A global risk assessment of primates under climate and land use/cover scenarios

Primates are facing an impending extinction crisis, driven by extensive habitat loss, land use change and hunting. Climate change is an additional threat, which alone or in combination with other drivers, may severely impact those taxa unable to track suitable environmental conditions. Here, we investigate the extent of climate and land use/cover (LUC) change‐related risks for primates. We employed an analytical approach to objectively select a subset of climate scenarios, for which we then calculated changes in climatic and LUC conditions for 2050 across primate ranges (N = 426 species) under a best‐case scenario and a worst‐case scenario. Generalized linear models were used to examine whether these changes varied according to region, conservation status, range extent and dominant habitat. Finally, we reclassified primate ranges based on different magnitudes of maximum temperature change, and quantified the proportion of ranges overall and of primate hotspots in particular that are likely to be exposed to extreme temperature increases. We found that, under the worst‐case scenario, 74% of Neotropical forest‐dwelling primates are likely to be exposed to maximum temperature increases up to 7°C. In contrast, 38% of Malagasy savanna primates will experience less pronounced warming of up to 3.5°C. About one quarter of Asian and African primates will face up to 50% crop expansion within their range. Primary land (undisturbed habitat) is expected to disappear across species' ranges, whereas secondary land (disturbed habitat) will increase by up to 98%. With 86% of primate ranges likely to be exposed to maximum temperature increases >3°C, primate hotspots in the Neotropics are expected to be particularly vulnerable. Our study highlights the fundamental exposure risk of a large percentage of primate ranges to predicted climate and LUC changes. Importantly, our findings underscore the urgency with which climate change mitigation measures need to be implemented to avert primate extinctions on an unprecedented scale.     

Distinguished primatologist address—moving from advocacy to activism: Changing views of primate field research and conservation over the past 40 years

Field studies of wild nonhuman primates have grown exponentially over the past 40 years and our knowledge of primate behavior, ecology, and social, and mating systems has expanded greatly. However, we are facing a major extinction crisis with some 60% of all primate species listed as threatened and more than 75% of species with declining populations. The primary factor driving primate population decline is human population increase, which over the past 50 years has resulted in the unsustainable conversion and degradation of natural landscapes for industrial agriculture, the production of nonagricultural commodities for international trade, pastureland for cattle, dam construction, fossil fuel exploration, mining, and the construction of road networks and infrastructure to support large urban centers. Recent ecological modeling predicts that by the end of the century, the four primate‐richest countries in the world will lose 32–78% of their existing primate habitat to agricultural expansion, and nine of the top 15 primate‐richest countries are expected to have 80–100% of their primate species extinct or threatened with extinction. If we are going to save the world's primates, the time to act is now! Not only should all primate field research include a strong conservation component, but in addition we must actively join with our professional societies, zoos and research facilities, universities, conservation organizations, concerned business leaders, global citizens, like‐minded political leaders, and grassroots organizations to inform, demand and direct governments, multinational corporations, and international organizations to engage in transformational change to protect biodiversity and seek environmental justice against those entities that actively destroy our planet. As the chief academic discipline dedicated to the study of primates, we must organize and collectively move from being advocates for primate conservation to becoming activists for primate conservation. This is a call to action.     

Human-modified landscapes driving the global primate extinction crisis

The world's primates have been severely impacted in diverse and profound ways by anthropogenic pressures. Here, we evaluate the impact of various infrastructures and human-modified landscapes on spatial patterns of primate species richness, at both global and regional scales. We overlaid the International Union for the Conservation of Nature (IUCN) range maps of 520 primate species and applied a global 100 km² grid. We used structural equation modeling and simultaneous autoregressive models to evaluate direct and indirect effects of six human-altered landscapes variables (i.e., human footprint [HFP], croplands [CROP], road density [ROAD], pasture lands [PAST], protected areas [PAs], and Indigenous Peoples' lands [IPLs]) on global primate species richness, threatened and non-threatened species, as well as on species with decreasing and non-decreasing populations. Two-thirds of all primate species are classified as threatened (i.e., Critically Endangered, Endangered, and Vulnerable), with ~86% experiencing population declines, and ~84% impacted by domestic or international trade. We found that the expansion of PAST, HFP, CROP, and road infrastructure had the most direct negative effects on primate richness. In contrast, forested habitat within IPLs and PAs was positively associated in safeguarding primate species diversity globally, with an even stronger effect at the regional level. Our results show that IPLs and PAs play a critical role in primate species conservation, helping to prevent their extinction; in contrast, HFP growth and expansion has a dramatically negative effect on primate species worldwide. Our findings support predictions that the continued negative impact of anthropogenic pressures on natural habitats may lead to a significant decline in global primate species richness, and likely, species extirpations. We advocate for stronger national and international policy frameworks promoting alternative/sustainable livelihoods and reducing persistent anthropogenic pressures to help mitigate the extinction risk of the world's primate species.     

