Elevated glucocorticoids during the ovarian follicular phase predict conception in wild female chacma baboons.

Mating related behavior during ovarian cycling can be energetically demanding and constitute a significant stressor, requiring physiological responses to mediate investment in reproduction. To better understand the proximate mechanisms underlying these responses, we examine hormonal and behavioral variation across the ovarian cycle during conceptive and nonconceptive cycles in wild female chacma baboons (Papio ursinus). We quantified immunoreactive fecal estradiol, progesterone, and cortisol metabolites for 21 adult females, and calculated activity budgets and rates of received aggression from over 5000 15-min behavioral samples. We found conception to be associated with higher concentrations of both estradiol and cortisol during the follicular phase, but no difference in progesterone between conceptive and nonconceptive cycles for either the follicular or luteal phase. While females spent less time feeding during the follicular compared to the luteal phase, we found no difference in time spent feeding, moving, or copulating between conceptive and nonconceptive cycles of the same phase. Rates of received aggression also were similar across the ovarian cycle, with no difference between conceptive and nonconceptive cycles. Finally, we found positive associations between cortisol and estradiol, indicating that glucocorticoids (GCs) do not suppress hypothalamic-pituitary-gonadal (HPG) activity and reproductive function in this context. Overall, our results suggest that elevated GCs may play an adaptive role in mobilizing energy during sexually receptive periods of ovarian cycling.

Enamel chipping and its ecological correlates in African papionins: Implications for hominin feeding behavior.

African papionins are classic paleoecological referents for fossil hominins. Enamel chips on the teeth of baboons and hominins are argued to represent responses to similar dietary habits; however, a comprehensive analysis of modern papionin chipping is lacking, leaving open the question of analog suitability. Here, we investigate patterns of antemortem enamel chipping across a diverse set of African papionin species occupying a range of ecological niches. We compare papionin chipping frequencies to estimates for Plio-Pleistocene hominins to address hypotheses of habitat and/or dietary similarities. Antemortem chips in seven African papionin species were scored on intact postcanine teeth (P3-M3) using established protocols. Chip size was scored on a tripartite scale. Papio hamadryas and Papio ursinus-two common paleoecological referents-display higher levels of chipping than Plio-Pleistocene hominin taxa (Australopithecus and Paranthropus) posited to have similar dietary habits. Papio populations occupying dry or highly seasonal habitats accumulate more large chips than Papio taxa occupying more mesic habitats, and terrestrial papionins chip their teeth more often than closely related taxa occupying arboreal niches. Chipping is present on the teeth of all Plio-Pleistocene hominins; however, chipping in baboons (P. ursinus and P. hamadryas) consistently exceeds most hominin taxa. Chipping frequencies on their own do not reliably sort taxa into major dietary groupings. We conclude that the large differences in chipping frequency may instead reflect habitat use and food processing idiosyncrasies. Less chipping in Plio-Pleistocene hominin teeth compared to modern Papio is more likely attributable to differences in dental morphology rather than diet.

Respect for males amid suppression of females: Selective use of aggression and fitness correlates in the male-dominated society of hamadryas baboons.

Hamadryas baboons (Papio hamadryas) are a useful model for human social evolution for multiple reasons, including their multilevel society, intense cross-sex bonds, and intermale tolerance. Their most stable social grouping, the one-male unit (OMU)-comprising a leader male, females, and sometimes follower males-is formed via successive takeovers of individual females by males. While takeovers occur via both aggressive and non-aggressive mechanisms, aggressive herding is common during and after takeovers and appears crucial in maintaining OMUs. Here we use behavioral and demographic data from Filoha, Ethiopia to examine the relationship between aggressive takeovers and fitness correlates. We found no relationship between a male's percentage of takeovers that were aggressive and his presumed number of infants sired, nor his number of females or followers. However, we did find that a leader male's average intensity of aggression toward both other males and females around the time of a takeover was negatively related to his presumed number of infants sired. In addition, a leader male's average intensity of aggression toward other males was negatively related to his maximum number of followers. Finally, leader males exhibited more intense aggression toward females in interband, compared to intraband, takeovers. Our findings suggest that (1) leader males who limit their aggression toward other males may have greater success in attracting followers, thereby increasing their fitness via enhanced defense of the OMU; (2) exceptionally aggressive takeovers may lead to lower birth rates via female reproductive suppression; and (3) the extent to which males use aggression toward females depends on the context in which the takeover occurs. Overall, these results both suggest that hamadryas males use aggression selectively and underscore the ubiquity of intermale tolerance and female suppression in the hamadryas social system. This study lends insight into the interplay between male-female and male-male social dynamics during human evolution.

