

Enamel chipping in Taï Forest cercopithecids: Implications for diet reconstruction in paleoanthropological contexts.

Antemortem enamel chipping in living and fossil primates is often interpreted as evidence of hard-object feeding (i.e., 'durophagy'). Laboratory analyses of tooth fracture have modeled the theoretical diets and loading conditions that may produce such chips. Previous chipping studies of nonhuman primates tend to combine populations into species samples, despite the fact that species can vary significantly in diet across their ranges. Chipping is yet to be analyzed across population-specific species samples for which long-term dietary data are available. Here, we test the association between enamel chipping and diet in a community of cercopithecid primates inhabiting the Taï Forest, Ivory Coast. We examined fourth premolars and first molars (n = 867) from naturally deceased specimens of Cercocebus atys, Colobus polykomos, Piliocolobus badius,Procolobus verus, and three species of Cercopithecus. We found little support for a predictive relationship between enamel chipping and diet across the entire Taï monkey community. Cercocebus atys, a dedicated hard-object feeder, exhibited the highest frequencies of (1) chipped teeth and (2) chips of large size; however, the other monkey with a significant degree of granivory, Co. polykomos, exhibited the lowest chip frequency. In addition, primates with little evidence of mechanically challenging or hard-food diets-such as Cercopithecus spp., Pi. badius, and Pr. verus-evinced higher chipping frequencies than expected. The equivocal and stochastic nature of enamel chipping in the Taï monkeys suggests nondietary factors contribute significantly to chipping. A negative association between canopy preference and chipping suggests a role of exogenous particles in chip formation, whereby taxa foraging closer to the forest floor encounter more errant particulates during feeding than species foraging in higher strata. We conclude that current enamel chipping models may provide insight into the diets of fossil primates, but only in cases of extreme durophagy. Given the role of nondietary factors in chip formation, our ability to reliably reconstruct a range of diets from a gradient of chipping in fossil taxa is likely weak.

Male social rank and food competition in a primate multi-level society.

OBJECTIVES: Social animals often have dominance hierarchies, with high rank conferring preferential access to resources. In primates, competition among males is often assumed to occur predominantly over reproductive opportunities. However, competition for food may occur during food shortages, such as in temperate species during winter. Higher-ranked males may thus gain preferential access to high-profitability food, which would enable them to spend longer engaged in activities other than feeding. MATERIALS AND METHODS: We performed a field experiment with a breeding band of golden snub-nosed monkeys, a species that lives in a multi-level society in high-altitude forests in central China. We provisioned monkey's high-profitability food during winter when natural foods are limited, and then recorded the times individual adult males spent engaged in different behaviors. RESULTS: Higher-ranking males spent less time feeding overall and fed on provisioned foods at a higher rate than lower-ranking males. Higher-ranking males therefore had more time to spend on alternative behaviors. We found no significant difference according to rank in times spent moving or resting. However, high-ranking males spend significantly longer on affiliative behaviors with other members of their social sub-units, especially grooming and being groomed, behaviors known to promote social cohesion in primates. DISCUSSION: We show that preferential access to high-profitability foods likely relaxes time-budget constraints to higher-ranking males. High-ranking males thus spend more time on non-feeding activities, especially grooming, which may enhance social cohesion within their social sub-unit. We discuss the potential direct and indirect benefits to high-ranking males associated with preferential access to high-value food during winter.

Eye Preference for Emotional Stimuli in Sichuan Snub-Nosed Monkeys.

