CD4 receptor diversity represents an ancient protection mechanism against primate lentiviruses.

Infection with human and simian immunodeficiency viruses (HIV/SIV) requires binding of the viral envelope glycoprotein (Env) to the host protein CD4 on the surface of immune cells. Although invariant in humans, the Env binding domain of the chimpanzee CD4 is highly polymorphic, with nine coding variants circulating in wild populations. Here, we show that within-species CD4 diversity is not unique to chimpanzees but found in many African primate species. Characterizing the outermost (D1) domain of the CD4 protein in over 500 monkeys and apes, we found polymorphic residues in 24 of 29 primate species, with as many as 11 different coding variants identified within a single species. D1 domain amino acid replacements affected SIV Env-mediated cell entry in a single-round infection assay, restricting infection in a strain- and allele-specific fashion. Several identical CD4 polymorphisms, including the addition of N-linked glycosylation sites, were found in primate species from different genera, providing striking examples of parallel evolution. Moreover, seven different guenons (Cercopithecus spp.) shared multiple distinct D1 domain variants, pointing to long-term trans-specific polymorphism. These data indicate that the HIV/SIV Env binding region of the primate CD4 protein is highly variable, both within and between species, and suggest that this diversity has been maintained by balancing selection for millions of years, at least in part to confer protection against primate lentiviruses. Although long-term SIV-infected species have evolved specific mechanisms to avoid disease progression, primate lentiviruses are intrinsically pathogenic and have left their mark on the host genome.

The evolution of masturbation is associated with postcopulatory selection and pathogen avoidance in primates.

Masturbation occurs throughout the animal kingdom. At first glance, however, the fitness benefits of this self-directed behaviour are unclear. Regardless, several drivers have been proposed. Non-functional hypotheses posit that masturbation is either a pathology, or a byproduct of high underlying sexual arousal, whereas functional hypotheses argue an adaptive benefit. The Postcopulatory Selection Hypothesis states that masturbation aids the chances of fertilization, while the Pathogen Avoidance Hypothesis states that masturbation helps reduce host infection by flushing pathogens from the genital tract. Here, we present comprehensive new data documenting masturbation across the primate order and use these, in conjunction with phylogenetic comparative methods, to reconstruct the evolutionary pathways and correlates of masturbation. We find that masturbation is an ancient trait within the primate order, becoming a more common aspect of the haplorrhine behavioural repertoire after the split from tarsiers. Our analyses provide support for both the Postcopulatory Selection and Pathogen Avoidance Hypotheses in male primates, suggesting that masturbation may be an adaptive trait, functioning at a macroevolutionary scale.

Discovery of Novel Herpes Simplexviruses in Wild Gorillas, Bonobos, and Chimpanzees Supports Zoonotic Origin of HSV-2.

Viruses closely related to human pathogens can reveal the origins of human infectious diseases. Human herpes simplexvirus type 1 (HSV-1) and type 2 (HSV-2) are hypothesized to have arisen via host-virus codivergence and cross-species transmission. We report the discovery of novel herpes simplexviruses during a large-scale screening of fecal samples from wild gorillas, bonobos, and chimpanzees. Phylogenetic analysis indicates that, contrary to expectation, simplexviruses from these African apes are all more closely related to HSV-2 than to HSV-1. Molecular clock-based hypothesis testing suggests the divergence between HSV-1 and the African great ape simplexviruses likely represents a codivergence event between humans and gorillas. The simplexviruses infecting African great apes subsequently experienced multiple cross-species transmission events over the past 3 My, the most recent of which occurred between humans and bonobos around 1 Ma. These findings revise our understanding of the origins of human herpes simplexviruses and suggest that HSV-2 is one of the earliest zoonotic pathogens.

Nocturnal activity in wild chimpanzees (Pan troglodytes): Evidence for flexible sleeping patterns and insights into human evolution.

