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Multilevel societies (MLS), which have two or more levels of organization, are among the most complex primate social systems. MLS has only been seen in some primates, including humans. The aim of this study was to identify MLS in proboscis monkeys in Sabah, Malaysia, through direct observation (>32 months) and genetic analysis, based on feces collected from ~200 individuals, to estimate the degree of kinship. The results revealed that the social system of the proboscis monkey exhibited a form of MLS, with several core reproductive units and the bachelor group woven together into a higher-level band. Genetic analysis revealed that the females migrated randomly over short and long distances; however, the males tended to migrate relatively shorter distances than females. Combined with the results of direct observations, we concluded that proboscis monkeys form an MLS with a double or triple structure and patrilineal basis. The results of this study may have implications for the evolution of human societies, as to why patrilineal MLS, an important step in the evolutionary path of human societies, emerged in phylogenetically distant primates, such as the proboscis monkey.

Decades of research have led to a solid understanding of the social systems of gregarious apes: chimpanzees, bonobos, gorillas, and gibbons. As field studies have increasingly collected data from multiple neighboring habituated groups, genetic and social interconnections have been revealed. These findings provide a more nuanced picture of intergroup relations in apes, and have led to claims in the literature that some ape taxa have multilevel societies. A multilevel society is defined as a nested collection of social entities comprising at least two discernible levels of social integration between the individual and the population. We argue that the evidence for multilevel sociality sensu stricto in apes is currently inconclusive and that it is premature to abandon the traditional classification of ape social systems. However, available findings appear to be consistent with the existence of some degree of higher social grouping patterns. We propose the term supra-group organization which may adequately capture ape social systems when viewed from a top-down perspective.

Objectives: Although fermented food use is ubiquitous in humans, the ecological and evolutionary factors contributing to its emergence are unclear. Here we investigated the ecological contexts surrounding the consumption of fruits in the late stages of fermentation by wild primates to provide insight into its adaptive function. We hypothesized that climate, socioecological traits, and habitat patch size would influence the occurrence of this behavior due to effects on the environmental prevalence of late-stage fermented foods, the ability of primates to detect them, and potential nutritional benefits. Materials and methods: We compiled data from field studies lasting at least 9 months to describe the contexts in which primates were observed consuming fruits in the late stages of fermentation. Using generalized linear mixed-effects models, we assessed the effects of 18 predictor variables on the occurrence of fermented food use in primates. Results: Late-stage fermented foods were consumed by a wide taxonomic breadth of primates. However, they generally made up 0.01%–3% of the annual diet and were limited to a subset of fruit species, many of which are reported to have mechanical and chemical defenses against herbivores when not fermented. Additionally, latestage fermented food consumption was best predicted by climate and habitat patch size. It was more likely to occur in larger habitat patches with lower annual mean rainfall and higher annual mean maximum temperatures. Discussion: We posit that primates capitalize on the natural fermentation of some fruits as part of a nutritional strategy to maximize periods of fruit exploitation and/or access a wider range of plant species. We speculate that these factors contributed to the evolutionary emergence of the human propensity for fermented foods. Canadian Institute for Advanced Research/[]/CIFAR/Canadá UCR::Vicerrectoría de Docencia::Ciencias Básicas::Facultad de Ciencias::Escuela de Biología

Traditionally, the genus Rhinopithecus (Milne-Edwards, 1872, Primates, Colobinae) included four allopatric species, restricted in their distributions to China and Vietnam. In 2010, a fifth species, the black snub-nosed monkey (Rhinopithecus strykeri) was discovered in the Gaoligong Mountains located on the border between China and Myanmar. Despite the remoteness, complex mountainous terrain, dense fog, and armed conflict that characterizes this region, over this past decade Chinese and Myanmar scientists have begun to collect quantitative data on the ecology, behavior and conservation requirements of R. strykeri. In this article, we review the existing data and present new information on the life history, ecology, and population size of R. strykeri. We discuss these data in the context of past and current conservation challenges faced by R. strykeri, and propose a series of both short-term and long-term management actions to ensure the survival of this Critically Endangered primate species. Specifically, we recommend that the governments and stakeholders in China and Myanmar formulate a transboundary conservation agreement that includes a consensus on bilateral exchange mechanisms, scientific research and monitoring goals, local community development, cooperation to prevent the hunting of endangered species and cross-border forest fires. These actions will contribute to the long-term conservation and survival of this Critically Endangered species.

