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Baboons are well studied in savannah but less so in more closed habitats. We investigated predation on mammals by olive baboons (Papio anubis) at a geographical and climatic outlier, Gashaka Gumti National Park (Nigeria), the wettest and most forested site so far studied. Despite abundant wildlife, meat eating was rare and selective. Over 16 years, baboons killed 7 bushbuck (Tragelaphus scriptus) and 3 red-flanked duiker (Cephalophus rufilatus), mostly still-lying ‘parked' infants. Taking observation time into account, this is 1 predation per group every 3.3 months - far lower than at other sites. Some features of meat eating resemble those elsewhere; predation is opportunistic, adult males monopolize most prey, a targeted killing bite is lacking and begging or active sharing is absent. Carcass owners employ evasive tactics, as meat is often competed over, but satiated owners may tolerate others taking meat. Other features are unusual; this is only the second study site with predation records for bushbuck and the only one for red-flanked duiker. The atypical prey and rarity of eating mammals probably reflects the difficulty of acquiring prey animals when vegetation cover is dense. Our data support the general prediction of the socioecological model that environments shape behavioural patterns, while acknowledging their intraspecific or intrageneric plasticity.

Spatial position within a group affects the value of group-living benefits such as reduced predation risk and improved foraging. The threat of predation, poor nutrition or increased competition from conspecifics can all cause stress. In many species, central positions are known to be more beneficial than peripheral positions in terms of reduced predation, vigilance and foraging. In this study, we examine whether spatial position within a group is associated with stress and anxiety in a troop of olive baboons (Papio anubis). We predicted that the benefits of occupying central positions would be reflected by a reduction in stress and anxiety for animals who spent the most time in the centre of the group. The study subjects appeared to compete actively for the centre of the group. Physiological stress measures (faecal glucocorticoid concentrations) were positively correlated with time spent in central positions. Time spent in central positions was positively correlated with proximity but negatively correlated with vigilance behaviours (alarm barks). Vigilance rates were positively correlated with measures of anxiety (self-scratch frequency). It is suggested that individuals experience chronic stress due to proximity to conspecifics in central positions, whilst perceived predation risk causes anxiety, with perceived predation risk experienced more by individuals on the periphery.

A female mammal's reproductive function and output are limited by the energy she is able to extract from her environment. Previous studies of the interrelationships between energetic circumstances and reproductive function in a variety of mammal species have produced varied results, which do not all support the common assumption that higher female reproductive hormone levels, specifically progesterone, indicate better ovarian function and greater reproductive potential, and are associated with lower energetic stress. In the present study faecal progesterone and glucocorticoid levels were assessed in two troops of olive baboons (Papio anubis) in the same population. They face similar ecological challenges, except that one troop crop-raids, potentially affecting its energetic intake and stress levels. The energy intake of individual females was assessed by combining detailed feeding observations with nutritional analysis of food samples. The crop-raiding troop experienced 50% higher energy intake rates and 50% lower glucocorticoid levels compared to the non-crop-raiding troop alongside substantially lower progesterone levels. This suggests that energetic stress is associated with elevated progesterone levels and may be the cause of the non-crop-raiding troop's lower reproductive output. By comparing groups which differ little, except in terms of food access, and also by directly assessing energy intake, our study addresses some of the design limitations of previous research investigating variation in progesterone levels and energetic stress. It therefore has the potential to contribute to greater understanding of the factors affecting differences in reproductive and stress hormone levels and reproductive function in mammals experiencing different energetic circumstances.