OBJECTIVES: We investigated occurrences and patterns of terrestrial nocturnal activity in wild chimpanzees (Pan troglodytes) and modelled the influence of various ecological predictors on nocturnal activity. METHODS: Data were extracted from terrestrial camera-trap footage and ecological surveys from 22 chimpanzee study sites participating in the Pan African Programme: The Cultured Chimpanzee. We described videos demonstrating nocturnal activity, and we tested the effects of the percentage of forest, abundance of predators (lions, leopards and hyenas), abundance of large mammals (buffalos and elephants), average daily temperature, rainfall, human activity, and percent illumination on the probability of nocturnal activity. RESULTS: We found terrestrial nocturnal activity to occur at 18 of the 22 study sites, at an overall average proportion of 1.80% of total chimpanzee activity, and to occur during all hours of the night, but more frequently during twilight hours. We found a higher probability of nocturnal activity with lower levels of human activity, higher average daily temperature, and at sites with a larger percentage of forest. We found no effect of the abundance of predators and large mammals, rainfall, or moon illumination. DISCUSSION: Chimpanzee terrestrial nocturnal activity appears widespread yet infrequent, which suggests a consolidated sleeping pattern. Nocturnal activity may be driven by the stress of high daily temperatures and may be enabled at low levels of human activity. Human activity may exert a relatively greater influence on chimpanzee nocturnal behavior than predator presence. We suggest that chimpanzee nocturnal activity is flexible, enabling them to respond to changing environmental factors.

A Closed-Form Error Model of Straight Lines for Improved Data Association and Sensor Fusing.

Linear regression is a basic tool in mobile robotics, since it enables accurate estimation of straight lines from range-bearing scans or in digital images, which is a prerequisite for reliable data association and sensor fusing in the context of feature-based SLAM. This paper discusses, extends and compares existing algorithms for line fitting applicable also in the case of strong covariances between the coordinates at each single data point, which must not be neglected if range-bearing sensors are used. Besides, in particular, the determination of the covariance matrix is considered, which is required for stochastic modeling. The main contribution is a new error model of straight lines in closed form for calculating quickly and reliably the covariance matrix dependent on just a few comprehensible and easily-obtainable parameters. The model can be applied widely in any case when a line is fitted from a number of distinct points also without a priori knowledge of the specific measurement noise. By means of extensive simulations, the performance and robustness of the new model in comparison to existing approaches is shown.

Raw material procurement for termite fishing tools by wild chimpanzees in the Issa valley, Western Tanzania.

OBJECTIVES: Chimpanzee termite fishing has been studied for decades, yet the selective processes preceding the manufacture of fishing tools remain largely unexplored. We investigate raw material selection and potential evidence of forward planning in the chimpanzees of Issa valley, western Tanzania. MATERIALS AND METHODS: Using traditional archaeological methods, we surveyed the location of plants from where chimpanzees sourced raw material to manufacture termite fishing tools, relative to targeted mounds. We measured raw material abundance to test for availability and selection. Statistics included Chi-Squared, two-tailed Wilcoxon, and Kruskall-Wallace tests. RESULTS: Issa chimpanzees manufactured extraction tools only from bark, despite availability of other suitable materials (e.g., twigs), and selected particular plant species as raw material sources, which they often also exploit for food. Most plants were sourced 1-16 m away from the mound, with a maximum of 33 m. The line of sight from the targeted mound was obscured for a quarter of these plants. DISCUSSION: The exclusive use of bark tools despite availability of other suitable materials indicates a possible cultural preference. The fact that Issa chimpanzees select specific plant species and travel some distance to source them suggests some degree of selectivity and, potentially, forward planning. Our results have implications for the reconstruction of early hominin behaviors, particularly with regard to the use of perishable tools, which remain archaeologically invisible.

Sustained myrmecophagy in Nigerian chimpanzees: Preferred or fallback food?