Competition for food is often a cost associated with living in a group, and can occur in an indirect (scramble) or direct (contest) form. We investigated feeding competition in a supergroup of Rwenzori black-and-white colobus monkeys (Colobus angolensis ruwenzorii) in Rwanda, with the aim of establishing whether freedom from scramble competition allows these monkeys to form supergroups. We used the patch depletion method, measuring intake rate coupled with movement rate, to assess if food patches become depleted over the occupancy period. Resource depletion was evident when the colobus fed on young leaves, but not when feeding on mature leaves. Scramble competition was inferred from a negative correlation between group size and change in intake rate over patch occupancy. Between-group contest competition was inferred from displacement from patches. Although feeding competition exists for select resources, limited competition for mature leaves may enable Rwenzori colobus to live in a supergroup of hundreds of individuals in this montane forest.

Multilevel societies (MLSs), stable nuclear social units within a larger collective encompassing multiple nested social levels, occur in several mammalian lineages. Their architectural complexity and size impose specific demands on their members requiring adaptive solutions in multiple domains. The functional significance of MLSs lies in their members being equipped to reap the benefits of multiple group sizes. Here, we propose a unifying terminology and operational definition of MLS. To identify new avenues for integrative research, we synthesise current literature on the selective pressures underlying the evolution of MLSs and their implications for cognition, intersexual conflict, and sexual selection. Mapping the drivers and consequences of MLS provides a reference point for the social evolution of many taxa, including our own species.

When preferred foods are scarce, one strategy employed by primates is to switch to an alternative food item of lower quality or preferability, i.e., a fallback food. In the montane rainforest of Nyungwe National Park in southwestern Rwanda, Rwenzori black-and-white colobus (Colobus angolensis ruwenzorii) (hereafter Rwenzori colobus) form a supergroup comprising hundreds of individuals. Over 13 mo we investigated how this supergroup uses resources in periods of resource abundance vs. periods of resource scarcity. Based on 5603 feeding records we first identified preferred foods and then identified fallback foods as those food categories or species whose consumption increases when preferred foods, or preferred food categories, are less available. When the availability of 19 preferred food items was low, fruticose lichens (Usnea sp.) contributed >50% of the monthly diet for the Rwenzori colobus. Moreover, consumption of lichens was significantly negatively related to the availability of preferred foods. Fruticose lichens can therefore be considered a fallback food for Rwenzori colobus that sustains the supergroup during periods of reduced food availability. This result, in combination with previous findings that mature foliage in Nyungwe is of high quality and does not elicit feeding competition, points to the importance of resources in facilitating supergroup formation. However, several other montane forests in Eastern Africa also harbor fruticose lichens and yet support only small groups of Angolan colobus, suggesting that additional factors such as sufficient forest size and limited fragmentation and hunting pressure by humans are required for supergroups to form.

In mammals characterized by a mating system in which a single male monopolizes reproduction, infanticide is reported to occur following a male take-over, often resulting in females returning to oestrus more rapidly than if their infant has survived. However, over the course of a 17-year study of golden snub-nosed monkeys, Rhinopithecus roxellana, a polygynous colobine species, we found no behavioural or suspected evidence of infanticide. Based on social network and genetic analyses, we found that more than half of the infants were sired as a result of extrapair matings (EPMs). Female golden snub-nosed monkeys initiated EPMs with bachelor or neighbouring resident males. These were considered ‘sneaky’ copulations because they did not occur in the presence of the leader male of the female's one-male unit (OMU). We suggest that through a process of paternity confusion, females decreased the likelihood of infanticide after a male take-over. In contrast, in a group of the same species in which the number of bachelor males was artificially reduced, EPMs were infrequent and four cases of infanticide were observed, three of them following a male take-over. Golden snub-nosed monkeys reside in a large multilevel society that develops by the fusion of several independent OMUs to form a breeding band, which is characterized by increased social cohesion and intrasexual tolerance among leader males. Females play an active role in mate choice, and engage in a diverse set of mating tactics, including counterstrategies such as EPMs to avoid infanticide. These results offer new insights into the diversity of behavioural strategies facilitated by complex social structures in nonhuman primates, as well as other social mammals.