Distribution and Population Densities of Diurnal Primates in the Karst Forests of Phong Nha – Ke Bang National Park,Quang Binh Province,Central Vietnam

We studied the status and distribution of the diurnal primates in the Phong Nha – Ke Bang National Park (PNKB NP) from April to August 2007. In the past, researchers reported 9 primate species and subspecies for the karst forests of PNKB NP, which constitutes the most important protected area for the endangered Hatinh langur (Trachypithecus hatinhensis) in Vietnam. All 9 primate taxa are threatened due to hunting pressure and habitat loss. We applied line transect sampling in 4 areas. During a total of 117 transect inspections along 12 different transect routes, we confirmed 5 primate taxa and the analysis of population densities resulted in 2143 (±467) Hatinh langurs, 1316 (±871) red-shanked douc langurs (Pygathrix nemaeus), 930 (±489) stump-tailed macaques (Macaca arctoides), 986 (±883) eastern Assamese macaques (M. a. assamensis), and 18 (±18) southern white-cheeked crested gibbons (Nomascus siki) in the whole PNKB NP, which covers an area of ca. 85,000 ha. We could not detect the 2 nocturnal lorises, Bengal slow lorises (Nycticebus bengalensis) and pygmy slow lorises (N. pygmaeus), as well as rhesus macaques (Macaca mulatta) and northern pig-tail macaques (M. leonina). The distribution of the primates predominantly depended on human impact. We could not recognize a correlation between habitat constitution and abundance of primates. The population density estimates showed a much higher density of the Hatinh langur than previously assumed. Thus the importance of the PNKB NP for the conservation of this endangered langur increased significantly.     

Putting the Spotlight on Internally Displaced Animals (IDAs): A Survey of Primate Sanctuaries in Africa,Asia, and the Americas

As anthropogenic activity makes deeper incursions into forests, fragmenting habitat, wildlife is forced into closer proximity to humans leading to increased incidences of human–wildlife conflict and wildlife displacement. These same incursions facilitate poaching for the commercial trade in dead and live animals. As a direct result, the number of sanctuaries and internally displaced animals (IDAs) in need of sanctuary placement and rehabilitation are increasing. We focus on internally displaced primates given the prevalence of primate‐focused facilities and anthropomorphic considerations surrounding this taxonomic group. Surveys were distributed globally to map the extent and range of native primate sanctuaries and species. Over 70 facilities care for more than 6,000 native primates comprising 64 species, with almost half listed as endangered or critically endangered. As not all sanctuaries were identified at the time of the survey distribution, we estimate that the actual number of facilities is closer to double this number with a captive population in excess of 10,000 individual primates. Native primate sanctuaries hold significant numbers of primates in long‐term captive care, with less than half (37%) identified as candidates for release. The surveyed sanctuary population accounts for 35% of the world's captive primates, as compared to ISIS‐registered (where ISIS is International Species Information System) zoological facilities, although we estimate that the actual population is closer to 58%. For some species, the sanctuary population represents the only population in captivity. We discuss the prevalence of range‐state sanctuaries and their primate populations, and issues surrounding their future development and management. Am. J. Primatol. 75:116‐134, 2013. © 2012 Wiley Periodicals, Inc.     