OBJECTIVE: At some sites across Africa, chimpanzees consume army ants, often aided by plant tools, although consumption frequencies vary greatly. Other populations do not eat these insects at all, despite apparent abundance. The relative importance of this type of myrmecophagy for chimpanzee diet therefore remains unclear. We investigate if army ants constitute a preferred food or a fallback resource for chimpanzees at Gashaka, Nigeria, where army ants are consumed much more frequently than elsewhere. METHODS: Long-term records on temporal variation of climate and availability of fruit as the chimpanzees' preferred food are compared with rates of recovered army ant dipping wands and army ant remains in feces. RESULTS: Despite strict seasonality of rainfall and fruit abundance, myrmecophagy does not negatively correlate with fruit availability. Instead, army ant eating is sustained year round at high levels, with 44% of feces containing remains. CONCLUSIONS: Results contradict the fallback hypothesis and support the hypothesis that ants are a preferred food. Nevertheless, compared with fruit, ant-meals can normally provide only negligible amounts of nutrients. At Gashaka, however, nutritional yield may be significant, given that ant-dipping sessions provide on average 13 mg of dry weight to a chimpanzee. The species exclusively eaten here, Dorylus rubellus, might be particularly aggressive, thus resulting in greater harvesting success than elsewhere. Army ants may hence serve as a diet supplement or complement in terms of macro- or micronutrients.

Not eating like a pig: European wild boar wash their food.

Carrying food to water and either dunking or manipulating it before consumption has been observed in various taxa including birds, racoons and primates. Some animals seem to be simply moistening their food. However, true washing aims to remove unpleasant surface substrates such as grit and sand and requires a distinction between items that do and do not need cleaning as well as deliberate transportation of food to a water source. We provide the first evidence for food washing in suids, based on an incidental observation with follow-up experiments on European wild boar (Sus scrofa) kept at Basel Zoo, Switzerland. Here, all adult pigs and some juveniles of a newly formed group carried apple halves soiled with sand to the edge of a creek running through their enclosure where they put the fruits in the water and pushed them to and fro with their snouts before eating. Clean apple halves were never washed. This indicates that pigs can discriminate between soiled and unsoiled foods and that they are able to delay gratification for long enough to transport and wash the items. However, we were unable to ascertain to which degree individual and/or social learning brought this behaviour about.

Antelope Predation by Nigerian Forest Baboons: Ecological and Behavioural Correlates.

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.

Apes finding ants: Predator-prey dynamics in a chimpanzee habitat in Nigeria.

Some chimpanzee populations prey upon army ants, usually with stick tools. However, how their prey's subterranean nesting and nomadic lifestyle influence the apes' harvesting success is still poorly understood. This is particularly true for chimpanzees (Pan troglodytes ellioti) at Gashaka/Nigeria, which consume army ants (Dorylus rubellus) with much higher frequency than at other sites. We assessed various harvesting and search options theoretically available to the apes. For this, we reconstructed annual consumption patterns from feces and compared the physical characteristics of exploited ant nests with those that were not targeted. Repeated exploitation of a discovered nest is viable only in the short term, as disturbed colonies soon moved to a new site. Moreover, monitoring previously occupied nest cavities is uneconomical, as ants hardly ever re-used them. Thus, the apes have to detect new nests regularly, although colony density is relatively low (1 colony/1.3 ha). Surprisingly, visual search cues seem to be of limited importance because the probability of a nest being exploited was independent of its conspicuousness (presence of excavated soil piles, concealing leaf-litter or vegetation). However, chimpanzees preferentially targeted nests in forests or at the base of food trees, that is, where the apes spend relatively more time and/or where ant colony density is highest. Taken together, our findings suggest that, instead of employing a search strategy based on visual cues or spatial memory, chimpanzee predation on army ants contains a considerable opportunistic element.

Molecular diversity of entodiniomorphid ciliate Troglodytella abrassarti and its coevolution with chimpanzees.