Morphological adaptations related to food processing generally reflect those elements of the diet that represent the greatest biomechanical challenge or that numerically dominate the diet. However, in periods of the annual cycle when the availability of such foods is low, items to which a species has low apparent morphological adaptation may be included in the diet. Here we test the responses of a diet-specialist primate to limitations in the supply of the resource it is specialized to exploit. Uacaris are primarily predators of immature seeds, in seasonally flooded forests in Amazonian Brazil, and have dental specializations to open hard-shelled fruits. We investigated the importance of arthropods in the diet of golden-backed uacaris (Cacajao melanocephalus ouakary), examining their seasonal importance in the uacari diet, and the ways C. m. ouakary used to access them. Using scan and ad libitum sampling of feeding and phenology from botanical study plots to assess fruit availability, we conducted an 18-mo study in Jau National Park, Amazonas State, Brazil. We recorded arthropod predation 298 times, with Cacajao melanocephalus ouakary feeding on 26 invertebrate taxa in ≥11 families and 9 different orders. Uacaris extracted wood-boring beetles dentally from rotting wood and smaller larvae from twigs, stems, and petioles, but this food class did not predominate. This food class (encapsulated foods) constituted 23.4 % of the arthropod records. The majority of arthropod food items were either manually removed from substrates (ants, beetle larvae, caterpillars, fulgorid bugs, grasshoppers, mayflies, spiders, termites, wasps, and a whip-scorpion) or plucked from the air (volant Lepidoptera). Uacaris appeared to avoid toxic caterpillars. Insectivory was most frequent when fruit and seeds were least available. Arthropods seem to be seasonally important to this primate, supplementing or making up for shortfalls in the hard fruits and immature seeds for which uacaris have highly developed dental, and possibly intestinal, adaptations.

Individuals under stress and those of intrinsically low quality may have insufficient resources to invest in developing equally on both sides. Such individuals may thus display higher levels of fluctuating asymmetry (FA)—small, random departures from perfect bilateral symmetry. These small deviations from symmetry are often less than 1 % of trait size, similar to variation in symmetry generated by measurement error. Studies regarding FA must thoroughly assess measurement error if reliable conclusions regarding FA and fitness parameters are to be drawn. Although many studies regarding facial symmetry in humans have been conducted, few have been carried out on non-human primates. The primary aim of this study was to assess whether facial FA in a non-human primate, the olive baboon (Papio anubis) can be reliably measured from digital photographs. Facial FA was measured in three sets of bilaterally symmetrical landmarks from digital images of 35 olive baboons at Gashaka-Gumti National Park, Nigeria. Measurement error was found to be low indicating a high level of reliability based on comparisons of values from two photos of the same individual (r = 0.85). Measurement error was found to be related to the size of the face. This suggests that orientation error increases in large faces and could potentially influence FA scores. These findings highlight the need for FA studies to measure and control error caused by orientation and position of subjects if FA is to be of utility to behavioural ecology and conservation (e.g. if FA reflects whether individuals in a population are suffering from environmental or genetic stress).

We analyse the behaviour of Cacajao melanocephalus ouakary feeding at patches of germinating seedlings in dried-out flooded forest. Seedlings of Eschweilera tenuifolia (Lecythidaceae) were the most commonly eaten (88.9%). Some seed patches were revisited over several days, while others were consistently ignored. We tested 3 predictions relating uacari terrestrial foraging behaviour to: (1) arboreal escape route proximity, (2) seed patch size choice and (3) temporal patterns of repeat exploitation. Comparison of fed-at and ignored patches revealed significant preferences for larger patches, and for those close to arboreal refuges but distant from dense ground-based vegetation. Support for these predictions is interpreted as evidence for predation risk-sensitive foraging.

Predation of tropical arboreal mammals is rarely observed. Here, we report the first observation of predation of a member of the genus Cacajao, and add to the knowledge of eagle predation in the genus Cebus. We compare responses to predation by Cacajao ouakary and Cebus albifrons with previous studies of congenerics and with other close relatives, and compare the alarm vocalizations of C. ouakary with those of other pitheciines. Reactions to non-predating raptors are reported. Constraints on primate anti-predator responses in flooded and non-flooded habitats are considered.

Sexual selection has driven the evolution and elaboration of a wide variety of displays and ornaments in male nonhuman primates, including capes, cheek flanges, and sexual coloration. Among the most sexually dimorphic of all primates is the drill (Mandrillus leucophaeus), the males of which can be 3 times the mass of females, possess large canines, and exhibit extremely bright sexual skin coloration. However, the function of male coloration in this species has never been examined. Here, we present data on male color (measured objectively using digital photography), dominance rank, measures of male-female association, and key sexual behaviors, of adult male drills (n = 17) living in four semi free-ranging enclosures at the Drill Rehabilitation and Breeding Center in Nigeria. We test the hypothesis that male coloration is a badge of status, indicating dominance rank, and the hypothesis that male coloration attracts females. We found that male coloration did indicate rank, and that high ranking, strongly colored males were more likely to associate with adult females, and more specifically with fully tumescent females. These males also engaged in more sexual activity. However, measures of male-female association and sexual behaviors were not related to male color once rank had been taken into consideration; i.e., for males of a given rank, females did not prefer those that were more colorful. We discuss the results in light of what is known about the wild drill social system, in which unfamiliar individuals may come together in “supergroups,” and in which quality indicators of competitive ability may be particularly important.