Noninvasive Saliva Collection for DNA Analyses From Free‐Ranging Tibetan Macaques (Macaca thibetana)

Cryptic and endangered fauna, including many primate taxa, pose challenges for noninvasive collection of biomaterials. As a result, application of noninvasive genotyping to primates has been limited to the use of samples such as feces and hair for the extraction of PCR‐amplifiable DNA. We present a method for noninvasive collection of saliva from habituated, free‐ranging monkeys. The method utilizes a low‐cost apparatus that controls for contamination and is usable with individual, free‐ranging primates. Saliva samples were collected from 18 individuals in a population of Tibetan macaques (Macaca thibetana) in the Valley of Wild Monkeys in Huangshan, People's Republic of China. DNA was extracted from these samples and PCR‐amplified for both mitochondrial and nuclear genes, Cytochrome B and MHC‐DR Beta 1, respectively. These results indicate this is an effective technique for the noninvasive collection of saliva across age and sex class, and dominance rank in a free‐ranging, terrestrial primate species. This device could have wide application for obtaining high‐quality saliva samples from free‐ranging primate populations for use in epidemiological studies, hormonal analyses of HPA axis function, pathogen screening, noninvasive genotyping, and behavioral genetics. Am. J. Primatol. 74:1064‐1070, 2012. © 2012 Wiley Periodicals, Inc.     

Diet of the Critically Endangered Brown Spider Monkey (Ateles hybridus) in an Inter‐Andean Lowland Rainforest in Colombia

Brown spider monkeys (Ateles hybridus) are one of the least known and more threatened primates in the Neotropics. Recognized as a species about a decade ago, field studies on these endangered primates have mainly focused on estimating local population densities. Since 2006, we habituated a group of wild brown spider monkeys at Serranía de Las Quinchas, Colombia, and studied their feeding ecology during 2.5 years using focal “subgroup” sampling, and conducted phenological surveys in order to estimate habitat‐wide fruit availability. Based on 847 hr of behavioral follows, brown spider monkeys spent approximately 25% of their time in feeding activities, and fed from fruits and leaves on at least 123 plant species. Ripe fruits were the most important item in the diet of A. hybridus at Las Quinchas comprising 92% of their feeding time. Probably due to the minor variation in the monthly proportion of fruits in brown spider monkey's diet throughout this study, there was no relation between habitat‐wide fruit availability and the proportion of fruit included in their monthly diet. The diet of brown spider monkeys at Las Quinchas is toward the high end of fruit intake, even within other wild spider monkeys’ populations, suggesting that these endangered primates might also be facing the challenges of being a large bodied fruit specialist under a regional scenario of habitat loss and fragmentation. Am. J. Primatol. 74:1097‐1105, 2012. © 2012 Wiley Periodicals, Inc.     

Use of agroecosystem matrix habitats by mammalian carnivores (Carnivora): a global‐scale analysis

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Restoration of tree lines in an agricultural landscape: their effectiveness as a conservation management tool

As remnant vegetation covers <15% of the Australian sheep‐wheat belt, it is important to identify conservation strategies suitable for use in agricultural landscapes. Tree lines are widespread ecological structures in rural areas, and are now the subject of government subsidy schemes in New South Wales. However, the contribution of tree lines to biodiversity conservation is poorly understood. To identify the conservation value of tree lines, the bird communities in 36 tree lines in Cowra Shire, New South Wales, were surveyed 4 times each. The results demonstrated that tree lines were used by a large number of species, six of which were threatened. Different taxa were associated with different physical tree line attributes, with tree line age an important predictor of species occurrence. While mature tree lines tended to support more species, as was reflected in higher Shannon Diversity Index scores, they also harboured relatively more introduced species and nest predators, while young tree lines provided the best habitat for threatened species. However, these tree lines will mature, and as they do so they will provide increasingly suitable habitat for the hyperaggressive native honeyeater, the Noisy Miner (Manorina melanocephala), and introduced species. Therefore, tree lines in agricultural landscapes may only be serving an important role for conservation on short time scales, and the suite of threatened species young tree lines currently support appear likely to decline further in the future.     