The entodiniomorphid ciliate Troglodytella abrassarti is a colonic mutualist of great apes. Its host specificity makes it a suitable model for studies of primate evolution. We explored molecular diversity of T. abrassarti with regard to large geographical distribution and taxonomic diversity of its most common host, the chimpanzee. We found a very low diversification of T. abrassarti in chimpanzees across Africa. Distribution of two types of T. abrassarti supports evolutionary separation of the Western chimpanzee, P. t. verus, from populations in Central and East Africa. Type I T. abrassarti is probably a derived form, which corresponds with the Central African origin of chimpanzees and a founder event leading to P. t. verus. Exclusivity of the respective types of T. abrassarti to Western and Central/Eastern chimpanzees corroborates the difference found between an introduced population of presumed Western chimpanzees on Rubondo Island and an autochthonous population in mainland Tanzania. The identity of T. abrassarti from Nigerian P. t. ellioti and Central African chimpanzees suggests their close evolutionary relationship. Although this contrasts with published mtDNA data, it corroborates current opinion on the exclusive position of P. t. verus within the chimpanzee phylogeny. The type of T. abrassarti occurring in Central and East African common chimpanzee was confirmed also in bonobos. This may point to the presence of an ancestral Type II found throughout the Lower Guinean rainforest dating back to the common Pan ancestor. Alternatively, the molecular uniformity of T. abrassarti may imply a historical overlap of the species' distribution ranges.

Obtaining raw material: plants as tool sources for Nigerian chimpanzees.

We investigated the acquisition of plant materials from which Nigerian chimpanzees manufacture wooden tools to harvest insects and honey from nests of army ants, honey bees and stingless bees. Slender trunks of juvenile trees and branches are most commonly used, and bendable vines rarely, probably reflecting the need to work with relatively sturdy tools to extract resources. While several tools are sometimes sourced from the same plant, there is also evidence for a depletion effect, as multiple tool sources at the same site are often spaced several metres apart. Identified tool sources belong to 27 species of at least 13 families. Honey-gathering implements are often chewed upon by chimpanzees. Interestingly, twigs of the most commonly used honey-gathering species possess antibacterial propensities and are favoured by Nigerians as chewing sticks. This suggests that extractive tools might possess associated medicinal or stimulatory properties. We do not know if chimpanzees actively select specific plant parts or species as we cannot compare observed with expected frequencies. Nevertheless, about three quarters of tools are picked from plants more than 6 m away from the extraction site, potentially indicating some degree of forward planning.

Performance of human groups in social foraging: the role of communication in consensus decision making.

Early hominids searched for dispersed food sources in a patchy, uncertain environment, and modern humans encounter equivalent spatial-temporal coordination problems on a daily basis. A fundamental, but untested assumption is that our evolved capacity for communication is integral to our success in such tasks, allowing information exchange and consensus decisions based on mutual consideration of pooled information. Here we examine whether communication enhances group performance in humans, and test the prediction that consensus decision-making underlies group success. We used bespoke radio-tagging methodology to monitor the incremental performance of communicating and non-communicating human groups (small group sizes of two to seven individuals), during a social foraging experiment. We found that communicating groups (n = 22) foraged more effectively than non-communicating groups (n = 21) and were able to reach consensus decisions (an 'agreement' on the most profitable foraging resource) significantly more often than non-communicating groups. Our data additionally suggest that gesticulations among group members played a vital role in the achievement of consensus decisions, and therefore highlight the importance of non-verbal signalling of intentions and desires for successful human cooperative behaviour.

Hybrid apes in the Anthropocene: Burden or asset for conservation?

Conservationists often view hybrid animals as problematic, at least if anthropogenic influence caused the intermixing to occur. However, critics propose that humans should respect non-human autonomy, reject and accept the creatures they have helped to create.Based on two case studies of our own ethological, genetic and ethnographic research about chimpanzee and orangutan subspecies hybrids, we assess what, if anything, should be done about such animals. We consider problems posed by cross-bred apes relating to: (a) Breeding-Do hybrids really experience reduced reproductive success? How are population-level concerns and welfare of individual animals balanced in conservation breeding? (b) Essentialism-Are anti-hybrid arguments based on essentialist or purist thinking? Does essentialism vary by conservation context? (c) Pragmatism-How do socio-economic circumstances influence whether hybrids are embraced or ignored? Does the erosion of 'untouched nature' render hybrids more important?We show that answers to these questions are complex and context-specific, and that therefore decisions should be made on a case-by-case basis. For example, we find that anti-hybrid arguments are essentialist in some cases (e.g. ape management in zoos) but not in others (e.g. ape reintroduction). Thus, rather than present recommendations, we conclude by posing nine questions that conservationists should ask themselves when making decisions about taxonomic hybrids. ​ A free Plain Language Summary can be found within the Supporting Information of this article.