Costs of seasonality at a southern latitude: Behavioral endocrinology of female baboons in the Cape Peninsula of South Africa.

Environmental challenges in the form of temperature extremes and unusual precipitation, which may lead to prolonged periods outside the thermoneutral zone, can be detrimental to animal physiology. Chacma baboons in the Cape Peninsula of South Africa, one of the highest latitudes at which nonhuman primates are found, experience extremes of both temperature and rainfall, as well as seasonal differences in day length that require animals to condense their daily routine into dramatically reduced daylight hours. Here we examine the effects of these climatic factors on the behavior (activity budgets and foraging patterns) and physiology (fecal glucocorticoid concentrations) of adult females (N = 33) in three groups of chacma baboons (Papio ursinus) inhabiting the Cape Peninsula, where temperatures ranged from 7 to 39 °C, monthly rainfall ranged from 2 to 158 mm, and day length varied by 4.5 h across seasons. Climatic variables showed a clear relationship to female baboon glucocorticoid concentrations, which significantly increased with lower temperatures, higher rainfall and shorter day lengths. Activity budgets also differed between summer and winter, with females generally spending less time socializing, moving and resting in the winter compared to summer, with some differences between troops in their feeding-related activities. Cold temperatures accompanied by rainfall and short day lengths may thus represent an ecological constraint for this population. This study highlights the potential impact of anthropogenic climate change on the physiology, behavior, and, ultimately, survival of wildlife populations.

Predictors of helminth parasite infection in female chacma baboons (Papio ursinus).

Helminth parasite infection can impose major consequences on host fitness. Several factors, including individual characteristics of hosts, environmental conditions, and patterns of coinfection, are thought to drive variation in parasite risk. Here, we report on four key drivers of parasite infection-phase of reproduction, steroid hormone profiles, rainfall, and patterns of coinfection-in a population of wild female chacma baboons (Papio ursinus) in South Africa. We collected data on reproductive state and hormone profiles over a 3-year span, and quantified helminth parasite burdens in 2955 fecal samples from 24 female baboons. On a host level, we found that baboons are sensitive to parasite infection during the costliest phases of the reproductive cycle: pregnant females harbored higher intensities of Protospirura eggs than cycling and lactating females; lactating and cycling females had a higher probability of Oesophagostomum infection than pregnant females; and cycling females exhibited lower Trichuris egg counts than pregnant and lactating females. Steroid hormones were associated with both immunoenhancing and immunosuppressive properties: females with high glucocorticoid concentrations exhibited high intensities of Trichuris eggs but were at low risk of Oesophagostomum infection; females with high estrogen and progestagen concentrations exhibited high helminth parasite richness; and females with high progestagen concentrations were at high risk of Oesophagostomum infection but exhibited low Protospirura egg counts. We observed an interaction between host reproductive state and progestagen concentrations in infection intensity of Protospirura: pregnant females exhibited higher intensities and non-pregnant females exhibited lower intensities of Protospirura eggs with increasing progestagen concentrations. At a population level, rainfall patterns were dominant drivers of parasite risk. Lastly, helminth parasites exhibited positive covariance, suggesting that infection probability increases if a host already harbors one or more parasite taxa. Together, our results provide a holistic perspective of factors that shape variation in parasite risk in a wild population of animals.

Home range, sleeping site use, and band fissioning in hamadryas baboons: Improved estimates using GPS collars.

Variation in spatial and temporal distribution of resources drives animal movement patterns. Links between ecology and behavior are particularly salient for the multilevel society of hamadryas baboons, in which social units cleave and coalesce over time in response to ecological factors. Here, we used data from GPS collars to estimate home range size and assess temporal patterns of sleeping site use in a band of hamadryas baboons in Awash National Park, Ethiopia. We used GPS data derived from 2 to 3 collared baboons over three 8-12-month collaring intervals to estimate annual and monthly home ranges using kernel density estimators (KDEs) and minimum convex polygons (MCPs). The 95% KDE home range was 64.11 km2 for Collaring Interval I (July 2015-March 2016), 85.52 km2 for Collaring Interval II (October 2016-October 2017), 76.43 km2 for Collaring Interval III (July 2018-May 2019), and 75.25 km2 across all three collaring intervals. MCP home ranges were 103.46 km2 for Collaring Interval I, 97.90 km2 for Collaring Interval II, 105.22 km2 for Collaring Interval III, and 129.33 km2 overall. Ninety-five percent KDE home range sizes did not differ across months, nor correlate with temperature or precipitation, but monthly MCP home ranges increased with monthly precipitation. Our data also revealed a southward home range shift over time and seven previously unknown sleeping sites, three of which were used more often during the wet season. Band cohesion was highest during dry months and lowest during wet months, with fissioning occurring more frequently at higher temperatures. One pair of collared individuals from Collaring Interval III spent 95% of nights together, suggesting they were members of the same clan. Our results both suggest that previous studies have underestimated the home range size of hamadryas baboons and highlight the benefits of remote data collection.