Group-living animals face a number of threats from extragroup conspecifics: from individuals seeking mating opportunities to rival groups attempting to access limited resources. The consequences of intergroup interactions can therefore include loss of mates, increased energy expenditure, and injury or death. There is increasing evidence that aggressive intergroup interactions can affect subsequent intragroup behavior, and that such post-conflict behavior may be directly related to the threats posed by different opponent groups (e.g., familiar vs. unfamiliar). However, empirical evidence for changes in intragroup behavior following intergroup conflict in social animals is limited. We compared the proximity and behavior of group members before and after 84 aggressive intergroup interactions involving 14 study groups of wild, habituated mountain gorillas (Gorilla beringei beringei) in Volcanoes National Park, Rwanda. Intergroup conflict affected subsequent intragroup behavior in several ways. We found that immediate movement away from the opponent following intergroup conflict was less likely between familiar groups compared to unfamiliar groups. However, both winning and losing groups spent twice as much time moving post-conflict, and losing groups spent less time resting. There was an increase in female intragroup affiliative interactions and a decrease in male intragroup agonistic interactions post-conflict. These results demonstrate that such intergroup contests can influence intragroup dynamics beyond the immediate period of interaction.

Investment in social interaction and affiliative behaviour is often related to variation in sex-specific dispersal patterns among species but can also vary within species in response to local environmental conditions and feeding competition. Chimpanzee (Pan troglodytes) association patterns have been studied primarily in low and mid-elevation tropical forests. This study adds to our understanding of association patterns in chimpanzees by presenting the first data on association patterns in a high-elevation population at Nyungwe in equatorial Rwanda. Using data on co-occurrence in social parties (party association) and calculating gregariousness indices, we quantified levels of associations among age-sex classes relative to food (fruit) distribution and the presence of oestrous females. Overall, adult females in Nyungwe are less gregarious than males. Female but not male gregariousness increased when feeding on clumped foods, indicating that resource characteristics impact female association tendencies more intensely than males. Despite high elevation forests representing a habitat with relatively fewer fruit resources for chimpanzees, at an edge of range location, we found that gregariousness patterns were driven by similar factors as those in mid and low elevation chimpanzee populations. – We quantified association patterns in a high elevation population of chimpanzees at Nyungwe (Rwanda)– Adult females were less gregarious than males– Female but not male gregariousness increased when feeding on clumped foods, indicating that resource characteristics impact female association tendencies more intensely than males – We quantified association patterns in a high elevation population of chimpanzees at Nyungwe (Rwanda) – Adult females were less gregarious than males – Female but not male gregariousness increased when feeding on clumped foods, indicating that resource characteristics impact female association tendencies more intensely than males

Both observational and indirect evidence are widely used to determine the diets of wild animals. Direct observations are often assumed to provide the most comprehensive reflection of diet, but many wild animals are logistically challenging to observe. Despite the regular use of observational and indirect methods for inferring diet in wild animals, they have rarely been compared in detail for the same study population. Over 12 months this study assessed the congruence of methods in estimating the diet of a montane community of eastern chimpanzees Pan troglodytes schweinfurthii in Nyungwe National Park, Rwanda using observational scan samples and macroscopic fecal inspection. The assessment of the number of food species consumed each month was comparable between methods, but the estimation of the composition of items in the diet differed significantly. Most notably, the fecal samples significantly underestimated the consumption of flowers, and certain fruit species, which based on direct behavioral observations were seasonally consumed at very high rates. Conversely, direct observations underestimated the consumption of leaves and pith in comparison to results present in the fecal samples. These results suggest that combining methods where possible is most useful for accurate monitoring of dietary trends, particularly for species that experience significant seasonal shifts in their diet.