This study presents the first evidence of effects of applying both positive and negative stimuli simultaneously on visual laterality in Old World monkeys. Thirteen captive individuals of Sichuan snub-nosed monkeys (Rhinopithecus roxellana) were chosen as focal subjects in the monocular box task. In total, 4 emotional categories (the preferred, the novel, the neutral, and the fearful) of visual stimuli were applied, and eye preference was recorded when individuals looked at each stimulus through an observation hole in the box. We found evidence of visual laterality at the individual level, but not at the group level for each stimulus. For the preferred stimulus, 9 individuals showed significant right-eye preference while 4 individuals showed significant left-eye preference. For the other 3 stimuli, 7 individuals displayed significant right-eye preference while 6 individuals displayed significant left-eye preference. Totally, 11 of 13 individuals showed consistency in the visual laterality direction (7 right-eye preference and 4 left-eye preference) across the 4 stimuli. The remaining 2 individuals displayed right-eye preference for the preferred stimulus while they showed left-eye preference for the other 3 stimuli. There was no significant difference among various stimuli regarding the direction of visual laterality. However, there was a significant difference in the strength of visual laterality among various stimulus categories. The strength of visual laterality for the preferred stimulus was significantly lower than that for the other 3 stimuli. The strength of visual laterality for the fearful stimulus was significantly higher than that for the novel stimulus and the neutral stimulus. Furthermore, the looking duration for the preferred stimulus was significantly higher than that for the other 3 stimuli. The looking duration for the novel stimulus was significantly higher than that for the neutral stimulus and the fearful stimulus. The looking duration for the neutral stimulus was significantly higher than that for the fearful stimulus. Our findings indicate emotional valence of stimuli significantly influence eye looking duration and the strength of visual laterality but not for the direction of visual laterality in this species. Taken together, emotional valence of stimuli plays an important role in the eye use of R. roxellana.

Effect of Group Size on Time Budgets and Ranging Behavior of White-Headed Langurs in Limestone Forest, Southwest China.

The ecological constraints model is well supported by data from most frugivorous primates; however, the prediction power of the model is weak for folivorous primates. From September 2016 to August 2017, we collected comparative data on time budgets, daily path lengths and diets of four groups of white-headed langurs (Trachypithecus leucocephalus), including two large groups (G-DS and G-ZWY) and two small groups (G-LZ and G-NN) in Chongzuo White-Headed Langur National Nature Reserve, Guangxi Province, Southwest China. The aim was to obtain evidence of foraging competition and to test the ecological constraints model on this highly folivorous primate in its karst habitat. The results showed that langurs in the larger groups spent more time traveling, less time resting, and had a longer average daily path length than those in the small groups. Diet composition and dietary diversity were not significantly different between the large and small groups. Our study demonstrates that langurs from large groups suffer scramble competition in limestone forests and supports the validity of the ecological constraints model for folivores.

Comparison of Activity Budgets, Diet, and Habitat Utilization between Provisioned and Wild Groups of the François' Langur (Trachypithecus francoisi) in Mayanghe National Nature Reserve, China.

The endangered François' langur (Trachypithecus francoisi) is only found from southern China to northeastern Vietnam. This study evaluates behavioral differences between provisioned (Ts, 7 individuals) and wild (Tn, 13 individuals) family groups in Mayanghe National Nature Reserve, China. Scan sampling every 5 min for 12 h daily between the months of March 2013 and February 2014 was used to record behaviors in seven categories. Tn exhibited three main feeding periods daily, whereas Ts showed more variable behavioral patterns that relied on provisions. The Tn alpha male showed twice the amount of guarding behavior compared with the Ts alpha male. The proportion of each habitat type utilized differed significantly between Tn and Ts (χ2[4] = 17,131.4, p < 0.01). Additionally, dietary diversity differed between the two groups: Tn fed on 61 plant species, whereas Ts fed on 43 plant species. Tn rarely ate sweet potato and corn other than discarded remnants of human food (0.2% of their total food sources), whereas these foods represented 21.8% of Ts food sources. We compared data for the two troops to understand this species' activities under human interference, in particular to determine whether provisioning is a suitable strategy for their conservation.

Reproductive Parameters of Wild Rhinopithecus bieti.