Multilevel Organisation of Animal Sociality.

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

Anthropogenic effects on the physiology and behaviour of chacma baboons in the Cape Peninsula of South Africa.

As animals increasingly occupy habitats in proximity to humans, with home ranges a mosaic of natural and anthropogenic landscapes, it becomes imperative from a conservation perspective to understand the impacts of human activities on wildlife. Many non-human primates share habitats with humans, an ability stemming largely from shared ecological needs due to our close evolutionary relationship. Such proximity, however, is often accompanied by direct conflict between humans and wildlife, leading to higher stress levels, injuries, mortality and behavioural changes, with detrimental effects on long-term health and fitness. Glucocorticoid (GC) hormones, which are released in response to ecological and social challenges, are increasingly employed to understand responses to anthropogenic disturbance. Here we investigate physiological and behavioural responses of female chacma baboons (Papio ursinus) to variation in spatial overlap and conflict with humans in their natural home range. The baboons resided in the Tokai Section of Table Mountain National Park in the Cape Peninsula of South Africa, where their home range included four types of habitats: protected, suburban, agricultural and mosaic areas. We studied the largest group of baboons in Tokai (70 individuals) and examined the effects of ranging in these different habitats on the faecal GCs and behaviour of all adult females in the troop (N = 16). We found time spent ranging in more anthropogenic habitats to be associated with higher levels of GCs, more aggression, less time socializing and shorter grooming bouts. Self-directed behaviour, however, varied and did not necessarily reflect physiological measures of stress. Taken together, the results of this study highlight the risks associated with ranging in anthropogenic environments and point to the need for a multifaceted approach to studying the negative impacts of human activities on animals so as to better inform conservation practices.

Biodiversity of protists and nematodes in the wild nonhuman primate gut.

Documenting the natural diversity of eukaryotic organisms in the nonhuman primate (NHP) gut is important for understanding the evolution of the mammalian gut microbiome, its role in digestion, health and disease, and the consequences of anthropogenic change on primate biology and conservation. Despite the ecological significance of gut-associated eukaryotes, little is known about the factors that influence their assembly and diversity in mammals. In this study, we used an 18S rRNA gene fragment metabarcoding approach to assess the eukaryotic assemblage of 62 individuals representing 16 NHP species. We find that cercopithecoids, and especially the cercopithecines, have substantially higher alpha diversity than other NHP groups. Gut-associated protists and nematodes are widespread among NHPs, consistent with their ancient association with NHP hosts. However, we do not find a consistent signal of phylosymbiosis or host-species specificity. Rather, gut eukaryotes are only weakly structured by primate phylogeny with minimal signal from diet, in contrast to previous reports of NHP gut bacteria. The results of this study indicate that gut-associated eukaryotes offer different information than gut-associated bacteria and add to our understanding of the structure of the gut microbiome.

Insights into the evolution of social systems and species from baboon studies.

Baboons, members of the genus Papio, comprise six closely related species distributed throughout sub-Saharan Africa and southwest Arabia. The species exhibit more ecological flexibility and a wider range of social systems than many other primates. This article summarizes our current knowledge of the natural history of baboons and highlights directions for future research. We suggest that baboons can serve as a valuable model for complex evolutionary processes, such as speciation and hybridization. The evolution of baboons has been heavily shaped by climatic changes and population expansion and fragmentation in the African savanna environment, similar to the processes that acted during human evolution. With accumulating long-term data, and new data from previously understudied species, baboons are ideally suited for investigating the links between sociality, health, longevity and reproductive success. To achieve these aims, we propose a closer integration of studies at the proximate level, including functional genomics, with behavioral and ecological studies.