We compared parasitic infection in a crop-raiding and a wild-foraging troop of olive baboons, Papio anubis, in Gashaka Gumti National Park, Nigeria, to quantify how crop raiding may have influenced primate-parasite interactions. We recovered gastrointestinal parasites from fecal samples from all adult individuals in both troops and processed them via formal-ether sedimentation. We compared parasitic species richness, prevalence, output, and load across troops. We recovered 9 parasite taxa. The wild-foraging troop had a significantly higher mean output than the crop-raiding troop for Physaloptera sp., Trichuris sp., and also a significantly higher total helminth load. The crop-raiding troop had a significantly higher mean output for the protozoan Balantidium coli and also showed a higher parasitic species richness, with 9 species recovered compared to the 7 recorded for the wild-foraging individuals. The changes in nutritional intake, behavior, and human proximity caused by crop raiding may have important epidemiological impacts on wild primate populations, and the nature of such impacts may vary across different taxa of parasites.

In order to maintain regulatory processes, animals are expected to be adapted to the range of environmental stressors usually encountered in their environmental niche. The available capacity of their stress responses is termed their reactive scope, which is utilised to a greater or lesser extent to deal with different stressors. Typically, non-invasive hormone assessment is used to measure the physiological stress responses of wild animals, but, for methodological reasons, such measurements are not directly comparable across studies, limiting interpretation. To overcome this constraint, we propose a new measure of the relative strength of stress responses, ‘demonstrated reactive scope’, and illustrate its use in a study of ecological correlates (climate, food availability) of faecal glucocorticoid (fGC) levels in two forest-living troops of baboons. Results suggest the wild-feeding troop experiences both thermoregulatory and nutritional stress, while the crop-raiding troop experiences only thermoregulatory stress. This difference, together with the crop-raiding troop’s lower overall physiological stress levels and lower demonstrated fGC reactive scope, may reflect nutritional stress-buffering in this troop. The relatively high demonstrated fGC reactive scope levels of both troops compared with other baboons and primate species, may reflect their extreme habitat, on the edge of the geographic range for baboons. Demonstrated reactive scope provides a means of gauging the relative strengths of stress responses of individuals, populations, or species under different conditions, enhancing the interpretive capacity of non-invasive studies of stress hormone levels in wild populations, e.g. in terms of animals’ adaptive flexibility, the magnitude of their response to anthropogenic change, or the severity of impact of environmental conditions.

Few mammalian orders carry their infants clinging to the mother's fur. I investigated the evolution of carrying behavior in primates and the life-history and ecological correlates of infant care patterns. Primates are ideal for the study as there is variation in infant care patterns. Primate infants are left hidden in nests or parked in trees, both of which strategies I term parking, and are carried orally or ride clinging to the mother's fur: riding. Infant carrying has evolved several times in the Primates and, once evolved, it has been conserved. Significant energetic costs of riding are indicated as riding species maintain smaller home ranges than those of non-riders of the same body size. With body size and phylogenetic influences taken into account, riders appear to incur a reproductive cost by weaning and breeding later than parkers. Although riders do not have lower birth rates than those of parkers, their later age at first reproduction leads to their having a lower reproductive rate, measured by the intrinsic rate of population increase. Precociality of infants is not correlated with either riding or nesting behavior. Although non-nesting species have larger litter sizes, their infants are not significantly smaller, nor are their neonatal brains relatively smaller. Although riding may have some energetic and reproductive costs, its repeated evolution in the Primates suggests that it also has some benefits, the most likely being a reduced mortality risk for carried infants.

