Nutritional Characteristics of Wild and Cultivated Foods for Chimpanzees (<Emphasis Type="Italic">Pan troglodytes</Emphasis>) in Agricultural Landscapes

Primate habitats are being transformed by human activities such as agriculture. Many wild primates include cultivated foods (crops) in their diets, calling for an improved understanding of the costs and benefits of crop feeding. We measured the macronutrient and antifeedant content of 44 wild and 21 crop foods eaten by chimpanzees (Pan troglodytes schweinfurthii) in a mosaic habitat at Bulindi, Uganda, to evaluate the common assertion that crops offer high nutritional returns compared to wild forage for primates. In addition, we analyzed 13 crops not eaten at Bulindi but that are consumed by chimpanzees elsewhere to assess whether nutritional aspects explain why chimpanzees in Bulindi ignored them. Our analysis of their wild plant diet (fruit, leaves, and pith) corresponds with previous chemical analyses of primate plant foods. Compared to wild food equivalents, crops eaten by the chimpanzees contained higher levels of digestible carbohydrates (mainly sugars) coupled with lower amounts of insoluble fiber and antifeedants. Cultivated fruits were relatively nutritious throughout the ripening process. Our data support the assumption that eating cultivated foods confers energetic advantages for primates, although crops in our sample were low in protein and lipids compared to some wild foods. We found little evidence that crops ignored by the chimpanzees were less nutritious than those that they did eat. Nonnutritional factors, e.g., similarity to wild foods, probably also influence crop selection. Whether cultivated habitats can support threatened but flexible primates such as chimpanzees in the long term hinges on local people’s willingness to share their landscape and resources with them.     

Use of wild and cultivated foods by chimpanzees at Bossou,Republic of Guinea: feeding dynamics in a human‐influenced environment

Increased human population growth and more conversions of natural habitat to agricultural land have resulted in greater proximity between humans and nonhuman primate species. Consequent increases in resource competition including crop‐raiding are a by‐product of both natural resources becoming less available and the nutritional benefits of cultivated foods becoming more known to the nonhuman primates. Chimpanzees at Bossou in the Republic of Guinea, West Africa, consume 17 different types of cultivated foods that are grown extensively throughout their small, fragmented home range. Direct observations of feeding behavior conducted over an 18‐month period revealed that during specific months crops account for up to one quarter of chimpanzee feeding time, with higher overall crop‐raiding levels throughout the periods of wild fruit scarcity. Some cultivated foods, especially sugar fruits, are mostly fallback foods, whereas others, such as rice pith (Oryza sp.) and maize (Zea mays), are consumed according to their availability even when wild foods are abundant. These findings highlight the importance of both crop choice by farmers and a thorough understanding of the ecology of resident primate species when establishing land management techniques for alleviating human–primate conflict. Am. J. Primatol. 71:636–646, 2009. © 2009 Wiley‐Liss, Inc.     

Dietary responses to fruit scarcity of wild chimpanzees at Bossou,Guinea: Possible implications for ecological importance of tool use

A 13-month ecological study was conducted at Bossou, Guinea, West Africa, to elucidate how a community of wild chimpanzees (Pan troglodytes verus) deals with the scarcity of main foods. During the study period, fruit availability fluctuated radically. The chimpanzees were confirmed to depend heavily on three “keystone resources” which were available when their main foods (fruit pulp) were scarce. These were fruits of Musanga cecropioides, oil-palm (Elaeis guineensis) nuts, and oil-palm pith. These are abundant in the chimpanzees' home range and their nutritional contents compensate for a decrease in nutritional intake from fruit pulp. The presence of these excellent backup foods may explain the high reproductive performance of Bossou chimpanzees. Here, chimpanzees consumed two of the three keystone foods using two types of tool behavior: nut-cracking for oil-palm nuts and pestle-pounding for oil-palm pith. These tool-using behaviors accounted for 31.9% of the total feeding time spent in June (the month in which the highest frequency occurred) and 10.4% in total for the year. It is suggested that the Bossou chimpanzees depend strongly on tools for their subsistence. This implies a possible function for tool technology in the evolution of our human ancestors. Am J Phys Anthropol 106:283–295, 1998. © 1998 Wiley-Liss, Inc.     