Convergence of human and Old World monkey gut microbiomes demonstrates the importance of human ecology over phylogeny.

BACKGROUND: Comparative data from non-human primates provide insight into the processes that shaped the evolution of the human gut microbiome and highlight microbiome traits that differentiate humans from other primates. Here, in an effort to improve our understanding of the human microbiome, we compare gut microbiome composition and functional potential in 14 populations of humans from ten nations and 18 species of wild, non-human primates. RESULTS: Contrary to expectations from host phylogenetics, we find that human gut microbiome composition and functional potential are more similar to those of cercopithecines, a subfamily of Old World monkey, particularly baboons, than to those of African apes. Additionally, our data reveal more inter-individual variation in gut microbiome functional potential within the human species than across other primate species, suggesting that the human gut microbiome may exhibit more plasticity in response to environmental variation compared to that of other primates. CONCLUSIONS: Given similarities of ancestral human habitats and dietary strategies to those of baboons, these findings suggest that convergent ecologies shaped the gut microbiomes of both humans and cercopithecines, perhaps through environmental exposure to microbes, diet, and/or associated physiological adaptations. Increased inter-individual variation in the human microbiome may be associated with human dietary diversity or the ability of humans to inhabit novel environments. Overall, these findings show that diet, ecology, and physiological adaptations are more important than host-microbe co-diversification in shaping the human microbiome, providing a key foundation for comparative analyses of the role of the microbiome in human biology and health.

Social evolution in Plio-Pleistocene hominins: Insights from hamadryas baboons and paleoecology.

Reconstructions of hominin evolution have long benefited from comparisons with nonhuman primates, especially baboons and chimpanzees. The hamadryas baboon (Papio hamadryas) is arguably one of the best such models, as it exhibits both the male kin bonding and the cross-sex pair bonding thought to have been important in hominin evolution. Here we link processes of behavioral evolution in hamadryas baboons with those in a Plio-Pleistocene hominin, provisionally identified as Homo erectus (sensu lato) - a pivotal species in that its larger body and brain size and wider ranging patterns increased female costs of reproduction, increasing the importance of sociality. The combination of these higher costs of reproduction and shifts in diet and food acquisition have previously been argued to have been alleviated either via strengthening of male-female bonds (involving male provisioning and the evolution of monogamy) or via the assistance of older, post-reproductive females (leading to post-reproductive longevity in females, i.e., the grandmother hypothesis). We suggest that both arrangements could have been present in Plio-Pleistocene hominins if they lived in multilevel societies. Here we expand on our earlier scenario with two sets of recent data in support of it, (1) archaeological data from the 2 million year old Oldowan site of Kanjera South, Kenya and other sites that are suggestive of tool dependent foraging on nutrient dense resources (animal carcasses and plant underground storage organs), cooperation, and food sharing; and (2) a pattern of genetic variation in hamadryas baboons that suggests the operation of kin selection among both males and females at multiple levels of society. Taken together, these two sets of data strengthen our model and support the idea of a complex society linked by male-male, male-female, and female-female bonds at multiple levels of social organization in Plio-Pleistocene hominins.

Meat eating in wild hamadryas baboons: Opportunistic trade-offs between insects and vertebrates.

The consumption of meat may provide herbivorous animals with important nutrients that are scarce in their plant-based diet. Seasonal variation in plant food availability has been suggested to motivate dietary flexibility in a range of species and thus primates may seek more prey when key plant resources are unavailable. Alternatively, prey encounter rate may drive meat eating. Here we investigate patterns of meat eating in hamadryas baboons (Papio hamadryas) at Filoha, Awash National Park, Ethiopia. The Filoha baboons rely largely on doum palm fruit (Hyphaene thebaica), which are available most months of the year, and the young leaves of Acacia senegal, which are more abundant during the wet season. We hypothesized that the baboons would consume more meat when H. thebaica and A. senegal were less available, which we tested by comparing meat eating and consumption of these plant food species from March 2005 through February 2006. Our results reveal a high rate of vertebrate meat eating at Filoha (0.028/hour of observation) compared with other hamadryas sites. We found no relationship, however, between meat eating (either insect or vertebrate) and either rainfall or consumption of H. thebaica or A. senegal, indicating that availability of preferred plant resources does not drive meat consumption. Vertebrate consumption and time spent feeding were significantly negatively associated; there was no relationship, however, between the consumption of animal matter and either home range size or daily path length. Vertebrate and insect consumption alternated throughout the year such that the baboons maintained a small amount of animal matter in their diet year-round. Our results suggest that the baboons do not often actively seek animal matter, but consume it opportunistically, with the presence of locust and dragonfly swarms driving insect consumption, and both prey availability and the availability of feeding time shaping vertebrate predation.