A Global-Scale Evaluation of Primate Exposure and Vulnerability to Climate Change

Human-induced climate change poses many potential threats to nonhuman primate species, many of which are already threatened by human activities such as deforestation, hunting, and the exotic pet trade. Here, we assessed the exposure and potential vulnerability of all nonhuman primate species to projected future temperature and precipitation changes. We found that overall, nonhuman primates will experience 10 % more warming than the global mean, with some primate species experiencing >1.5 °C for every °C of global warming. Precipitation changes are likely to be quite varied across primate ranges (from >7.5 % increases per °C of global warming to >7.5 % decreases). We also identified individual endangered species with existing vulnerabilities (owing to their small range areas, specialized diet, or restricted habitat use) that are expected to experience the largest climate changes. Finally, we defined hotspots of primate vulnerability to climate changes as areas with many primate species, high concentrations of endangered species, and large expected climate changes. Although all primate species will experience substantial changes from current climatic conditions, our hotspot analysis suggests that species in Central America, the Amazon, and southeastern Brazil, as well as portions of East and Southeast Asia, may be the most vulnerable to the anticipated impacts of global warming. It is essential that impacts of human-induced climate change be a priority for research and conservation planning in primatology, particularly for species that are already threatened by other human pressures. The vulnerable species and regional hotspots that we identify here represent critical priorities for conservation efforts, as existing challenges are expected to become increasingly compounded by the impacts of global warming.     

Species diversity and postcranial anatomy of eocene primates from Shanghuang,China

The middle Eocene Shanghuang fissure‐fillings, located in southern Jiangsu Province in China near the coastal city of Shanghai (Fig. 1 ), contain a remarkably diverse array of fossil primates that provide a unique window into the complex role played by Asia during early primate evolution. 1 Compared to contemporaneous localities in North America or Europe, the ancient primate community sampled at the Shanghuang fissure‐fillings is unique in several ways. Although Shanghuang has some typical Eocene primates (Omomyidae and Adapoidea), it also contains the earliest known members of the Tarsiidae and Anthropoidea (Fig. 2 ), and some new taxa that are not as yet known from elsewhere. It exhibits a large number of primate species, at least 18, most of which are very small (15‐500 g), including some of the smallest primates that have ever been recovered. 2 - 4 © 2012 Wiley Periodicals, Inc.     

Low Primate Diversity and Abundance in Northern Amazonia and its Implications for Conservation

Large areas of the Rio Negro basin in Amazonia are covered by continuous tracts of tropical forest, but have few primate species. This is anomalous considering the general relationship between area and number of species. One possibility is that much of the forest is unsuitable habitat for most primates and the area of suitable habitat is much less than the forested area. This has important consequences for the design of reserves and predictions of the consequences of climate change, which tend to be based on broad categories based on satellite images, and not on information of species distributions within those broad categories. The study was conducted through diurnal and nocturnal line‐transect surveys in the Biodiversity Research Program 25‐km² permanent grid in Viruá National Park, which has vegetation associations typical of much of northern Amazonia. The highest primate diversity and abundances occurred in tall terra firme forests (58%), whereas inundated forests and scrublands, which cover 42 percent of the survey grid and 90.8 percent of the Viruá National Park, have virtually no primates. This suggests that parks and reserves in northwestern Amazonia will have to be very large to maintain viable populations of most primates and their ecological interactions, and that very broad habitat categories are not sufficient to make predictions about actual and future suitability of areas for primate conservation.     

Primates in Human-Modified and Fragmented Landscapes: The Conservation Relevance of Modelling Habitat and Disturbance Factors in Density Estimation

Accurate density estimations of threatened animal populations is essential for management and conservation. This is particularly critical for species living in patchy and altered landscapes, as is the case for most tropical forest primates. In this study, we used a hierarchical modelling approach that incorporates the effect of environmental covariates on both the detection (i.e. observation) and the state (i.e. abundance) processes of distance sampling. We applied this method to already published data on three arboreal primates of the Udzungwa Mountains of Tanzania, including the endangered and endemic Udzungwa red colobus (Procolobus gordonorum). The area is a primate hotspot at continental level. Compared to previous, ‘canonical’ density estimates, we found that the inclusion of covariates in the modelling makes the inference process more informative, as it takes in full account the contrasting habitat and protection levels among forest blocks. The correction of density estimates for imperfect detection was especially critical where animal detectability was low. Relative to our approach, density was underestimated by the canonical distance sampling, particularly in the less protected forest. Group size had an effect on detectability, determining how the observation process varies depending on the socio-ecology of the target species. Lastly, as the inference on density is spatially-explicit to the scale of the covariates used in the modelling, we could confirm that primate densities are highest in low-to-mid elevations, where human disturbance tend to be greater, indicating a considerable resilience by target monkeys in disturbed habitats. However, the marked trend of lower densities in unprotected forests urgently calls for effective forest protection.     