Forest fragments become farmland: Dietary Response of wild chimpanzees (Pan troglodytes) to fast-changing anthropogenic landscapes

Behavioral flexibility, including an ability to modify feeding behavior, is a key trait enabling primates to survive in forest fragments. In human-dominated landscapes, unprotected forest fragments can become progressively degraded, and may be cleared entirely, challenging the capacity of primates to adjust to the changes. We examined responses of wild chimpanzees (Pan troglodytes schweinfurthii) to major habitat change: that is, clearance of forest fragments for agriculture. Over 7 years, fragments in Bulindi, Uganda, were reduced in size by 80%. We compared the chimpanzees’ diet at the start and end of this period of rapid deforestation, using data derived mainly from fecal analysis. Similar to other long-term study populations, chimpanzees in Bulindi have a diverse diet comprising over 169 plant foods. However, extensive deforestation seemed to impact their feeding ecology. Dietary changes after fragment clearance included reduced overall frugivory, reduced intake of figs (Ficus spp.; formerly a dietary “staple” for these chimpanzees), and reduced variety of fruits in fecal samples. Nevertheless, the magnitude of most changes was remarkably minor given the extent of forest loss. Agricultural fruits increased in dietary importance, with crops accounting for a greater proportion of fruits in fecal samples after deforestation. In particular, cultivated jackfruit (Artocarpus heterophyllus) became a “staple” food for the chimpanzees but was scarcely eaten before fragment clearance. Crops offer some nutritional benefits for primates, being high in carbohydrate energy and low in hard-to-digest fiber. Thus, crop feeding may have offset foraging costs associated with loss of wild foods and reduced overall frugivory for the chimpanzees. The adaptability of many primates offers hope for their conservation in fragmented, rural landscapes. However, long-term data are needed to establish whether potential benefits (i.e. energetic, reproductive) of foraging in agricultural matrix habitats outweigh fitness costs from anthropogenic mortality risk for chimpanzees and other adaptable primates.     

Chimpanzees share forbidden fruit

The sharing of wild plant foods is infrequent in chimpanzees, but in chimpanzee communities that engage in hunting, meat is frequently used as a 'social tool' for nurturing alliances and social bonds. Here we report the only recorded example of regular sharing of plant foods by unrelated, non-provisioned wild chimpanzees, and the contexts in which these sharing behaviours occur. From direct observations, adult chimpanzees at Bossou (Republic of Guinea, West Africa) very rarely transferred wild plant foods. In contrast, they shared cultivated plant foods much more frequently (58 out of 59 food sharing events). Sharing primarily consists of adult males allowing reproductively cycling females to take food that they possess. We propose that hypotheses focussing on 'food-for-sex and -grooming' and 'showing-off' strategies plausibly account for observed sharing behaviours. A changing human-dominated landscape presents chimpanzees with fresh challenges, and our observations suggest that crop-raiding provides adult male chimpanzees at Bossou with highly desirable food commodities that may be traded for other currencies.     

Flexible feeding on cultivated underground storage organs by rainforest-dwelling chimpanzees at Bossou, West Africa

It has been proposed that exploitation of underground storage organs (USOs) played an important role in the evolution of the genus Homo, these items serving as ‘fallback foods’ during periods of low food availability. The use of USOs as food by wild chimpanzees is infrequent and seen mostly in populations inhabiting relatively arid environments, such as the savanna. Here, we specifically test the hypothesis that chimpanzees (Pan troglodytes verus) inhabiting tropical wet forest at Bossou (Republic of Guinea, West Africa) exploit USOs as a fallback food during periods of fruit scarcity. Chimpanzees were never observed feeding on wild USOs, that is, those that were never cultivated, and rarely on other underground plant parts. However, direct observations revealed regular consumption of the USOs of cultivated cassava (Manihot esculenta), a spatially abundant and continuously available plant, although the chimpanzees did not use tools when acquiring and feeding on cassava. In agreement with the fallback foods hypothesis, our results show that chimpanzees exploited cassava USOs more frequently when both wild and cultivated fruits were scarce, and consumption patterns of cassava paralleled those of wild fallback foods. These seasonal extractive USO foraging strategies by chimpanzees can strengthen attempts to construct a clearer picture of the importance of USO feeding in hominoid evolution.     

