Less bins, less baboons: reducing access to anthropogenic food effectively decreases the urban foraging behavior of a troop of chacma baboons (Papio hamadryas ursinus) in a peri-urban area.

In South Africa, chacma baboons (Papio hamadryas ursinus) living near peri-urban areas may forage on anthropogenic food. Baboons have been recorded to damage crops, scatter waste from trash bins, and damage homes. A number of methods have been tested over the past 20 years to solve these problems, but none proved successful over the long-term or involved considerable costs. An efficient management system requires a detailed knowledge of how baboon troops proceed and organize during these urban foraging actions. This study examines the response of a troop of baboons to an experimental reduction of anthropogenic food sources in a peri-urban environment, the George campus of Nelson Mandela University (SA). We gradually suppressed access to waste food in trash cans, reducing the amount of anthropogenic food available. This change in food availability led baboons to modify their urban foraging strategy. They compensated for the lack of anthropogenic food by spending more time foraging on natural food and less time in urban areas. However, the troop still exploited waste-free areas during the experiment and even more when the conditions were normal again. Overall, these results show the ability of baboons to adapt to changes in anthropogenic food availability but also that they are highly dependent on this type of resource. Limiting its access is a mitigation strategy that humans must absolutely develop for reaching a high level of coexistence with baboons.

Evolutionary Determinants of Nonseasonal Breeding in Wild Chacma Baboons.

AbstractAnimal reproductive phenology varies from strongly seasonal to nonseasonal, sometimes among closely related or sympatric species. While the extent of reproductive seasonality is often attributed to environmental seasonality, this fails to explain many cases of nonseasonal breeding in seasonal environments. We investigated the evolutionary determinants of nonseasonal breeding in a wild primate, the chacma baboon (Papio ursinus), living in a seasonal environment with high climatic unpredictability. We tested three hypotheses proposing that nonseasonal breeding has evolved in response to (1) climatic unpredictability, (2) reproductive competition between females favoring birth asynchrony, and (3) individual, rank-dependent variations in optimal reproductive timing. We found strong support for an effect of reproductive asynchrony modulated by rank: (i) birth synchrony is costly to subordinate females, lengthening their interbirth intervals; (ii) females alter their reproductive timings (fertility periods and conceptions) in relation to previous conceptions in the group; and (iii) the reported effect of birth synchrony on interbirth intervals weakens the intensity of reproductive seasonality at the population level. This study emphasizes the importance of sociality in mediating the evolution of reproductive phenology in group-living organisms, a result of broad significance for understanding key demographic parameters driving population responses to increasing climatic fluctuations.

Intolerant baboons avoid observer proximity, creating biased inter-individual association patterns.

Social network analysis is an increasingly popular tool for behavioural ecologists exploring the social organisation of animal populations. Such analyses require data on inter-individual association patterns, which in wild populations are often collected using direct observations of habituated animals. This assumes observers have no influence on animal behaviour; however, our previous work showed that individuals in a habituated group of chacma baboons (Papio ursinus griseipes) displayed consistent and individually distinct responses to observer approaches. We explored the implications of our previous findings by measuring the inter-individual association patterns of the same group of chacma baboons at different observer distances. We found a strong positive association between individual tolerance levels (towards observers) and how often an animal appeared as a neighbour to focal animals when observers were nearer, and a neutral relationship between the same variables when the observer was further away. Additionally, association matrices constructed from different observation distances were not comparable within any proximity buffer, and neither were the individual network metrics generated from these matrices. This appears to be the first empirical evidence that observer presence and behaviour can influence the association patterns of habituated animals and thus have potentially significant impacts on measured social networks.

Nonlethal management of baboons on the urban edge of a large metropole.