The amount of non-maternal care (allocare) found in primates varies greatly from species to species. Our paper examines this variation and focuses on possible reasons why mothers in some anthropoid primate species are prepared to relinquish their infants to other carers whereas others are not. We use data collected from the literature to test a number of hypotheses that attempt to explain the observed variation in non-maternal care. Analyses were carried out using comparative methods that control for the influence of both body size and phylogeny on life-history and ecological variation. The results support previous studies of both primates and other mammals in finding a clear link between the amount of allocare provided and female reproductive rates. Species with high allocare levels grow rapidly post-natally and wean their infants at a younger age (but at the same relative size) than species of the same body size with lower allocare levels. This early weaning allows high allocare species to support higher birth rates than low allocare species but does not result in earlier maturation. Our results, therefore, suggest that mothers allow non-maternal care of their infants in order to increase their own reproductive output. It is not clear whether such a strategy also benefits the infant, as we could find no link between the presence of allocare and early maturation (which would increase the infant’s reproductive output) or between allocare levels and infant survival (as measured by vulnerability to infanticide). This suggests that mothers and infants might come into conflict over transfer to other carers, as the benefits to the mother may not be accompanied by benefits to the infant. However, although mothers may benefit from allocare in some circumstances, they will not be expected to allow allocare if the costs are high (e.g. if there is a high risk to the infant).

Simple Summary Wild chimpanzee populations are dramatically declining due to anthropogenic pressure. One way of increasing wild population numbers and/or repopulating areas where local extinction has occurred is to release captive animals. HELP Congo was the first project to successfully release wild-born orphan chimpanzees in their natural environment. We studied the behaviour of eight released chimpanzees over eight years. Over time, they modified their behaviour, suggesting long-term behavioural and ecological adaptations. This suggests that successful release programmes may reinforce existing populations of endangered species. Abstract Wild chimpanzee populations are still declining due to logging, disease transmission and hunting. The bushmeat trade frequently leads to an increase in the number of orphaned primates. HELP Congo was the first project to successfully release wild-born orphan chimpanzees into an existing chimpanzee habitat. A collection of post monitoring data over 16 years now offers the unique opportunity to investigate possible behavioural adaptations in these chimpanzees. We investigated the feeding and activity patterns in eight individuals via focal observation techniques from 1997–1999 and 2001–2005. Our results revealed a decline in the number of fruit and insect species in the diet of released chimpanzees over the years, whereas within the same period of time, the number of consumed seed species increased. Furthermore, we found a decline in time spent travelling, but an increase in time spent on social activities, such as grooming, as individuals matured. In conclusion, the observed changes in feeding and activity patterns seem to reflect important long-term behavioural and ecological adaptations in wild-born orphan released chimpanzees, demonstrating that the release of chimpanzees can be successful, even if it takes time for full adaptation.

We carried out a study of Hanuman langurs in Sariska Tiger Reserve, Rajasthan, India during three months in 1990. The area surveyed included habitat disturbed by human activity, with both the grazing of livestock and firewood collecting resulting in degraded forest. Langur population density was estimated to be between 19 and 36 animals/km2. The density of langurs in disturbed areas was significantly lower than in undisturbed areas. Both tree cover, total tree, and shrub cover were positively correlated with langur density, suggesting that a lack of trees in disturbed areas may have caused the lower langur densities. Although bisexual groups were significantly more common in undisturbed areas the distribution of all male groups did not appear to be affected by human disturbance, suggesting that male bands are likely to inhabit more disturbed habitats than bisexual groups. We suggest that either a lack of food trees or the high density of predators in Sariska may prevent bisexual groups from inhabiting areas where tree cover is low, although the more mobile male groups can survive in these areas. Although Hanuman langurs are a relatively common species in India, these results suggest that they are not, as is commonly assumed, immune to the effects of human disturbance. In areas where langurs appear to survive in disturbed habitats the different response of bisexual and all male groups to disturbance may still result in serious disruption to the population structure.

A take-over and infanticide were observed in a South Indian population of Hanuman langurs (Presbytis entellus). A one-male, bisexual troop of langurs (Troop B1) was attacked by a band of two adult males and one subadult male. During the take-over, the resident male and one elderly female were driven from the troop. The youngest infant, about two months old, was fatally wounded by an attack from the dominant incoming male (S1). However, other young infants in Troop B1 were left unharmed. All three incoming males remained in Troop B1 but the troop appeared to remain functionally one-male, as S1 did not allow the other adult male to copulate. There was no sign that S1 attempted to eject the other two males from troop B1. © 1993 Wiley-Liss, Inc.