Activity and Habitat Use of Chimpanzees (<Emphasis Type="Italic">Pan troglodytes verus</Emphasis>) in the Anthropogenic Landscape of Bossou,Guinea, West Africa

Many primate populations inhabit anthropogenic landscapes. Understanding their long-term ability to persist in such environments and associated real and perceived risks for both primates and people is essential for effective conservation planning. Primates in forest–agricultural mosaics often consume cultivars to supplement their diet, leading to potentially negative encounters with farmers. When crossing roads, primates also face the risk of encounters with people and collision with vehicles. Chimpanzees (Pan troglodytes verus) in Bossou, Guinea, West Africa, face such risks regularly. In this study, we aimed to examine their activity budget across habitat types and the influence of anthropogenic risks associated with cultivated fields, roads, and paths on their foraging behavior in noncultivated habitat. We conducted 6-h morning or afternoon follows daily from April 2012 to March 2013. Chimpanzees preferentially used forest habitat types for traveling and resting and highly disturbed habitat types for socializing. Wild fruit and crop availability influenced seasonal habitat use for foraging. Overall, chimpanzees preferred mature forest for all activities. They showed a significant preference for foraging at >200 m from cultivated fields compared to 0–100 m and 101–200 m, with no effect of habitat type or season, suggesting an influence of associated risk. Nevertheless, the chimpanzees did not actively avoid foraging close to roads and paths. Our study reveals chimpanzee reliance on different habitat types and the influence of human-induced pressures on their activities. Such information is critical for the establishment of effective land use management strategies in anthropogenic landscapes.     

Dietary Response of Chimpanzees and Cercopithecines to Seasonal Variation in Fruit Abundance. II. Macronutrients

In a continuation of our study of dietary differentiation among frugivorous primates with simple stomachs, we present the first comparison of differences in dietary macronutrient content between chimpanzees and cercopithecine monkeys. Previously we have shown that chimpanzee and monkey diets differ markedly in plant part and species content. We now examine whether this diet diversity is reflected in markedly different dietary macronutrient levels or the different feeding strategies yield the same macronutrient levels in their diets. For each primate group we calculated the total weighted mean dietary content of 4 macronutrients: crude lipid (lipid), crude protein (CP), water-soluble carbohydrates (WSC), and total nonstructural carbohydrates (TNC). We also calculated 4 fiber fractions: neutral-detergent fiber (NDF), which includes the subfractions hemicellulose (HC), cellulose (Cs), and sulfuric acid lignin (Ls). The HC and Cs are potentially fermentable fibers and would contribute to the energy provided by plant food, depending on the hind gut fermenting capacity of the individual primate species. The chimpanzee diet contained higher levels of WSC and TNC because during times of fruit abundance the chimpanzees took special advantage of ripe fruit, while the monkeys did not. The monkey diets contained higher levels of CP because the monkeys consumed a constant amount of leaf throughout the year. All four primate species consumed diets with similar NDF levels. However, the chimpanzees also took advantage of periods of ripe fruit abundance to decrease their Ls levels and to increase their HC levels. Conversely, the monkey diets maintained constant levels of the different fiber fractions thoughout the year. Nevertheless, despite these differences, the diets of the 4 frugivores were surprisingly similar, considering the substantial differences in body size. We conclude that the chimpanzee diet is of higher quality, particularly of lower fiber content, than expected on the basis of their body size.     