The transformation of natural land for agricultural and urban use has displaced baboons from large parts of their historical distribution. Abundant resources within transformed areas, however, continue to attract baboons back into these human-dominated areas resulting in chronic levels of conflict throughout much of Africa. In the city of Cape Town, chacma baboons (Papio ursinus) have been raiding human-derived foods for over 200 years. In the last 20 years, the management of this conflict has shifted from predominantly lethal to nonlethal methods. In this study, we assess the success of field rangers to deter baboons from urban areas and investigate whether such management affects the diet and activity patterns of baboons. We opportunistically sampled baboon behavior and movement when field rangers were temporarily absent from managed troops in 2008. We also compared the intensity of baboon management by comparing one troop on days when the field rangers aggressively herded the troop away from the urban edge versus passively monitored them in 2004/2005. Our results reveal that when field rangers were absent, the two troops spent 70% and 80% of their time within the urban edge compared to 3% and 19% when they were present. Both troops also consumed more human-derived foods when field rangers were absent. There was no significant change in the activity budget or daily distance traveled for either troop with and without field rangers. The intensity of herding did have an impact on baboon activity and high levels of herding significantly reduced time spent feeding and increased time spent traveling, socializing, and resting. Habitat use and dietary composition did not differ between high- and low-herding days. Our results suggest that field rangers are a successful nonlethal method for reducing spatial overlap between baboons and urban areas but that intensive, unsystematic herding of the troop does have measurable impacts on behavior and should be prevented.

Exploring individual variation in associative learning abilities through an operant conditioning task in wild baboons.

Cognitive abilities underpin many of the behavioural decisions of animals. However, we still have very little understanding of how and why cognitive abilities vary between individuals of the same species in wild populations. In this study, we assessed the associative learning abilities of wild chacma baboons (Papio ursinus) across two troops in Namibia with a simple operant conditioning task. We evaluated the ability of individuals to correctly associate a particular colour of corn kernels with a distasteful flavour through repeated presentations of two small piles of corn dyed different colours, one of which had been treated with a non-toxic bitter substance. We also assessed whether individual variation in learning ability was associated with particular phenotypic traits (sex, social rank and neophilia) and states (age and prior vigilance). We found no evidence of learning the association either within each trial or across trials, nor any variation based on individuals' phenotypes. This appeared to be due to a high tolerance for bitter foods leading to similar acceptance of both palatable and unpalatable kernels. Earlier avoidance of the bitter kernels during pilot trials suggests this higher tolerance may have been largely driven by a drought during the experiments. Overall, our findings highlight the potential influence of current environmental challenges associated with conducting cognitive tests of animals in the wild.

Jealous females? Female competition and reproductive suppression in a wild promiscuous primate.

Female-female competition over paternal care has rarely been investigated in promiscuous mammals, where discreet forms of male care have recently been reported despite low paternity certainty. We investigated female competition over paternal care in a wild promiscuous primate, the chacma baboon (Papio ursinus), where pregnant and lactating females establish strong social bonds (friendships) with males that provide care to their offspring. We tested whether pregnant and lactating females interfere with the sexual activity of their male friend to prevent new conceptions that might lead to the subsequent dilution of his paternal care. We found that pregnant and lactating females were more aggressive towards oestrous females when they had recently conceived themselves, and when the oestrous female was mate-guarded by, and showed greater sexual activity with, their male friend. This aggression also reduced the likelihood of conception of the targeted female. These findings indicate that females can aggressively prevent further conceptions with their offspring's carer through reproductive suppression. Competition over access to paternal care may play an important and underestimated role in shaping female social relationships and reproductive strategies in promiscuous mammalian societies.

Isotopic assessment of marine food consumption by natural-foraging chacma baboons on the Cape Peninsula, South Africa.