Simple Summary The size of primate ‘harems’ varies considerably, both inter- and intra-specifically. Previous studies have shown that females prefer high-quality males and that high-quality males are superior in inter-male competition, leading to them having a larger harem size. Based on eleven years of observations of Yunnan snub-nosed monkeys (Rhinopithecus bieti), we documented longitudinal stability in the distribution of harem sizes between 2010 and 2020. These demographic properties are the outcome of male and female social investment decisions that affect their reproductive performance and success. Male reproductive success was positively related to harem size, while constraints on individual social benefits and social investments limited harem size. Our findings advance our understanding of the socioecological determinants of harem size variation in polygynous primates. Abstract We used long-term data on the variation in harem size in Yunnan snub-nosed monkeys to research the effects of harem size on reproductive success and the ratio of grooming received to given (RGRG). The results suggest that harem holders derive reproductive benefits commensurate with harem size, whereas the females’ reproductive success is unaffected by harem size. Males of larger harems groomed less and had higher RGRG than males of smaller harems. In the case of females, grooming given increased, and RGRG decreased with an increase in harem size. The males’ reproductive success seems to be a driver of harem size maximization. From the females’ perspective, dwindling social benefits appear to set the upper limit for harem enlargement. We also showed that males of monogamous units (‘single-female harems’) invested more into grooming their female, presumably to prevent unit disintegration and loss of mating privileges.

Animal communication has long been thought to be subject to pressures and constraints associated with social relationships. However, our understanding of how the nature and quality of social relationships relates to the use and evolution of communication is limited by a lack of directly comparable methods across multiple levels of analysis. Here, we analysed observational data from 111 wild groups belonging to 26 non-human primate species, to test how vocal communication relates to dominance style (the strictness with which a dominance hierarchy is enforced, ranging from 'despotic' to 'tolerant'). At the individual-level, we found that dominant individuals who were more tolerant vocalized at a higher rate than their despotic counterparts. This indicates that tolerance within a relationship may place pressure on the dominant partner to communicate more during social interactions. At the species-level, however, despotic species exhibited a larger repertoire of hierarchy-related vocalizations than their tolerant counterparts. Findings suggest primate signals are used and evolve in tandem with the nature of interactions that characterize individuals' social relationships. Publisher PDF

The conservation of endangered species can benefit from a clear understanding of the quantity and distribution of their main foods. The population of mountain gorillas (Gorilla beringei beringei) living in the Virunga Massif of Rwanda, Uganda, and Democratic Republic of Congo has doubled in size since the 1980s, due to success in conservation efforts in and around their habitat. However, this increase in population size along with pressures on gorilla habitat raises concerns about spatial-temporal changes in the gorillas’ food plants. This study modelled the abundance and distribution of gorilla food species in the Virunga Massif. A total of 1050 vegetation recordings were collected on five plant species that are known to be frequently consumed by gorillas in one region of the Virungas, the Karisoke area. Two types of datasets collected along vegetation zones were combined: one with plant abundance expressed with Braun-Blanquet scores; and the other with abundance expressed as biomass. Moreover, ecological characteristics of locations where these species occur were extracted from satellite imagery. Analysis of variance and linear regression models were used to examine relationships between food species abundances and predictor variables. Subsequently, maps for the food species were created using boosted regression trees (BRTs). The abundance of species differed across vegetation zones, and the differences were statistically significant among vegetation zones with enough species observations. The accuracy of the BRTs indicated greater than random predictions (AUC > 0.65). This study shows the suitable areas for these gorilla food species and relevant ecological variables determining their distribution. The results provide insights into habitat occupancy by mountain gorillas, and help to design a baseline for monitoring changes in the abundance of gorilla food species under changing climate and anthropogenic pressure.

Urban living is often thought to promote incivility, but the existing sociological evidence paints a mixed picture. We aimed to examine the urban incivility phenomenon from an evolutionist’s perspective. Small communities are expected to show a higher incidence of helping because the applicability of theories such as kin selection, direct reciprocity and indirect reciprocity to acts of cooperative behaviour is augmented in small-scale demographic settings. Smaller communities have a reduced total pool of individuals to interact with, increasing the likelihood of encountering any given individual multiple times. This makes it easier for individuals to form cooperative relationships with one another, which may facilitate prosociality within smaller communities. Using the lost letter technique, our results show that city dwelling, compared with rural residence, per se does not negatively influence prosociality. This contradicts the expected erosion of cooperative behaviour in anonymous cities and adds to our understanding of the interplay between human macroecology and individual behavioural tendencies.