In Amazonian seasonally flooded forest (igapo), golden-backed uacaris, Cacajao melanocephalus ouakary, show high selectivity for sleeping trees. Of 89 tree species in igapo, only 16 were used for sleeping (18%). Hydrochorea marginata (Fabaceae) and Ormosia paraensis (Fabaceae) were used most frequently (41% of records) despite being uncommon (Ivlev electivity ratios were 0.76, and 0.84, respectively), though the third most commonly used species (11%), Amanoa oblongifolia (Euphorbiaceae), was selected at near parity. All three species have broad, open canopies with large horizontal limbs and uncluttered interiors. Compared with random trees, sleeping trees had above average diameter at breast height (DBH) and height, lacked lianas and wasp nests, and were more frequently within 5 m of open water. Uacaris generally slept one adult per tree or widely separated in the same canopy and on the outer third of the branch. These behaviours are interpreted as maximising detection of both aerial and arboreal predators.

Although many studies have analyzed the causes and consequences of social relationships, few studies have explicitly assessed how measures of social relationships are affected by the choice of behaviors used to quantify them. The use of many behaviors to measure social relationships in primates has long been advocated, but it was analytically difficult to implement this framework into primatological work. However, recent advances in social network analysis (SNA) now allow the comparison of multiple networks created from different behaviors. Here we use our database of baboon social behavior (Papio anubis, Gashaka Gumti National Park, Nigeria) to investigate (i) to what extent social networks created from different behaviors overlap, (ii) to what extent individuals occupy similar social positions in these networks and (iii) how sex affects social network position in this population of baboons. We used data on grooming, aggression, displacement, mounting and presenting, which were collected over a 15-month period. We calculated network parameters separately for each behavior. Networks based on displacement, mounting and presenting were very similar to each other, whereas grooming and aggression networks differed both from each other and from mounting, displacement and presenting networks. Overall, individual network positions were strongly affected by sex. Individuals central in one network tended to be central in most other networks as well, whereas other measures such as clustering coefficient were found to vary depending on the behavior analyzed. Thus, our results suggest that a baboon's social environment is best described by a multiplex network based on affiliative, aggressive and sexual behavior. Modern SNA provides a number of useful tools that will help us to better understand animals' social environment. We also discuss potential caveats related to their use. Am. J. Primatol. 73:775–789, 2011. © 2011 Wiley-Liss, Inc.

Macaques are found both in broadleaf evergreen forest and in more variable habitats. The former group might be expected to be subject to less variability in their environment and hence to suffer lower rates of density independent mortality. Life history evolution models predict that species in such conditions will have lower rates of development and breeding than those found in more variable habitats where density independent mortality is high. This prediction is tested here by comparing the breeding and development rates of nine species of macaque. Although measures of developmental rate are not found to vary in a predictable way with habitat, measures of breeding rate do correlate with habitat categories used. As predicted, species that are found in more variable habitats tend to have higher birth rates and a higher intrinsic rate of natural increase than do species in more stable, forest habitats. Contrary to prediction selection does not always act to produce an early age at first reproduction in macaques living in seasonal environments. This is discussed with relation to physiological and environmental constraints.

The relationship between basal metabolic rate (BMR), body weight and diet is examined for primates. Contrary to the results reported in several recent works, there is no strong evidence that diet is directly linked to BMR, although a low BMR, relative to body weight, may be found in species with folivorous diets. There is some evidence that nocturnal haplorhine species have a relatively low BMR, but strepsirhines appear to have a uniformly low relative BMR regardless of their primary activity period. The evolution of BMR in primates is discussed in the light of these findings. Some predictions are made about the relative BMRs that should be found for other species whose BMR is, as yet, unknown.