Dispersal of a Human-Cultivated Crop by Wild Chimpanzees (<Emphasis Type="Italic">Pan troglodytes verus</Emphasis>) in a Forest–Farm Matrix

With the conversion of natural habitats to farmland, nonhuman primates (hereafter primates) are increasingly exposed to agricultural crops. Although frugivorous primates are important seed dispersers that sometimes feed on agricultural fruits, evidence for dispersal of crops by primates is lacking. Here, we examine flexible feeding on cacao (Theobroma cacao) fruit and seed dispersal patterns by chimpanzees (Pan troglodytes verus) at Bossou in Guinea, and consequent cacao germination and survival. From direct observations, we confirm that cacao fruit is not an important food to chimpanzees, representing 0.23 % of focal animal feeding time. Chimpanzees ingest cacao pulp and either spit out the large seeds intact from unripe cacao fruit or swallow the seeds from ripe cacao fruits, which are consequently deposited in feces. From ecological surveys we show that chimpanzees distributed cacao extensively throughout their home range, at a mean distance of 407 m?±?SE 0.6 (N?=?90 clusters, range: 4–1130 m) from cacao plantations. As distance from the cacao plantation increased, cacao plants were more likely to survive. Other factors, including number of cacao plants in a cluster, plant height, and openness of the understory did not predict short-term cacao survival. Cacao plants within the forest did not produce fruit. By contrast, when chimpanzees deposited seeds in a plantation, cacao plants produced fruits as a result of farmers’ maintenance of the area. Our local-scale findings emphasize the complex behavioral and ecological interconnections between coexisting humans and primates in agricultural landscapes and generate interesting questions regarding primate niche construction and crop “ownership” related to who “plants” the crop.     

The role of non-natural foods in the nutritional strategies of monkeys in a human-modified mosaic landscape

Many tropical animals inhabit mosaic landscapes including human-modified habitat. In such landscapes, animals commonly adjust feeding behavior, and may incorporate non-natural foods. These behavioral shifts can influence consumers' nutritional states, with implications for population persistence. However, few studies have addressed the nutritional role of non-natural food. We examined nutritional ecology of wild blue monkeys to understand how dietary habits related to non-natural foods might support population persistence in a mosaic landscape. We documented prevalence and nutritional composition of non-natural foods in monkey diets to assess how habitat use influenced their consumption, and their contribution to nutritional strategies. While most energy and macronutrients came from natural foods, subjects focused non-natural feeding activity on five exotic plants, and averaged about a third of daily calories from non-natural foods. Most non-natural food calories came from non-structural carbohydrates and least from protein. Consumption of non-natural foods related to time in human-modified habitats, which two groups used non-randomly. Non-natural and natural foods were similar in nutrients, and the amount of non-natural food consumed drove variation in nutritional strategy. When more daily calories came from non-natural foods, females consumed a higher ratio of non-protein energy to protein (NPE:P). Females also prioritized protein while allowing NPE:P to vary, increasing NPE while capitalizing on non-natural foods. Overall, these tropical mammals achieved a similar nutrient balance regardless of their intake of non-natural foods. Forest and forest-adjacent areas with non-natural vegetation may provide adequate nutrient access for consumers, and thus contribute to wildlife conservation in mosaic tropical landscapes.     

Nutrient composition of plants consumed by black and white ruffed lemurs,Varecia variegata,in the Betampona Natural Reserve,Madagascar

The purpose of this study was to quantify the concentrations of crude protein, fat, ash, neutral detergent fiber, acid detergent fiber, lignin, nonstructural carbohydrates, and gross energy in plant foods consumed by wild black and white ruffed lemurs (Varecia variegata). Calcium, phosphorous, magnesium, potassium, sodium, iron, zinc, copper, manganese, molybdenum, and selenium concentrations were also determined. A total of 122 samples from 33 plant families and more than 60 species were collected and analyzed for their nutritional content. The specific nutrient needs of black and white ruffed lemurs are unknown, but quantifying the nutritional composition of the foods they consume in the wild will help nutritionists and veterinarians formulate more appropriate diets for captive ruffed lemurs. This information will also supply information on how man‐induced habitat changes affect the nutritional composition of foods consumed by free‐ranging lemurs. Zoo Biol 29:375–396, 2010. © 2009 Wiley‐Liss, Inc.     