Animal life activities are rhythmic and affected by seasonal periodicity. Based on 9 years of observations, we estimated the reproductive parameters of a wild, but provisioned Yunnan snub-nosed monkey (Rhinopithecus bieti) group at Xiangguqing in Baimaxueshan National Nature Reserve, Yunnan Province, China. We observed 84 infants (43 males and 41 females) from 41 females between 2010 and 2018. We found the birth sex ratio was 1:1, the female age at first birth was 6.13 years and infant mortality was about 15.5%. Nine years of data showed that the maximum birth season lasted 126 days, and the average length per year was 98.57 ± 18.71 days. R. bieti,characterized by strictly seasonal reproduction, started giving birth on February 1, and this ended on June 7, with a peak reached from March 4 to March 11 (10th week). The mean birth date was March 20 (79.21 ± 29.54 days), and the median birth date was March 11 (71st day). The mean interbirth interval (IBI) was approximately 2 years, and the IBIs among females whose infants had survived for 1 year were found to be significantly longer than those found in females who lost their infant within 1 year. Comparing the reproduction parameters among Asian and African colobines, we concluded that Asian and African colobines tend to have an IBI of 2 years or more, females tend to give birth at the age of 5-6 years, Rhinopithecus species had a strict seasonal reproductive pattern concentrated in February to April. Seasonal changes in food resources and climatic factors may be the main reasons for the variation in reproductive parameters in intraspecific and interspecific comparisons of Asian and African colobines.

Positional Behaviours of François' Langur (Trachypithecus francoisi) in the Limestone Forest of Nonggang, Guangxi, South-West China.

Primates' positional behaviours provide information for understanding relationships among morphology, ecology and behavioural flexibility. From September 2005 to August 2006, we collected data via instantaneous scan sampling on positional behaviours in François' langurs at Nonggang Nature Reserve, south-west China. We predicted that these langurs would use (1) leaping as the predominant locomotor mode and (2) bipedal standing more frequently in the rainy season than in the dry season. Our result showed that leaping was the dominant locomotor mode (38.38%), followed by -quadrupedal walking (31.2%), vertical climbing (25.1%) and quadrupedal running (5.3%). The ground was the most frequently used stratum during movement (33.4%). Most locomotion through trees occurred on small- (48.7%) and medium-sized (47.6%) substrates. Locomotor mode, forest stratum use and substrate use during movement did not vary seasonally. When stationary, sitting was the most common posture (92.1%), followed by bipedal standing (3.7%), lying (3.5%), quadrupedal standing (0.6%), suspending (0.2%) and back-lying (<0.1%). Posture varied significantly with the season. During resting, langurs used sitting and bipedal standing more frequently in the dry season, while adopting lying more frequently in the rainy season. During feeding, sitting was adopted more frequently in the rainy than in the dry season, whereas bipedal standing was used more frequently in the dry season. Langurs spent more feeding time on the ground in the dry than in the rainy season. Locomotor patterns in François' langurs are likely linked to morphological and anatomical characteristics, along with the limestone forest's structure. Our result completely supported prediction 1 but not prediction 2. This study suggests that seasonal variation in positional behaviour might result from the temporal difference in spatial distribution of foods and behavioural thermoregulation strategy. We found that François' langurs adjusted positional behaviour in response to seasonality, and this behavioural flexibility allows them to survive in a variety of habitats, including limestone forests.

Individuality in coo calls of adult male golden snub-nosed monkeys (Rhinopithecus roxellana) living in a multilevel society.