OBJECTIVES: Stable isotope analysis has been used to investigate consumption of marine resources in a variety of terrestrial mammals, including humans, but not yet in extant nonhuman primates. We sought to test the efficacy of stable isotope analysis as a tool for such studies by comparing isotope- and observation-based estimates of marine food consumption by a troop of noncommensal, free-ranging chacma baboons. MATERIALS AND METHODS: We determined δ13 C and δ15 N values of baboon hair (n = 9) and fecal samples (n = 144), and principal food items (n = 362). These values were used as input for diet models, the outputs of which were compared to observation-based estimates of marine food consumption. RESULTS: Fecal δ13 C values ranged from -29.3‰ to -25.6‰. δ15 N values ranged from 0.9‰ to 6.3‰ and were positively correlated with a measure of marine foraging during the dietary integration period. Mean (± SD) δ13 C values of adult male and female baboon hairs were -21.6‰ (± 0.1) and -21.8‰ (± 0.3) respectively, and corresponding δ15 N values were 5.0‰ (± 0.3) and 3.9‰ (± 0.2). Models indicated that marine contributions were ≤10% of baboon diet within any season, and contributed ≤17% of dietary protein through the year. DISCUSSION: Model output and observational data were in agreement, both indicating that despite their abundance in the intertidal region, marine foods comprised only a small proportion of baboon diet. This suggests that stable isotope analysis is a viable tool for investigating marine food consumption by natural-foraging primates in temperate regions.

Extreme behavioural shifts by baboons exploiting risky, resource-rich, human-modified environments.

A range of species exploit anthropogenic food resources in behaviour known as 'raiding'. Such behavioural flexibility is considered a central component of a species' ability to cope with human-induced environmental changes. Here, we study the behavioural processes by which raiding male chacma baboons (Papio ursinus) exploit the opportunities and mitigate the risks presented by raiding in the suburbs of Cape Town, South Africa. Ecological sampling and interviews conducted with 'rangers' (employed to manage the baboons' space use) revealed that baboons are at risk of being herded out of urban spaces that contain high-energy anthropogenic food sources. Baboon-attached motion/GPS tracking collars showed that raiding male baboons spent almost all of their time at the urban edge, engaging in short, high-activity forays into the urban space. Moreover, activity levels were increased where the likelihood of deterrence by rangers was greater. Overall, these raiding baboons display a time-activity balance that is drastically altered in comparison to individuals living in more remote regions. We suggest our methods can be used to obtain precise estimates of management impact for this and other species in conflict with people.

Male Violence and Sexual Intimidation in a Wild Primate Society.

Sexual violence occurring in the context of long-term heterosexual relationships, such as sexual intimidation, is widespread across human populations [1-3]. However, its evolutionary origins remain speculative because few studies have investigated the existence of comparable forms of sexual coercion in animals [4, 5], in which repeated male aggression toward a female provides the aggressor with delayed mating benefits [6]. Here, we test whether male aggression toward females functions as sexual coercion in wild chacma baboons (Papio ursinus). We found support for all three main predictions of the sexual coercion hypothesis [7]: male aggression (1) is greatest against cycling females, (2) is costly and represents the main source of injuries for cycling females, and (3) increases male mating success with their victims in the future. Detailed analysis of chronological sequences between aggression and matings ruled out other coercive mechanisms, such as short-term harassment and punishment, by showing that aggression and matings are temporally decoupled. This decoupling may explain why some forms of sexual violence have been largely overlooked in well-studied animal populations despite their likely impact on the fitness of both sexes. Finally, we found no support for alternative hypotheses such as a female preference for aggressive males [8, 9]. This new, detailed study of the forms and intensity of sexual intimidation in a wild primate suggests that it may be widespread across mammalian societies, with important implications for understanding the evolution of mate choice and sexual conflict in mammals, as well as the origins of human sexual violence.

Strategic Use of Affiliative Vocalizations by Wild Female Baboons.