Population history of chimpanzees introduced to Lake Victoria's Rubondo Island.

Between 1966 to 1969, Bernhard Grzimek (Frankfurt Zoological Society, FZS) introduced chimpanzees (Pan troglodytes) previously held in European institutions to Rubondo Island in Lake Victoria in Tanzania. Earlier publications report various numbers of released animals and that all founders originated from West Africa. We revise these assumptions through consultation of archived FZS records and genetic analyses of surviving descendants. Accordingly, 17 chimpanzees were transported to Africa in four waves, with male-female ratios of 3:8, 1:0, 1:0 and 2:2; one female died in transit. Thus, 16 chimpanzees were released in total. FZS and studbook records allocate a West African provenance to only 19% of the founders and a generic "Africa" origin to 56%. Still, studbook records render it unlikely that any of the founders were captive-born. In addition, our genetic analyses based on biological samples from the current descendants trace the geographical origin of their ancestors back to West Africa (P. t. verus) and Central Africa (P. t. troglodytes). Based on counts of individuals and night nests, we estimate that the population, since 1969, grew to around 35 individuals in 2014 (annual increase 3.3%). Thus, chimpanzees released onto a large forested island free from predators or hunting pressure, habitat destruction and conspecific competition can form a self-sustaining island population without human support.

Developmental instability in wild Nigerian olive baboons (Papio anubis).

BACKGROUND: Developmental instability in archaeological samples can be detected through analysis of skeletal and dental remains. During life, disruptions to biological internal homeostasis that occur during growth and development redirect bodily resources to returning to homeostasis and away from normal processes such as symmetrical development. Because dental enamel does not remodel in life, any deviations from normal development are left behind. Even subtle disturbances to developmental trajectory may be detected in asymmetrical development of traits, specifically a random variation in sides termed fluctuating asymmetry. Human dental fluctuating asymmetry studies are common, but here we investigate the permanent dentition of a non-human primate Papio anubis, for potential fluctuating asymmetry relative to sex, weaning, and reproductive maturity. The sample stems from an outlier population that lives in the wettest and most humid habitat of any studied baboon group. METHODS: The skulls of adult baboons were collected after their natural death in Gashaka Gumti National Park, Nigeria. The permanent dentition of antimeric teeth (paired) were measured for maximum length and breadth using standard methods. The metrics were analyzed to assess the presence of fluctuating asymmetry in adult permanent mandibular and maxillary dentition. Measurement error and other forms of asymmetry (antisymmetry, directional asymmetry) were considered and dental measures expressing true fluctuating asymmetry were used to address three research questions. RESULTS: Males exhibit greater fluctuating asymmetry than females, suggesting that males experience greater overall instability during the developmental period. While weaning is not more stressful than other life history stages for males and females (using the first molar fluctuating asymmetry index as a proxy compared to other teeth), it is more stressful for females than males. The onset of reproduction is also not more stressful than other life history stages for males and females (using the third molar fluctuating asymmetry index as a proxy compared to other teeth), but it is more stressful for males than females. We explore possible explanations for these findings in the discussion.

Limited evidence of C4 plant consumption in mound building Macrotermes termites from savanna woodland chimpanzee sites.