Vocal individuality is a prerequisite for individual recognition, especially when visual and chemical cues are not available or effective. Vocalizations encoding information of individual identity have been reported in many social animals and should be particularly adaptive for species living in large and complexly organized societies. Here, we examined the individuality in coo calls of adult male golden snub-nosed monkeys (Rhinopithecus roxellana) living in a large and multilevel society. Coo calls are one of the most frequently occurring call types in R. roxellana and likely serve as the signals for contact maintenance or advertisement in various contexts including group movement, foraging, and resting. From April to October 2016, April to July 2017, and September to October 2017, we recorded a total of 721 coo calls from six adult males in a provisioned, free-ranging group and one adult male in captivity in Shennongjia National Park, China. We selected 162 high-quality recordings to extract 14 acoustic parameters based on the source-filter theory. Results showed that each of all parameters significantly differed among individuals, while pairwise comparisons failed to detect any parameter that was different between all pairs. Furthermore, a discriminant function analysis indicated that the correct assignment rate was 80.2% (cross-validation: 67.3%), greater than expected by chance (14.3%). In conclusion, we found evidence that coo calls of adult male R. roxellana allowed the reliable accuracy of individual discrimination complementarily enhanced by multiple acoustic parameters. The results of our study point to the selective pressures acting on individual discrimination via vocal signals in a highly gregarious forest-living primate.

Lichen as a Biomonitor for Vehicular Emission of Metals: A Risk Assessment of Lichen Consumption by the Sichuan Snub-Nosed Monkey (Rhinopithecus roxellana).

Two lichen species, Usnea aciculifera and Usnea luridorufa, were used as biomonitors for the deposition of traffic-related metals in China's Shennongjia National Nature Reserve. The suitability of the two lichen species for use as biomonitors was compared. The health threat to the Sichuan snub-nosed (aka golden) monkey (Rhinopithecus roxellana) from consuming lichen with elevated metal concentrations due to vehicular traffic was then assessed. Lichens, with large surface areas and neither roots nor stomata, efficiently absorb both particulate and gaseous air pollutants. The resulting data was used to assess the effect of heavy metal accumulation on the lichens as well as the health risk imposed on the monkeys as lichen is a primary food source. Lichen samples were collected in the core area of the reserve at three locations of varying traffic intensity. A forth site in the reserve, with no proximate traffic, was used as the control. Results show: (1) lichen from high traffic sites has significantly higher concentrations of Fe, Cd, Pb Zn, and Cr than lichen collected from the control site; (2) vehicular traffic is the primary source of metals in lichen; (3) U. luridorufa collected at high traffic sites displayed decreased photosynthetic efficiency, an indication of stress; (4) intake of Cd and Pb from vehicle emissions in the Shennongjia National Nature Reserve could adversely affect snub-nosed monkey health. This research advances the science of biomonitoring, contributes to environmental protection efforts in China's nature reserves and helps improve food safety for Sichuan snub-nosed monkey, a national treasure of China.

A Short Cut to Mushrooms - Red Langur (Presbytis rubicunda) Consumption of Terrestrial Fungus.

We report the first in-depth evidence of targeted mushroom foraging in an Asian colobine. Using direct observations (2010-2018) and camera traps (2008-2018) in the Sebangau Forest, Central Kalimantan, Indonesian Borneo, we show how adult female red langurs (Presbytis rubicunda) are regularly descending to the ground to consume mushrooms. We recorded 82 counts (0.36% of all focal observations, n = 25,502) of the focal adult langur on the ground from direct observations of habituated groups, and 80 independent images/videos of red langurs on the ground were obtained from the camera traps representing 1.12% of total images (n = 7,145). Mushroom consumption took place in 4 families, representing 0.04% of total focal behaviour observations and 24.3% of total time feeding on the ground. From the camera trap photos, red langurs are spending 20% of time on the ground feeding. We speculate that mushrooms could be a supplementary food for adult female langurs as there is an increase in consumption in April and November.

A new fossil cercopithecid tibia from Laetoli and its implications for positional behavior and paleoecology.