Contexts and consequences of takeovers in hamadryas baboons: Female parity, reproductive state, and observational evidence of pregnancy loss.

The evolutionary consequences of sexual selection and sexual conflict are epitomized in the hamadryas baboon, a species characterized by strong sexual dimorphism and intense male-male competition. Hamadryas males coerce individual females into reproductively exclusive one-male units via aggressive takeovers, and infants involved in such takeovers are at least four times more likely to be killed (or otherwise die) after takeovers compared to other times. Here we examine female reproductive state before and after takeovers to further investigate the determinants of takeovers and their impact on female reproduction. Our data set consists of 162 takeovers over a period of 20 years in a population of wild hamadryas baboons at Filoha, Ethiopia. Results show that, in contrast to their polygynandrous relatives, hamadryas males show no bias with regard to female reproductive state at the time of the takeover, but are more likely to take over nulliparous compared to multiparous females. In addition, these results reinforce previous findings demonstrating the high likelihood of infant loss after hamadryas takeovers and suggest that females may use an additional strategy of pregnancy termination to curtail investment in offspring that are very likely to die anyway. Thus, hamadryas males may target females with high future reproductive value and hamadryas females may employ counterstrategies to help mitigate the effects of sexual coercion and infanticide on their own fitness.

The ties that bind: Maternal kin bias in a multilevel primate society despite natal dispersal by both sexes.

In many social animals, individuals derive fitness benefits from close social bonds, which are often formed among kin of the philopatric sex. Hamadryas baboons, however, exhibit a hierarchical, multilevel social system where both sexes disperse from their natal one-male-unit (OMU). Although this would seem to hinder maintenance of kin ties, both sexes appear largely philopatric at the higher order band and clan levels, possibly allowing for bonds with same sex kin by both males and females. In order to investigate the possibility of kin bonds in hamadryas baboons, we identified kin dyads in a band without known pedigree information using a large panel of genetic markers: 1 Y-linked, 4 X-linked, and 23 autosomal microsatellites and part of the mitochondrial hypervariable region I. With these data, we performed a kinship analysis while accounting for misclassification rates through simulations and determined kinship among two types of dyads: leader and follower males and female dyads within OMUs. Leader and follower males were maternal relatives more often than expected by chance, suggesting that kinship plays a role in the formation of these relationships. Moreover, maternal female relatives were found in the same OMU more often than expected by chance, indicating that females may be motivated to maintain post-dispersal contact with maternal female kin. Our results suggest that hamadryas baboons can recognize maternal kin and that kin selection has contributed to shaping their complex social system. This implies that an ancestral maternal kin bias has been retained in hamadryas society. Am. J. Primatol. 78:731-744, 2016. © 2016 Wiley Periodicals, Inc.

The adaptive value of secondary males in the polygynous multi-level society of hamadryas baboons.

OBJECTIVES: One-male social systems are usually characterized by polygyny and reproductive exclusion by a single resident male. Sometimes, however, secondary males join these groups, and this may carry fitness costs and/or benefits to the resident male. In hamadryas baboons (Papio hamadryas hamadryas), which live in one-male units (OMUs) with female defense polygyny within a multi-level social system, secondary "follower" males often reside in OMUs. Our aim here is to examine possible benefits of these secondary males to hamadryas resident males. MATERIALS AND METHODS: Using 6 years of data from 65 OMUs in a band of wild hamadryas baboons in Ethiopia, we compared demographic and reproductive parameters of OMUs with and without secondary "follower" males to assess whether their presence conferred any reproductive benefits to resident "leader" males. RESULTS: Leaders with followers had tenure lengths almost twice as long, acquired more than twice as many females, retained females longer, and had three times as many infants during their tenure compared to leaders without followers. DISCUSSION: Hamadryas follower males enabled leaders to retain females for longer periods of time-likely through unit defense, social relationships with OMU members, and/or infant protection. Hamadryas leaders appear to be able to monopolize access to females despite the presence of followers, and as such any enhanced reproduction derived from the presence of followers likely increases the fitness of the leader rather than the follower. Thus the relationship between leaders and followers in hamadryas society appears to be a mutually beneficial one and tolerance of secondary males may be an adaptive reproductive strategy characterizing hamadryas leader males.