Nutritional composition of the diet of the western gorilla (Gorilla gorilla): Interspecific variation in diet quality

To meet nutritional needs, primates adjust their diets in response to local habitat differences, though whether these dietary modifications translate to changes in dietary nutrient intake is unknown. A previous study of two populations of the mountain gorilla (MG: Gorilla beringei) found no evidence for intraspecific variation in the nutrient composition of their diets, despite ecological and dietary differences between sites. One potential explanation is that nutritional variability in primate diets requires greater ecological divergence than what was captured between MG sites, underpinning environmental differences in the nutrient quality of plant foods. To test whether Gorilla exhibits interspecific variation in dietary composition and nutrient intake, we studied the composition and macronutrients of the western gorilla (WG: Gorilla gorilla) staple diets and compared them with published data from the two MG populations. We recorded feeding time and food intake of four adult female WGs from one habituated group over a period of 11 months (December 2004–October 2005) at the Mondika Research Center, Republic of Congo, allowing for assessment of seasonal patterns of nutrient intake. Staple diets of WGs and MGs diverged in their dietary and macronutrient composition. Compared to MGs, the staple diet of WGs (by intake) contained higher proportions of fruit (43%) and leaf (12%) and a lower proportion of herb (39%), resulting in a higher intake of total nonstructural carbohydrate and fiber and a lower intake of crude protein. Staple gorilla fruits and herbs differed in nutrient quality between sites. Gorillas exhibit nutritional flexibility that reflects ecological variation in the nutrient quality of plant foods. Since dietary quality typically affects rates of growth and reproduction in primates, our results suggest that interspecific differences in nutrient intake and food quality may shape differences in gorilla nutrient balancing and female life history strategies.     

Macronutrient balancing in free‐ranging populations of moose

At northern latitudes, large spatial and temporal variation in the nutritional composition of available foods poses challenges to wild herbivores trying to satisfy their nutrient requirements. Studies conducted in mostly captive settings have shown that animals from a variety of taxonomic groups deal with this challenge by adjusting the amounts and proportions of available food combinations to achieve a target nutrient balance. In this study, we used proportions‐based nutritional geometry to analyze the nutritional composition of rumen samples collected in winter from 481 moose (Alces alces) in southern Sweden and examine whether free‐ranging moose show comparable patterns of nutrient balancing. Our main hypothesis was that wild moose actively regulate their rumen nutrient composition to offset ecologically imposed variation in the nutritional composition of available foods. To test this, we assessed the macronutritional composition (protein, carbohydrates, and lipids) of rumen contents and commonly eaten foods, including supplementary feed, across populations with contrasting winter diets, spanning an area of approximately 10,000 km². Our results suggest that moose balanced the macronutrient composition of their rumen, with the rumen contents having consistently similar proportional relationship between protein and nonstructural carbohydrates, despite differences in available (and eaten) foods. Furthermore, we found that rumen macronutrient balance was tightly related to ingested levels of dietary fiber (cellulose and hemicellulose), such that the greater the fiber content, the less protein was present in the rumen compared with nonstructural carbohydrates. Our results also suggest that moose benefit from access to a greater variety of trees, shrubs, herbs, and grasses, which provides them with a larger nutritional space to maneuver within. Our findings provide novel theoretical insights into a model species for ungulate nutritional ecology, while also generating data of direct relevance to wildlife and forest management, such as silvicultural or supplementary feeding practices.     

Plant foods consumed by Pan: Exploring the variation of nutritional ecology across Africa

It has been shown that differences in resource density and nutrient supply affect variation in ranging patterns, habitat use, and sociality. Among nonhuman primates, chimpanzees (Pan troglodytes) and bonobos (P. paniscus) have often been used as models for the link between social system and habitat ecology. Field reports suggest that resource density is higher in habitats occupied by bonobos (compared to chimpanzee habitats), and in the West (compared to the East) of the range of chimpanzees. In this study we compared diet quality at the level of species and populations using information from nutritional analyses of fruit and leaves consumed by chimpanzees (three) and bonobos (one population). Quality of plant foods was assessed on the basis of a) the concentration of macronutrients, fiber, and anti‐feedants, and b) associations of different nutrient components. Overall plant samples collected at each site differed in terms of macronutrient content. However, nutritious quality and gross energy content of food samples were similar suggesting that dietary quality reflects selectivity rather than habitat ecology. The quality of plant foods consumed by bonobos was within the range of chimpanzees and the quality of plant foods consumed by western chimpanzees was not higher than that of eastern chimpanzees. While the results showed significant variation across forests inhabited by Pan, they did not match with geographical patterns between and within Pan species as proposed in previous studies. This suggests that the nutritional quality of the habitat is not always a reliable predictor of the quality of the diet. Am J Phys Anthropol 2010. © 2009 Wiley‐Liss, Inc.     