Detailed analyses and comparisons of postcranial specimens of Plio-Pleistocene cercopithecids provide an opportunity to examine the recent evolutionary history and locomotor diversity in Old World monkeys. Studies examining the positional behavior and substrate preferences of fossil cercopithecids are also important for reconstructing the paleoenvironments of Plio-Pleistocene hominin sites. Here we describe a new fossil cercopithecid tibia (EP 1100/12) from the Australopithecus afarensis-bearing Upper Laetolil Beds (∼3.7 Ma) of Laetoli in northern Tanzania. The fossil tibia is attributed to cf. Rhinocolobus sp., which is the most common colobine at Laetoli. In addition to qualitative comparisons, the tibial shape of EP 1100/12 was compared to that of 190 extant cercopithecids using three-dimensional landmarks. Discriminant function analyses of the shape data were used to assess taxonomic affinity and shape variation relating to positional behavior. EP 1100/12 clustered with extant colobines, particularly the large-bodied genera Nasalis and Rhinopithecus. Comparisons reveal that EP 1100/12 belongs to a large-bodied monkey that engaged in arboreal pronograde quadrupedalism. These findings add further support to previous inferences that woodland and forest environments dominated the paleoenvironment of the Upper Laetolil Beds, which supported the diverse community of cercopithecids at Laetoli. The inferred paleoecology and the presence of large-bodied arboreally-adapted monkeys at Laetoli show that A. afarensis had access to a range of diverse habitats, including woodlands and forests. This supports the possibility that A. afarensis, with its potential range of positional capabilities, was able to utilize arboreal settings for food acquisition and refuge from predators.

Seasonal changes in social cohesion among males in a same-sex primate group.

Male-male interactions in mixed-sex groups of social mammals are typically characterized by a mix of hostility and affiliation, as a result of inherent conflicts over mating opportunities, and the costs and benefits of social alliances, co-operative behaviors, and coalitionary defense. In species of nonhuman primates that form all-male groups, it is still unclear how the tradeoffs between the benefits of forming an all-male group and the cost of male-male competition in seeking mating opportunities with females in bisexual groups influence social cohesion in different seasons. Here, we used social network analysis to quantify the impact of reproductive seasonality on social cohesion and clique size of bachelor males residing in an all-male unit (AMU) in wild black-and-white snub-nosed monkeys (Rhinopithecus bieti). These primates are strict seasonal breeders and live in a modular social system composed of a number of one-male units (OMUs) and an associated peripheral AMU. We found that the AMU social network had a significantly lower density, centralization, clustering coefficient, and smaller clique size during the mating season compared to the non-mating period. However, aggression among AMU males during both mating and non-mating periods was low. Our results suggest that network structure topology in male same-sex social units is modulated by seasonal changes. Bachelor males engage in two types of competition to gain reproductive success: first, which is analogous to contest competition, in which bachelor males act aggressively and challenge OMU leader males in an attempt to take over an OMU; and second, which is more analogous to scramble competition, in which bachelor males avoid aggressive interactions and instead engage in sneaky copulations with fertile females. Our work adds to an understanding of the maintenance of all-male groups in species that form a multilevel society.

Sleeping site and tree selection by Sichuan snub-nosed monkeys (Rhinopithecus roxellana) in Baihe Nature Reserve, Sichuan, China.

For non-human primates to optimize their survival chances, sleeping site selection is crucial as they spend much of their time sleeping. We studied sleeping site and tree selection by a group of wild Sichuan snub-nosed monkeys (Rhinopithecus roxellana) in the temperate forests of Baihe Nature Reserve, Sichuan, China, to assess if certain site and tree characteristics were selected. We identified a total of 39 sleeping sites and 111 sleeping trees over a period of 1 year. We compared nine sleeping site and six sleeping tree variables related to the environment, habitat, and vegetation structure. We found that certain characteristics of sleeping sites and sleeping trees predicted their selection by R. roxellana. On a larger spatial scale, sleeping sites were selected based mainly on four factors: canopy height, slope direction, slope gradient, and vegetation type. They also selected sites with trees that were taller and larger, had larger crown diameters and higher bole branches. On a smaller spatial scale, they selected larger trees with larger crown diameters. The selection of these characteristics could be explained in terms of predator avoidance and thermoregulation efficiency, although it was difficult to delineate which was more important. This could be due to some characteristics of R. roxellana, that is, large groups living in temperate climates, which required a balanced strategy for sleeping site and tree selection in order to optimize their survival chances.