Rarity in the tropics: biogeography and macroecology of the primates

Aim To describe rarity and elucidate its biology in a tropical mammalian order, the Primates. Location Africa, Central and South America, Asia, Madagascar. Methods A review of the literature, with some additional analyses using data from the literature. A variety of definitions of rarity are used in order to describe it and to investigate its biology by correlating the degree of rarity with a variety of biological traits indicative of resource use (e.g. size of annual home range), reproductive rate (e.g. birth interval)and specialization (e.g. number of habitat types used). Results Few primate taxa occur outside the tropics, and most taxa are rare (small geographical range size or latitudinal extent, low density or both). Latitudinal extent is narrower at lower latitudes in Africa and Asia, but the potential resultant packing of taxa appears not to explain the taxonomic diversity gradient. Whilst primate species do not show the common, positive density by range size relationship, primate genera show a significant shallow slope, and primate families/subfamilies a strongly positive slope. Rare taxa are specialized, but neither use more resources nor breed more slowly than common taxa. The correlation of rarity and specialization is via geographical range: taxa with small ranges, or small ranges for their density, are specialized, but not taxa at low density. Common taxa are generalized because they consist of more differently specialized subtaxa, not because each subtaxon is generalized. Main conclusions Most primate taxa are rare, in which case most are presumably likely to go extinct. Rare primates are specialized, but do not necessarily use more resources, nor breed more slowly. Specialization as an explanation for rarity appears to work via constriction of range size, not of density. Common primates might be common (large range size) not because subtaxa or individuals are generalized, but because they are composed of more subtaxa. A consequence could be that persistence of even common taxa will depend on conservation of several populations scattered across the taxon's geographical range.     

Threatened and Endangered Subspecies with Vulnerable Ecological Traits Also Have High Susceptibility to Sea Level Rise and Habitat Fragmentation

The presence of multiple interacting threats to biodiversity and the increasing rate of species extinction make it critical to prioritize management efforts on species and communities that maximize conservation success. We implemented a multi-step approach that coupled vulnerability assessments evaluating threats to Florida taxa such as climate change, sea-level rise, and habitat fragmentation with in-depth literature surveys of taxon-specific ecological traits. The vulnerability, adaptive capacity, and ecological traits of 12 threatened and endangered subspecies were compared to non-listed subspecies of the same parent species. Overall, the threatened and endangered subspecies showed high vulnerability and low adaptive capacity, in particular to sea level rise and habitat fragmentation. They also exhibited larger home ranges and greater dispersal limitation compared to non-endangered subspecies, which may inhibit their ability to track changing climate in fragmented landscapes. There was evidence for lower reproductive capacity in some of the threatened or endangered taxa, but not for most. Taxa located in the Florida Keys or in other low coastal areas were most vulnerable to sea level rise, and also showed low levels of adaptive capacity, indicating they may have a lower probability of conservation success. Our analysis of at-risk subspecies and closely related non-endangered subspecies demonstrates that ecological traits help to explain observed differences in vulnerability and adaptive capacity. This study points to the importance of assessing the relative contributions of multiple threats and evaluating conservation value at the species (or subspecies) level when resources are limited and several factors affect conservation success.     

Threatened woody flora as an ecological indicator of large herbivore introductions

Rewilding and translocations of large herbivores for conservation purposes have increased in recent times, with numerous introductions inside and outside their native range. This study aims to analyze the use of threatened plant taxa as a possible ecological indicator of large herbivore introductions. We examined the effects of a threatened large ungulate, the Barbary sheep (Ammotragus lervia), on both endangered and vulnerable woody taxa after its introduction in 1970. Contrary to our hypothesis, the herbivore impact on threatened woody species was higher than that found on widespread woody plants. The results reveal that 35.7% of the threatened species showed the highest possible level of herbivore damage in contrast to 6.5% for the widespread species. Threatened species were preferred over common plants, probably due to their greater palatability. Overall plant cover, including neighboring species, was also an important factor determining browsing damage and, thus, habitats with low ground cover should be particularly considered in conservation plans. Herbivore damage on common taxa should be taken with caution since they could mask unsustainable herbivore densities for threatened woody taxa or protected habitats. The use of threatened woody taxa through the studied ecological indicators (herbivore damage, plant preferences, habitat use and regeneration success) represented a useful tool to assess the sustainability of large herbivores introductions and to establish a priority conservation ranking for threatened plant species. These findings highlight the deleterious effects of overabundant ungulate populations regardless its origin (exotic or native) and the need of monitoring threatened woody taxa to better estimate the suitability and sustainability of large herbivore introductions.     