Correlational models are widely used to predict changes in species' distribution, but generally have failed to address the comprehensive effects of anthropogenic activities, climate change, habitat connectivity and gene flow on wildlife sustainability. Here, we used integrated approaches (MAXENT model, circuit model and genetic analysis) to assess and predict the effects of climate change and anthropogenic activities on the distribution, habitat connectivity, and genetic diversity of an endangered primate, Rhinopithecus bieti, from 2000 to 2050. We created six scenarios: climatic factors only (scenario-a), anthropogenic activities only (scenario-b), climatic factors and anthropogenic activities (scenario-c), plus three additional scenarios that included climatic factors and anthropogenic activities but controlled for individual anthropogenic activities (scenario-d: grazing, scenario-e: collecting, and scenario-f: grazing and collecting). The results indicate that areas of suitable habitat for R. bieti are expected to decline by 8.0%–22.4% from 2000 to 2050, with the collection of local forest products and the grazing of domesticated cattle as the primary drivers of landscape fragmentation and range contraction. If these anthropogenic activities are strictly controlled, however, the area of suitable habitat is predicted to increase by10.4%–14.3%. We also found that habitats vulnerable to human disturbance were principally located in areas of low habitat connectivity resulting in limited migration opportunities and increased loss of genetic diversity among R. bieti living in these isolated subpopulations. Thus, we suggest that effective management policies to protect this species include prohibiting both livestock grazing and the collecting of forest products. Although our study focuses on a single primate species, the conservation modeling approaches we presented have wide applicability to a broad range of threatened mammalian and avian taxa that currently inhabit a limited geographic range and are affected by anthropogenic activities (e.g. collecting, grazing, hunting), loss of habitat connectivity, reduced genetic diversity, and the effects of climate change.

In species with flexible grouping dynamics (i.e., fission-fusion), party (or subgroup) size is often shaped by available resources. Food resources are thought to limit party size in a range of mammalian species, reflecting a strategy of reducing feeding competition. In montane habitats, where food is highly seasonal, we may expect to see strong effects of ecological constraints on party size. In the montane forest of Nyungwe National Park, Rwanda, we quantified changes in chimpanzee (Pan troglodytes schweinfurthii) party size. We used path analysis to analyze the direct and indirect effects of (i) ecological variables and (ii) a reproductive variable (estrous females) on party size. Our path analysis adds precision and directionality to the hypothesis that food availability and estrous females influence party size. We found that the presence of estrous females had the strongest effect on party size. Interestingly, the availability and distribution of important and preferred fruits did not directly influence party size but did influence the presence of estrous females. These findings indicate that fruit distribution may be the ecological precondition that attracts estrous females, which is the main driver of larger parties. Party size was, however, positively correlated with patch size, indicating that specific fruit species may be of particular importance to the ecology of these chimpanzees. In montane habitats with lower fruit tree density and diversity than lowland sites, it may be expected that ecological factors play a more important role in fission-fusion grouping patterns than reproductive factors. This is because the cost of competition for food among group members may be higher in a more resource-limited (montane) environment. We investigated the ecological and reproductive factors that influence grouping patterns of chimpanzees in a montane forest that encompasses the upper limit of this species’ altitudinal range. Our results showed that the presence of estrous females had the strongest influence on party size. These results have implications for evaluating ecological and reproductive drivers of chimpanzee party size across differing habitats.

Additional co-authors: Fabiano R de MELO, P Fan, Cyril C Grueter, Diana C Guzman-Caro, Eckhard W Heymann, Ilka Herbinger, Minh D Hoang, Robert H Horwich, Tatyana Humle, Rachel A Ikemeh, Inaoyom S Imong, Leandro Jerusalinsky, Steig E Johnson, Peter M Kappeler, Maria Cecilia M Kierulff, Inza Kone, Rebecca Kormos, Khac Q LE, Baoguo Li, Andrew J Marshall, Erik Meijaard, Russel A Mittermeier, Yasuyuki Muroyama, Eleonora Neugebauer, Lisa Orth, Erwin Palacios, Sarah K Papworth, Andrew J Plumptre, Ben M Rawson, Johannes Refisch, Jonah Ratsimbazafy, Christian Roos, Joanna M Setchell, Rebecca K Smith, Tene Sop, Christoph Schwitzer, Kathy Slater, Shirley C Strum, William J Sutherland, Mauricio Talebi, Janette Wallis, Serge Wich, Roman M Wittig, Hjalmar S Kuhl