Primate Crop Feeding Behavior,Crop Protection,and Conservation

Many species across a range of primate genera, irrespective of dietary and locomotory specializations, can and will incorporate agricultural crops in their diets. However, although there is little doubt that rapid, extensive conversion of natural habitats to agricultural areas is significantly impacting primate populations, primate crop foraging behaviors cannot be understood solely in terms of animals shifting to cultivated crops to compensate for reduced wild food availability. To understand fully why, how, and when primates might incorporate crops in their dietary repertoire, we need to examine primate crop foraging behavior in the context of their feeding strategies and nutritional ecology. Here I briefly outline current debates about the use of terms such as human–wildlife conflict and crop raiding and why they are misleading, summarize current knowledge about primate crop foraging behavior, and highlight some key areas for future research to support human–primate coexistence in an increasingly anthropogenic world.     

Functional macronutritional generalism in a large omnivore,the brown bear

We combine a recently developed framework for describing dietary generalism with compositional data analysis to examine patterns of omnivory in a large widely distributed mammal. Using the brown bear (Ursus arctos) as a model species, we collected and analyzed data from the literature to estimate the proportions of macronutrients (protein, carbohydrate, and lipid) in the diets of bear populations. Across their range, bears consumed a diversity of foods that resulted in annual population diets that varied in macronutrient proportions, suggesting a wide fundamental macronutrient niche. The variance matrix of pairwise macronutrient log‐ratios indicated that the most variable macronutrient among diets was carbohydrate, while protein and lipid were more proportional or codependent (i.e., relatively more constant log‐ratios). Populations that consumed anthropogenic foods, such agricultural crops and supplementary feed (e.g., corn), had a higher geometric mean proportion of carbohydrate, and lower proportion of protein, in annual diets. Seasonally, mean diets were lower in protein and higher in carbohydrate, during autumn compared to spring. Populations with anthropogenic subsidies, however, had higher mean proportions of carbohydrate and lower protein, across seasons compared to populations with natural diets. Proportions of macronutrients similar to those selected in experiments by captive brown bears, and which optimized primarily fat mass gain, were observed among hyperphagic prehibernation autumn diets. However, the majority of these were from populations consuming anthropogenic foods, while diets of natural populations were more variable and typically higher in protein. Some anthropogenic diets were close to the proportions selected by captive bears during summer. Our results suggest that omnivory in brown bears is a functional adaptation enabling them to occupy a diverse range of habitats and tolerate variation in the nutritional composition and availability of food resources. Furthermore, we show that populations consuming human‐sourced foods have different dietary macronutrient proportions relative to populations with natural diets.     

Diet and Feeding Ecology of Chimpanzees (Pan troglodytes) in Bulindi, Uganda: Foraging Strategies at the Forest–Farm Interface