Although vocal production in non-human primates is highly constrained, individuals appear to have some control over whether to call or remain silent. We investigated how contextual factors affect the production of grunts given by wild female chacma baboons, Papio ursinus, during social interactions. Females grunted as they approached other adult females 28% of the time. Supporting previous research, females were much more likely to grunt to mothers with young infants than to females without infants. Grunts also significantly increased the likelihood of affiliative interactions among all partners. Notably, however, grunts did not simply mirror existing social bonds. Instead, they appeared to perform a very different function: namely, to serve as signals of benign intent between partners whose relationship is not necessarily close or predictable. Females were less likely to grunt to their mothers or adult daughters-the individuals with whom they shared the closest and least aggressive bonds-than to other females. In contrast, patterns of grunting between sisters were similar to those between nonkin, perhaps reflecting sisters' more ambivalent relationships. Females grunted at higher rates to lower-ranking, than to higher-ranking, females, supporting the hypothesis that grunts do not simply signal the signaler's level of arousal or anxiety about receiving aggression, but instead function as signals of benign intent. Taken together, results suggest that the grunts given by female baboons serve to reduce uncertainty about the likely outcome of an interaction between partners whose relationship is not predictably affiliative. Despite their limited vocal repertoire, baboons appear to be skilled at modifying call production in different social contexts and for different audiences.

Social behavior and patterns of testosterone and glucocorticoid levels differ between male chacma and Guinea baboons.

In multi-male, multi-female groups of mammals, males usually compete aggressively over access to females. However, species vary in the intensity of male contest competition, which has been linked to differences in testosterone and glucocorticoid profiles. Chacma (Papio ursinus) and Guinea (P. papio) baboons constitute an intriguing model to examine variation in male competition and male endocrine correlates, because of the differences in their social systems. Chacma baboons live in stable female-bonded groups with linear male dominance hierarchies and a high male mating skew, whereas Guinea baboons live in male-bonded, multi-level societies. We recorded male behavior and assayed testosterone (fT) and glucocorticoid metabolite (fGC) levels from fecal samples in one population of each species. Male chacma baboons were more frequently involved in agonistic interactions, and dominance relationships were more consistent than in Guinea baboons, where we could not detect linear hierarchies. Notably, male chacma baboons were also more aggressive towards females, indicating an overall higher aggressiveness in this species. In contrast, male Guinea baboons showed higher levels of affiliative interactions and spatial tolerance. High-ranking and consorting male chacma baboons showed elevated fGC levels and also tended to show elevated fT levels, but there was no effect of consortship in Guinea baboons. Agonism was not related to hormone levels in either species. Thus, predictors of fT and fGC levels in Guinea baboons seem to differ from chacma baboons. Our results support the view that different social systems create differential selection pressures for male aggression, reflected by different hormone profiles.

Characterizing Vocal Repertoires--Hard vs. Soft Classification Approaches.

To understand the proximate and ultimate causes that shape acoustic communication in animals, objective characterizations of the vocal repertoire of a given species are critical, as they provide the foundation for comparative analyses among individuals, populations and taxa. Progress in this field has been hampered by a lack of standard in methodology, however. One problem is that researchers may settle on different variables to characterize the calls, which may impact on the classification of calls. More important, there is no agreement how to best characterize the overall structure of the repertoire in terms of the amount of gradation within and between call types. Here, we address these challenges by examining 912 calls recorded from wild chacma baboons (Papio ursinus). We extracted 118 acoustic variables from spectrograms, from which we constructed different sets of acoustic features, containing 9, 38, and 118 variables; as well 19 factors derived from principal component analysis. We compared and validated the resulting classifications of k-means and hierarchical clustering. Datasets with a higher number of acoustic features lead to better clustering results than datasets with only a few features. The use of factors in the cluster analysis resulted in an extremely poor resolution of emerging call types. Another important finding is that none of the applied clustering methods gave strong support to a specific cluster solution. Instead, the cluster analysis revealed that within distinct call types, subtypes may exist. Because hard clustering methods are not well suited to capture such gradation within call types, we applied a fuzzy clustering algorithm. We found that this algorithm provides a detailed and quantitative description of the gradation within and between chacma baboon call types. In conclusion, we suggest that fuzzy clustering should be used in future studies to analyze the graded structure of vocal repertoires. Moreover, the use of factor analyses to reduce the number of acoustic variables should be discouraged.

Ontogenetic scaling of fore- and hind limb posture in wild chacma baboons (Papio hamadryas ursinus).