Vocal repertoire of free-ranging adult golden snub-nosed monkeys (Rhinopithecus roxellana).

Vocal signaling represents a primary mode of communication for most nonhuman primates. A quantitative description of the vocal repertoire is a critical step in in-depth studies of the vocal communication of particular species, and provides the foundation for comparative studies to investigate the selective pressures in the evolution of vocal communication systems. The present study was the first attempt to establish the vocal repertoire of free-ranging adult golden snub-nosed monkeys (Rhinopithecus roxellana) based on quantitative methods. During 8 months in Shennongjia National Park, China, we digitally recorded the vocalizations of adult individuals from a provisioned, free-ranging group of R. roxellana across a variety of social-ecological contexts. We identified 18 call types, which were easily distinguishable by ear, visual inspection of spectrograms, and quantitative analysis of acoustic parameters measured from recording samples. We found a great sexual asymmetry in the vocal repertoire size (females produced many more call types than males), likely due to the sex differences in body size and social role. We found a variety of call types that occurred during various forms of agonistic and affiliative interactions at close range. We made inference about the functions of particular call types based on the contexts in which they were produced. Studies on the vocal communication in R. roxellana are particularly valuable since they provide a case about how nonhuman primates, inhabiting forest habitats and forming complex social systems, use their vocalizations to interact with their social and ecological environments.

Identifying potential refugia and corridors under climate change: A case study of endangered Sichuan golden monkey (Rhinopithecus roxellana) in Qinling Mountains, China.

Climate change threatens endangered species and challenges current conservation strategies. Effective conservation requires vulnerability assessments for species susceptible to climate change and adaptive strategies to mitigate threats associated with climate. In this paper, we used the Maxent to model the impacts of climate change on habitat suitability of Sichuan golden monkey Rhinopithecus roxellana. Our results showed that (i) suitable habitat for Sichuan golden monkey was predicted to decrease by 37% in 2050s under climate change; (ii) the mean elevations of suitable habitat in the 2050s was estimated to shift 160 m higher; (iii) nature reserves protect 62% of current suitable habitat and 56% of future suitable habitat; and (iv) 49% of current suitable habitat was predicted to be vulnerable to future climate change. Given these results, we proposed conservation implications to mitigate the impacts of climate change on Sichuan golden monkey, including adjusting range of national park, establishing habitat corridors, and conducting long-term monitoring.

Seasonal variation in diet and nutrition of the northern-most population of Rhinopithecus roxellana.

There is a great deal of spatial and temporal variation in the availability and nutritional quality of foods eaten by animals, particularly in temperate regions where winter brings lengthy periods of leaf and fruit scarcity. We analyzed the availability, dietary composition, and macronutrients of the foods eaten by the northern-most golden snub-nosed monkey (Rhinopithecus roxellana) population in the Qinling Mountains, China to understand food choice in a highly seasonal environment dominated by deciduous trees. During the warm months between April and November, leaves are consumed in proportion to their availability, while during the leaf-scarce months between December and March, bark and leaf/flower buds comprise most of their diet. When leaves dominated their diet, golden snub-nosed monkeys preferentially selected leaves with higher ratios of crude protein to acid detergent fiber. While when leaves were less available, bark and leaf/flower buds that were high in nonstructural carbohydrates and energy, and low in acid detergent fiber were selected. Southern populations of golden snub-nosed monkey can turn to eating lichen, however, the population studied here in this lichen-absent area have adapted to their cool deciduous habitat by instead consuming buds and bark. Carbohydrate and energy rich foods appear to be the critical resources required for the persistence of this species in temperate habitat. The dietary flexibility of these monkeys, both among seasons and populations, likely contributes to their wide distribution over a range of habitats and environments.