Wild animals increasingly inhabit human-influenced environments such as forest fragments amid agricultural systems. Dietary studies provide a means of assessing wildlife responses to anthropogenic habitat changes. Chimpanzees are specialist frugivores that consume other plant parts, e.g., fibrous pith and leaves, in greater amounts during fruit shortages. I examined the plant diet and seasonal foraging strategies of chimpanzees inhabiting small forest fragments within a cultivated landscape in Uganda. I determined diet over 13 mo via systematic fecal analysis, supplemented by direct observation and feeding trace evidence. I identified important foods and examined their role as seasonal fallbacks. Diet composition and breadth were overall species typical. Chimpanzees were highly frugivorous and the fruit component of fecal samples exceeded that of nonfruit fiber in all months. Forest fruit availability fluctuated seasonally, including a 3-mo low fruiting season, when overall fruit intake declined. During this time chimpanzees pursued a mixed strategy of increasing fiber consumption and feeding more heavily on energy-rich cultivars, including those obtained through crop raiding. The data suggest that exploiting agricultural fruits helped chimpanzees maintain a fruit-dominated diet when forest fruit was scarce. No evidence suggested this disturbed forest–farm mosaic is a food-impoverished habitat for chimpanzees overall. Nevertheless, cultivar feeding creates conflict with people and the high nutritional quality of crops is likely offset by the inherent risk associated with obtaining them. This study adds to growing evidence of ecological and behavioral adaptability of Pan troglodytes in response to anthropogenic habitat alteration. Targeted conservation of key natural foods for wildlife —particularly fallbacks— would help reduce conflicts and improve the survival prospects of threatened species sharing environments with people.     

Ecological correlates of trophic status and frugivory in neotropical primates

Primates are among the most observable and best studied vertebrate order in tropical forest regions, with widespread attention dedicated to the feeding ecology of wild populations. In particular, primates play a key role as frugivores and seed‐dispersal agents for a myriad of tropical plants. Sampling effort by primatologists, however, has been unequally distributed, hampering quantitative comparisons of primate diets. We provide the first systematic review of primate diets, with an emphasis on frugivory, using a comprehensive compilation of 290 unique primate dietary studies from 164 localities in 17 countries across the entire Neotropical realm. We account for sampling effort (standardised as hours) in comparing the richness of fruiting plants recorded in primate diets, and the relative contribution of frugivory to the overall diet in relation to key life‐history traits, such as body mass. We find strong support for the long‐held hypothesis, based on Kay's Threshold, that body size imposes an upper limit on insectivory and a lower limit on folivory, and therefore that frugivory is most important at intermediate body sizes. However, the upper body mass limit of extant neotropical primates, truncated by the post‐Pleistocene megafaunal overkill, has implications for the extent of the frugivory–folivory continuum in extinct lineages. Contemporary threats faced by the largest primates serve as a further warning that the feeding ecology and diet of all neotropical primates remain severely undersampled with regard to the composition and richness of fruits consumed. Indeed, frugivorous primates expected to have the most species‐rich plant diets are amongst those most poorly sampled, exposing implications for our current understanding of primate–plant interaction networks.     

The ‘omnivorous badger dilemma’: towards an integration of nutrition with the dietary niche in wild mammals

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Estrogenic plant foods of red colobus monkeys and mountain gorillas in Uganda

Phytoestrogens, or naturally occurring estrogen-mimicking compounds, are found in many human plant foods, such as soybeans (Glycine max) and other legumes. Because the consumption of phytoestrogens may result in both health benefits of protecting against estrogen-dependent cancers and reproductive costs of disrupting the developing endocrine system, considerable biomedical research has been focused on the physiological and behavioral effects of these compounds. Despite this interest, little is known about the occurrence of phytoestrogens in the diets of wild primates, nor their likely evolutionary importance. We investigated the prevalence of estrogenic plant foods in the diets of two folivorous primate species, the red colobus monkey (Procolobus rufomitratus) of Kibale National Park and mountain gorilla (Gorilla beringei) of Bwindi Impenetrable National Park, both in Uganda. To examine plant foods for estrogenic activity, we screened 44 plant items (species and part) comprising 78.4% of the diet of red colobus monkeys and 53 plant items comprising 85.2% of the diet of mountain gorillas using transient transfection assays. At least 10.6% of the red colobus diet and 8.8% of the gorilla diet had estrogenic activity. This was mainly the result of the red colobus eating three estrogenic staple foods and the gorillas eating one estrogenic staple food. All estrogenic plants exhibited estrogen receptor (ER) subtype selectivity, as their phytoestrogens activated ERβ, but not ERα. These results demonstrate that estrogenic plant foods are routinely consumed by two folivorous primate species. Phytoestrogens in the wild plant foods of these two species and many other wild primates may have important implications for understanding primate reproductive ecology.