Abstract Two mechanisms have been proposed to explain why scramble competition can increase the travel requirements of individuals within larger groups. Firstly, individuals in larger groups may be more likely to encounter food sites where other group members have already eaten, leading to greater asynchronous “individual” travel to find fresh sites. Secondly, when food sites are aggregated into patches, larger groups may need to visit more patches to obtain the same amount of food per capita, leading to greater synchronous “group” travel between patches. If the first mechanism can be mitigated by increasing group spread, then we expect the second mechanism to be more sensitive to group size. Here, we examine the individual travel and group travel of the Virunga mountain gorillas, along with potential implications for the two mechanisms of scramble competition. Asynchronous individual travel accounted for 67% of the total travel time, and the remainder arose from group travel. Group spread increased significantly for larger groups, but not enough to prevent an increase in individual travel. Contrary to expectations, group travel decreased with size among most groups, and we found only limited evidence of patch depletion that would cause the second mechanism of scramble competition. Collectively, our results illustrate how the influence of group size can differ for individual travel versus group travel, just as it differs among species for overall travel. Studies that distinguish between the two mechanisms of scramble competition may enhance our understanding of ecological constraints upon group size, including potential differences between frugivores and folivores. Significance statement Feeding competition provides insight into how group size can influence the foraging patterns of social animals, but two key mechanisms are not typically compared. Firstly, larger groups may visit more patches to access the same amount of food per capita (group travel). Secondly, their individuals may also need to move past more spots where another member has already eaten (individual travel). Contrary to expectations, we found that group travel decreased with size for most groups of mountain gorillas, which may reflect extra travel by smaller groups to avoid larger groups. Individual travel increased with size in most groups, even though gorillas in larger groups compensated by spreading out over a broader area. The two mechanisms revealed patterns that were not apparent in our previous study of overall travel. Our approach may help to explain potential differences between folivores and frugivores.

Threats to biodiversity are well documented. However, to effectively conserve species and their habitats, we need to know which conservation interventions do (or do not) work. Evidence-based conservation evaluates interventions within a scientific framework. The Conservation Evidence project has summarized thousands of studies testing conservation interventions and compiled these as synopses for various habitats and taxa. In the present article, we analyzed the interventions assessed in the primate synopsis and compared these with other taxa. We found that despite intensive efforts to study primates and the extensive threats they face, less than 1% of primate studies evaluated conservation effectiveness. The studies often lacked quantitative data, failed to undertake postimplementation monitoring of populations or individuals, or implemented several interventions at once. Furthermore, the studies were biased toward specific taxa, geographic regions, and interventions. We describe barriers for testing primate conservation interventions and propose actions to improve the conservation evidence base to protect this endangered and globally important taxon.

Primates display broad diversity in their social organization. The social groups of a few primate species are organized in a multilevel fashion, with large groups composed of multiple, core one-male units (OMUs). A characteristic of multilevel societies is that the higher levels can include hundreds of individuals. The Rwenzori black-and-white colobus (Colobus angolensis ruwenzorii) in the montane forests of Rwanda form supergroups and have been suspected to exhibit multilevel social organization. Here we present the first data on the "anatomy" of a supergroup numbering 500+ individuals. We identified subgroups within the supergroup based on progression data, extracting the social network structure from the time-stamped spatiotemporal distribution of passing individuals identified to age-sex class, and selecting an optimal time window for each network using the two-step approach developed by Uddin, Choudhury, Farhad, and Rahman (2017). We detail the existence of core units-multi-male units (MMUs) with a mean of 1.7 adult males and 3.1 adult females, as well as OMUs, all-female units and bachelor units composed of adult and sub-adult males. More than two-thirds of units are MMUs. These grouping patterns conform to a multilevel society with predominantly multi-male core units, a social system that has recently also been described for a population of the same taxon in Uganda. Individual identification will be required to corroborate these interpretations.