Fine-scale genetic assessment of sex-specific dispersal patterns in a multilevel primate society.

Like humans, hamadryas baboons (Papio hamadryas) are unusual among primates in having a multilevel social system and stable pair bonds, and are thus a useful model for the evolution of human sociality. While the kinship structure and sex-biased dispersal patterns that underlie human social organization have been extensively elucidated, the impact of these factors on the social system of hamadryas baboons is currently unclear. Here we use genetic analysis of individuals to elucidate the patterns of male and female dispersal across multiple levels of society in a wild population of hamadryas baboons. We characterized 244 members of five hamadryas bands at Filoha, Ethiopia by genotyping one Y-linked and 23 autosomal microsatellite loci and sequencing part of the mitochondrial hypervariable control region I. We found both male and female dispersal to be limited at the level of the band, with more movement of females than males among bands. By integrating long-term behavioral data for Band 1, we also found evidence for male and female philopatry at the clan level. We speculate that male philopatry at the clan level and female dispersal across one-male units and clans may enable both kin-based cooperation among males and the maintenance of kin bonds among females after dispersal. This would mean that, as in humans, kin bonds within both sexes are a core feature of the hamadryas social system, thus contributing to our understanding of the evolution of social organization in humans.

The rise and fall of leader males in a multi-level society: takeovers and tenures of male Hamadryas baboons.

As one means to maximize access to females, males of some species are intolerant of other males in social units, resulting in female defense polygyny, a mating system in which one male monopolizes mating access to as many females as he can for as long as possible. In such a system, the length of a male's tenure and the number of females he is able to acquire are important predictors of his reproductive success. Hamadryas baboons differ from many other taxa with female defense polygyny in that they acquire and lose females individually, thus patterns of acquisition and loss of females over time are additional factors contributing to a male's fitness. Here, we describe longitudinal patterns of female acquisition and loss over a 9-year period in a group of 250 wild hamadryas baboons. Complete tenures of leader males ranged from 310 to 2,160 days (N = 13) and results from a survival analysis yielded a median tenure length of 2,160 days, or 6 years (N = 49). The total number of females acquired, which increased with tenure length, ranged from 1 to 14 and averaged 3.5, and leader males acquired females both opportunistically and via challenging other males. The interval between acquisition of successive females ranged from 0 to 1,196 days with a median of 203, and males acquired all of their females less than halfway into their tenure. Females from outside of a leader male's social sphere (clan and band) were acquired relatively later in their tenure compared to females from within a male's social sphere. Leaders typically lost females gradually during the latter part of their tenure or all (or most) at once, suggesting an inverted U-shaped longitudinal arc of male competitive ability.

30 days in the life: daily nutrient balancing in a wild chacma baboon.

For most animals, the ability to regulate intake of specific nutrients is vital to fitness. Recent studies have demonstrated nutrient regulation in nonhuman primates over periods of one observation day, though studies of humans indicate that such regulation extends to longer time frames. Little is known about longer-term regulation in nonhuman primates, however, due to the challenges of multiple-day focal follows. Here we present the first detailed study of nutrient intake across multiple days in a wild nonhuman primate. We conducted 30 consecutive all day follows on one female chacma baboon (Papio hamadryas ursinus) in the Cape Peninsula of South Africa. We documented dietary composition, compared the nutritional contribution of natural and human-derived foods to the diet, and quantified nutrient intake using the geometric framework of nutrition. Our focus on a single subject over consecutive days allowed us to examine daily dietary regulation within an individual over time. While the amounts varied daily, our subject maintained a strikingly consistent balance of protein to non-protein (fat and carbohydrate) energy across the month. Human-derived foods, while contributing a minority of the diet, were higher in fat and lower in fiber than naturally-derived foods. Our results demonstrate nutrient regulation on a daily basis in our subject, and demonstrate that she was able to maintain a diet with a constant proportional protein content despite wide variation in the composition of component foods. From a methodological perspective, the results of this study suggest that nutrient intake is best estimated over at least an entire day, with longer-term regulatory patterns (e.g., during development and reproduction) possibly requiring even longer sampling. From a management and conservation perspective, it is notable that nearly half the subject's daily energy intake derived from exotic foods, including those currently being eradicated from the study area for replacement by indigenous vegetation.