Welfare based primate rehabilitation as a potential conservation strategy: does it measure up?

Many primate species are threatened with extinction and are the focus of extensive conservation efforts including re-introduction, captive breeding and habitat conservation. Welfare-based rehabilitation (hereafter also ‘rehabilitation’) is a management strategy commonly used for primates, particularly those species targeted by the pet and bush meat trades. Rehabilitation of rescued primates typically has the dual motivation of welfare and conservation, but has not been assessed as a conservation strategy. As the species involved in rehabilitation are often endangered (e.g. chimpanzees, gorillas, orang-utans), it is important for rehabilitation projects to follow a ‘best practice’ model in order to increase positive outcomes. In this study, we compared the approaches of 28 welfare-based primate rehabilitation projects to the ‘IUCN guidelines for nonhuman primate re-introductions’, in addition to components of the ‘Best practice guidelines for the re-introduction of great apes’ in order to assess where additional work might be needed for released animals to contribute to conservation outcomes. Few projects examined complied with the guidelines for re-introduction, failing to incorporate important factors such as quarantine, long term post-release monitoring and training for predator awareness. Further development of species-specific rehabilitation guidelines may improve the outcomes of future rehabilitation projects. To support this, we recommend that detailed methods and results be published for all rehabilitation efforts, regardless of the outcome.     

Rethinking Primate Origins Again

In 1974, Cartmill introduced the theory that the earliest primate adaptations were related to their being visually oriented predators active on slender branches. Given more recent data on primate‐like marsupials, nocturnal prosimians, and early fossil primates, and the context in which these primates first appeared, this theory has been modified. We hypothesize that our earliest primate relatives were likely exploiting the products of co‐evolving angiosperms, along with insects attracted to fruits and flowers, in the slender supports of the terminal branch milieu. This has been referred to as the primate/angiosperm co‐evolution theory. Cartmill subsequently posited that: “If the first euprimates had grasping feet and blunt teeth adapted for eating fruit, but retained small divergent orbits…” then the angiosperm coevolution theory would have support. The recent discovery of Carpolestes simpsoni provides this support. In addition, new field data on small primate diets, and a new theory concerning the visual adaptations of primates, have provided further evidence supporting the angiosperm coevolution theory.Am. J. Primatol. 75:95‐106, 2013. © 2012 Wiley Periodicals, Inc.     

The primate extinction crisis in China: immediate challenges and a way forward

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.     

Interspecific Interactions between Primates,Birds, Bats,and Squirrels May Affect Community Composition on Borneo

For several decades, primatologists have been interested in understanding how sympatric primate species are able to coexist. Most of our understanding of primate community ecology derives from the assumption that these animals interact predominantly with other primates. In this study, we investigate to what extent multiple community assembly hypotheses consistent with this assumption are supported when tested with communities of primates in isolation versus with communities of primates, birds, bats, and squirrels together. We focus on vertebrate communities on the island of Borneo, where we examine the determinants of presence or absence of species, and how these communities are structured. We test for checkerboard distributions, guild proportionality, and Fox's assembly rule for favored states, and predict that statistical signals reflecting interactions between ecologically similar species will be stronger when nonprimate taxa are included in analyses. We found strong support for checkerboard distributions in several communities, particularly when taxonomic groups were combined, and after controlling for habitat effects. We found evidence of guild proportionality in some communities, but did not find significant support for Fox's assembly rule in any of the communities examined. These results demonstrate the presence of vertebrate community structure that is ecologically determined rather than randomly generated, which is a finding consistent with the interpretation that interactions within and between these taxonomic groups may have shaped species composition in these communities. This research highlights the importance of considering the broader vertebrate communities with which primates co‐occur, and so we urge primatologists to explicitly consider nonprimate taxa in the study of primate ecology. Am. J. Primatol. 75:170‐185, 2013. © 2012 Wiley Periodicals, Inc.