Stable isotope analysis is an increasingly used molecular tool to reconstruct the diet and ecology of elusive primates such as unhabituated chimpanzees. The consumption of C4 plant feeding termites by chimpanzees may partly explain the relatively high carbon isotope values reported for some chimpanzee communities. However, the modest availability of termite isotope data as well as the diversity and cryptic ecology of termites potentially consumed by chimpanzees obscures our ability to assess the plausibility of these termites as a C4 resource. Here we report the carbon and nitrogen isotope values from 79 Macrotermes termite samples from six savanna woodland chimpanzee research sites across equatorial Africa. Using mixing models, we estimated the proportion of Macrotermes C4 plant consumption across savanna woodland sites. Additionally, we tested for isotopic differences between termite colonies in different vegetation types and between the social castes within the same colony in a subset of 47 samples from 12 mounds. We found that Macrotermes carbon isotope values were indistinguishable from those of C3 plants. Only 5 to 15% of Macrotermes diets were comprised of C4 plants across sites, suggesting that they cannot be considered a C4 food resource substantially influencing the isotope signatures of consumers. In the Macrotermes subsample, vegetation type and caste were significantly correlated with termite carbon values, but not with nitrogen isotope values. Large Macrotermes soldiers, preferentially consumed by chimpanzees, had comparably low carbon isotope values relative to other termite castes. We conclude that Macrotermes consumption is unlikely to result in high carbon isotope values in either extant chimpanzees or fossil hominins.

Structure of Chimpanzee Gut Microbiomes across Tropical Africa.

Understanding variation in host-associated microbial communities is important given the relevance of microbiomes to host physiology and health. Using 560 fecal samples collected from wild chimpanzees (Pan troglodytes) across their range, we assessed how geography, genetics, climate, vegetation, and diet relate to gut microbial community structure (prokaryotes, eukaryotic parasites) at multiple spatial scales. We observed a high degree of regional specificity in the microbiome composition, which was associated with host genetics, available plant foods, and potentially with cultural differences in tool use, which affect diet. Genetic differences drove community composition at large scales, while vegetation and potentially tool use drove within-region differences, likely due to their influence on diet. Unlike industrialized human populations in the United States, where regional differences in the gut microbiome are undetectable, chimpanzee gut microbiomes are far more variable across space, suggesting that technological developments have decoupled humans from their local environments, obscuring regional differences that could have been important during human evolution. IMPORTANCE Gut microbial communities are drivers of primate physiology and health, but the factors that influence the gut microbiome in wild primate populations remain largely undetermined. We report data from a continent-wide survey of wild chimpanzee gut microbiota and highlight the effects of genetics, vegetation, and potentially even tool use at different spatial scales on the chimpanzee gut microbiome, including bacteria, archaea, and eukaryotic parasites. Microbial community dissimilarity was strongly correlated with chimpanzee population genetic dissimilarity, and vegetation composition and consumption of algae, honey, nuts, and termites were potentially associated with additional divergence in microbial communities between sampling sites. Our results suggest that host genetics, geography, and climate play a far stronger role in structuring the gut microbiome in chimpanzees than in humans.

Visual depictions of female genitalia differ depending on source.

Very little research has attempted to describe normal human variation in female genitalia, and no studies have compared the visual images that women might use in constructing their ideas of average and acceptable genital morphology to see if there are any systematic differences. The objective of the present work was to determine if visual depictions of the vulva differed according to their source so as to alert medical professionals and their patients to how these depictions might capture variation and thus influence perceptions of 'normality'. A comparative analysis was conducted by measuring (a) published visual materials from human anatomy textbooks in a university library, (b) feminist publications (print and online) depicting vulval morphology and (c) online pornography, focusing on the most visited and freely accessible sites in the UK. Post hoc tests showed that labial protuberance was significantly less (p<0.001, equivalent to approximately 7 mm) in images from online pornography compared to feminist publications. All five measures taken of vulval features were significantly correlated (p<0.001) in the online pornography sample, indicating a less varied range of differences in organ proportions than the other sources where not all measures were correlated. Women and health professionals should be aware that specific sources of imagery may depict different types of genital morphology and may not accurately reflect true variation in the population, and consultations for genital surgeries should include discussion about the actual and perceived range of variation in female genital morphology.

Living on the edge: life-history of olive baboons at Gashaka-Gumti National Park, Nigeria.

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.