Baboons are the most successful and ubiquitous African primates, renowned for their behavioral and reproductive flexibility, which enable them to inhabit a wide variety of habitat types. Owing to a number of long-term field studies, comparative behavioral, developmental, demographic, and life-history data are available from several populations, but study sites show a heavy bias toward South and East African savannahs, with little research in West or Central Africa. Life-history data from such areas are important if we are fully to understand the nature of the environmental factors that limit baboon distribution. Here, we present demographic data for olive baboons at Gashaka-Gumti National Park (GGNP), Nigeria, collected from December 2000-February 2006, and use these data to test comparative models of baboon life-history. The GGNP habitat, which includes large areas of rainforest, is an environment in which baboons are little studied, and rainfall is much higher than at previous study sites. GGNP troop size data are presented from censuses, as well as life-history data for two troops, one of which is within the park and wild-feeding (Kwano troop), whereas the other dwells at the park edge, and supplements its diet by crop-raiding (Gamgam troop). Troop sizes at GGNP are small compared with other field sites, but fit within previously suggested ranges for baboons under these climatic conditions. Inter-birth intervals in Kwano troop were long compared with most studied populations, and values were not as predicted by comparative models. Consistent with known effects of food enhancement, Gamgam troop experienced shorter inter-birth intervals and lower infant mortality than Kwano troop. We indicate some possible factors that exclude baboons from true rainforest, and suggest that the clearing of forests in Central and West Africa for agricultural land may allow baboons to extend their range into regions from which they are currently excluded.

Allometric methods are used to investigate the life history patterns of New World monkeys. These patterns are compared to those found in other haplorhines and links with ecology are explored. The life history characteristics of the cebid monkeys suggest that these species are primarily slow breeding and slow developing for primates of their size. I suggest that these characteristics arose during a long period of adaptation to stable rainforest habitats. The capuchins and squirrel monkeys are a notable exception to the general rule that genera found outside primary forests will have a “fast” life history. In the marmosets and tamarins selection for small size and high birth rate has occurred, probably as a result of their adapting to more secondary habitats. It is this selection for rapid breeding, combined with the preadaptation of paternal care, that has probably led to twinning in these species. The evidence presented indicates that it is unlikely that twinning arose as a direct result of the mechanical constraints of small size but is more probably due to selection for increased reproductive output.

Baboon researchers commonly use the timing of sexual swelling detumescence to infer the timing of ovulation. These estimates are then used for a variety of purposes, including the interpretation of male and female behaviour, assessment of likely paternity, and the calculation of gestation lengths. Although captive studies have measured the timing of ovulation with respect to detumescence, this has not been measured in wild baboons. Here, we use observational data on female sexual swellings, and hormonal data measured from faeces, to investigate the timing of ovulation with respect to detumescence in wild olive baboons. Our data show similar variability in the timing of ovulation with respect to detumescence as that produced by captive studies, although there are some notable differences, including the absence of any ovulations measured on or after the day of swelling detumescence. We discuss the importance of our results for baboon researchers using swellings to infer ovulation as part of their studies.

An all-male band of 15 Hanuman langurs alarm barked and mobbed a leopard in Sariska Tiger Reserve, a forest in Rajasthan, India. After initially alarm calling and barking in response to the leopard's presence five animals persistently followed and watched the leopard for about an hour. During this time they were barking and tooth grinding. Both the initial alarm calling and the following mobbing can be interpreted as altruistic behaviour. However, unlike the situation in bisexual troops, Hanuman langur males may not be related to other members of the group and the evolution of such behaviour cannot necessarily be explained by kin selection.

Many primate species have a natal coat coloured differently from the coat colour of adults and, in some cases, the natal coat is very different from that of older animals with the infant being conspicuous and/or brightly coloured. This study uses data from 82 anthropoid species to investigate possible correlates of conspicuous natal coat colour (CNC), using comparative, phylogenetically controlled analyses. Five hypotheses were tested using statistical tests on both species data and phylogenetically controlled data. The analyses show some support for the hypothesis that CNCs have evolved to promote allocare but the degree of CNC does not correlate with predation rates, arboreality, or relative testes size.

The intrinsic rate of natural increase (rmax) is calculated for 58 primate species. It is found that this parameter is negatively correlated with body weight, so that larger primate species consistently have a lower rmax than do smaller species. Although there is no apparent link between the raw value of rmax and environmental predictability, a relationship between a high rmax, relative to body weight, and an unpredictable environment is found to exist. However, there is no relationship between a predictable environment and either the raw value of rmax, or the relative rmax. After body size effects are removed, rmax is not correlated with basal metabolic rate (BMR). Pre-natal maternal investment (MI) is found to correlate highly with BMR, even after the removal of body weight effects. MI does not correlate either with rmax or with environmental predictability. Diet appears to have little influence either on rmax on MI, but there is some indication that folivores have both a high relative rmax and a high relative MI. These results are discussed in the light of theories of life-history strategy evolution.