Faecal Particle Size in Free-Ranging Proboscis Monkeys, Nasalis larvatus: Variation between Seasons.

Reducing the size of food particles is crucial for herbivores. Seasonal dietary changes are known to influence animals' chewing efficiency. Proboscis monkeys (Nasalis larvatus) are foregut fermenters, with a high chewing efficiency allowing them to achieve very fine faecal particles. In this study, we investigated how proboscis monkeys' chewing efficiency varies between wet and dry seasons, hypothesising differences possibly related to diet change. Faecal particle size analysis is an established approach to estimate chewing efficiency in mammalian herbivores. We analysed 113 proboscis monkey faecal samples collected in the Lower Kinabatangan Wildlife Sanctuary, between 2015 and 2017. By following standard sieve analysis protocols, we measured a mean particle size MPS0.025-8 of 0.45 ± 0.14 mm, and confirmed a previous result that proboscis monkeys have a very low faecal MPS. This study highlights a seasonal influence on proboscis monkeys' chewing efficiency, with smaller MPS (better chewing efficiency) during the wet season. During that time of the year, individuals may potentially change their diet, as all faecal samples contained intact seeds. Whether the seasonal MPS difference in proboscis monkeys is smaller than in other colobines due to their "rumination" strategy remains to be investigated.

Male-Male Mounting Behaviour in Free-Ranging Golden Snub-Nosed Monkeys (Rhinopithecus roxellana).

An all-male band of golden snub-nosed monkeys (Rhinopithecus roxellana) was observed for 3 months in the Qinling Mountains of China, in order to collect data on the frequencies and contextual significance of male-male mounting behaviour. Mounts occurred in a variety of affiliative, dominance-related and sexual contexts, which differed depending upon the ages of the males involved. Mounting behaviour in this group was mainly initiated by adults. Juveniles mounted each other in affiliative contexts (during play and prior to grooming). Adult males mounted subadult and juvenile partners in a greater variety of sociosexual contexts (dominance/rank-related interactions; reconciliation following agonistic encounters, and sometimes as a prelude to receiving grooming). However, subadults and juveniles were never observed to mount adults. In one dyad, involving an adult male and a subadult partner, mounting was more frequent and prolonged, and included bouts of deep pelvic thrusting. Two mounts resulted in anal intromissions and, in 1 case, the subadult partner exhibited seminal emission. Given that the study took place during the annual mating peak period of R. roxellana, it is possible that this unusual male-male sexual activity was related to the absence of mating opportunities for those adults that were excluded from 1-male units.

Male cooperation for breeding opportunities contributes to the evolution of multilevel societies.

A small number of primate species including snub-nosed monkeys (colobines), geladas (papionins) and humans live in multilevel societies (MLSs), in which multiple one-male polygamous units (OMUs) coexist to form a band, and non-breeding males associate in bachelor groups. Phylogenetic reconstructions indicate that the papionin MLS appears to have evolved through internal fissioning of large mixed-sex groups, whereas the colobine MLS evolved through the aggregation of small, isolated OMUs. However, how agonistic males maintain tolerance under intensive competition over limited breeding opportunities remains unclear. Using a combination of behavioural analysis, satellite telemetry and genetic data, we quantified the social network of males in a bachelor group of golden snub-nosed monkeys. The results show a strong effect of kinship on social bonds among bachelors. Their interactions ranged from cooperation to agonism, and were regulated by access to mating partners. We suggest that an 'arms race' between breeding males' collective defence against usurpation attempts by bachelor males and bachelor males' aggregative offence to obtain reproductive opportunities has selected for larger group size on both sides. The results provide insight into the role that kin selection plays in shaping inter-male cohesion which facilities the evolution of multilevel societies. These findings have implications for understanding human social evolution, as male-male bonds are a hallmark of small- and large-scale human societies.