Anthropogenic changes and fragmentation of natural habitats often exert a negative effect on resource availability and distribution, and the nutritional ecology and feeding behavior of nonhuman primates. The goals of this study are to examine food choice and to identify the nutritional profile of foods consumed by the Critically Endangered black snub-nosed monkey (Rhinopithecus strykeri). To accomplish our study goals, we presented cafeteria-style feeding trials of fresh food items collected in the home range of wild black snub-nosed monkeys to the only two captive R. strykeri, and compared the nutritional profiles of the leafy foods (buds, young, and mature leaves, 100 i23tems from 70 plant species) selected with those avoided (54 items from 48 plant species). Overall, the results indicate that captive R. strykeri selected foods that were higher in moisture (Mo; 77.7%), crude protein (CP; 21.2%), total nonstructural carbohydrates (TNC; 34.9%), and phosphorus (P; 0.37%) while tending to avoid foods with a neutral detergent fiber (NDF) content of greater than 46.8%. Leaves collected in autumn and selected by the monkeys were characterized by a slightly higher amount of metabolizable energy (ME) than those rejected (1,350 kJ/100 g vs. 1,268 kJ/100 g). In contrast, the protein content of foods collected and consumed during the spring was greater (22.9%) than in autumn (16.4%). Random Forests modeling, an ensemble learning method, indicated that the proportion of Mo, NDF, ME, CP, P, and TNC were among the most important factors in predicting which items were consumed by the captive R. strykeri during spring and autumn. On the basis of the nutritional profile of foods consumed across the two seasons, we identified 18 nutrient-rich native plant species that we recommend for use in ex- and in-situ conservation management and reforestation programs to provide long-term access to a nutritionally adequate diet.

In group-living species, encounters with extragroup rivals can be one of the riskiest actions in which individuals participate. Different group members often have different incentives to participate during intergroup interactions, and individuals with fewer payoffs of competition, including those of the smaller sex and/or lower rank, may ‘free-ride’ to avoid the costs of conflict. However, there is little evidence for how different types of intergroup interactions (e.g. interactions that do not involve conflict) can influence the participation of individuals. We examined the ecological, demographic and social predictors of individual participation in interactions between 14 fully habituated mountain gorilla groups in Volcanoes National Park, Rwanda from 2003 to 2015. The probability of an individual participating decreased with group size but remained relatively high in aggressive interactions and in multimale groups, illustrating the potential for ‘load lightening’ among group members. Males with fewer mating opportunities participated less often than males with more mating opportunities; however, male participation was significantly higher than female participation across all types of intergroup interactions. Females were more likely to be involved in aggressive interactions with solitary males, possibly to avoid the potential cost of infanticide if a resident male is killed or injured. Both sexes demonstrated more affiliative behaviours towards familiar groups, indicating a benefit of maintaining social relationships with familiar groups. Individuals show considerable variation in behaviour during intergroup interactions, and our results suggest that this variation is primarily driven by intergroup familiarity and individual reproductive benefits, both of which may have long-term consequences for individual fitness.

China is facing an unprecedented set of challenges in balancing the effects of economic development and global climate change with environmental protection and maintaining biodiversity. Although positive steps have been undertaken to remedy this situation, currently 80% of China’s 25 extant primate species are threatened, 15–18 species have population sizes of less than 3000 individuals, and two species of gibbons and one species of langur have been extirpated over the past few decades. Today, virtually all species of primates in China inhabit fragmented landscapes and are distributed in small isolated subpopulations with limited opportunities to exchange individuals or genetic information. Here we present a historical framework examining how human-induced environmental changes, particularly since the second half of the 20th century, accelerated primate population decline in China. In addition, we modeled the expected spatial conflict between agricultural expansion and primate distributions over the next 25–75 years and assessed the current overlap between protected areas and primate distributions. Depending on the assumptions of the spatial conflict model, primate distributions are expected to decline by an additional 51–87% by the year 2100. Thus, unless large-scale conservation policies are implemented immediately the current trend of primate population decline, local extirpation, and species extinctions will accelerate. To mitigate against such extinction scenarios, we advocate the creation of a Chinese national agency and repository of environmental information focused on public awareness and education, the implementation of targeted programs of habitat restoration designed to return impacted forests to a more natural state especially within and at the boundaries of nature reserves, the establishment of additional protect areas, and the construction of a latticework of corridors connecting isolated primate subpopulations. This comprehensive approach offers the most effective way to protect China’s animal and plant biodiversity, including its endangered primate populations.