Large-scale interspecific studies of mammals ranging between 0.04-280 kg have shown that larger animals walk with more extended limb joints. Within a taxon or clade, however, the relationship between body size and joint posture is less straightforward. Factors that may affect the lack of congruence between broad and narrow phylogenetic analyses of limb kinematics include limited sampling of (1) ranges of body size, and/or (2) numbers of individuals. Unfortunately, both issues are inherent in laboratory-based or zoo locomotion research. In this study, we examined the relationship between body mass and elbow and knee joint angles (our proxies of fore- and hind limb posture, respectively) in a cross-sectional ontogenetic sample of wild chacma baboons (Papio hamadryas ursinus) habituated in the De Hoop Nature Reserve, South Africa. Videos were obtained from 33 individuals of known age (12 to ≥ 108 months) and body mass (2-29.5 kg) during walking trials. Results show that older, heavier baboons walk with significantly more extended knee joints but not elbow joints. This pattern is consistent when examining only males, but not within the female sample. Heavier, older baboons also display significantly less variation in their hind limb posture compared to lighter, young animals. Thus, within this ontogenetic sample of a single primate species spanning an order of magnitude in body mass, hind limb posture exhibited a postural scaling phenomenon while the forelimbs did not. These findings may further help explain 1) why younger mammals (including baboons) tend to have relatively stronger bones than adults, and 2) why humeri appear relatively weaker than femora (in at least baboons). Finally, this study demonstrates how field-acquired kinematics can help answer fundamental biomechanical questions usually addressed only in animal gait laboratories.

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.

How do foragers decide when to leave a patch? A test of alternative models under natural and experimental conditions.

A forager's optimal patch-departure time can be predicted by the prescient marginal value theorem (pMVT), which assumes they have perfect knowledge of the environment, or by approaches such as Bayesian updating and learning rules, which avoid this assumption by allowing foragers to use recent experiences to inform their decisions. In understanding and predicting broader scale ecological patterns, individual-level mechanisms, such as patch-departure decisions, need to be fully elucidated. Unfortunately, there are few empirical studies that compare the performance of patch-departure models that assume perfect knowledge with those that do not, resulting in a limited understanding of how foragers decide when to leave a patch. We tested the patch-departure rules predicted by fixed rule, pMVT, Bayesian updating and learning models against one another, using patch residency times (PRTs) recorded from 54 chacma baboons (Papio ursinus) across two groups in natural (n = 6175 patch visits) and field experimental (n = 8569) conditions. We found greater support in the experiment for the model based on Bayesian updating rules, but greater support for the model based on the pMVT in natural foraging conditions. This suggests that foragers may place more importance on recent experiences in predictable environments, like our experiment, where these experiences provide more reliable information about future opportunities. Furthermore, the effect of a single recent foraging experience on PRTs was uniformly weak across both conditions. This suggests that foragers' perception of their environment may incorporate many previous experiences, thus approximating the perfect knowledge assumed by the pMVT. Foragers may, therefore, optimize their patch-departure decisions in line with the pMVT through the adoption of rules similar to those predicted by Bayesian updating.

Peripheral oxytocin in female baboons relates to estrous state and maintenance of sexual consortships.

The neuro-hypophysial hormone oxytocin (OT) has been implicated in female reproductive and maternal behaviors and in the formation of pair bonds in monogamous species. Here we measure variation in urinary OT concentrations in relation to reproductive biology and socio-sexual behavior in a promiscuously breeding species, the chacma baboon (Papio hamadryas ursinus). Subjects were members of a habituated group of baboons in the Okavango Delta, Botswana. We collected behavioral data and urine samples from n=13 cycling females across their estrous cycles and during and outside short-term, exclusive sexual consortships. Samples were analyzed via enzyme immunoassay (EIA) and we used linear mixed models (LMM) to explore the relationship between peripheral OT and a female's estrous stage and consortship status, her previous reproductive experience and fertility. We also used a Pearson's correlation to examine the relationship between OT concentrations of consorting females and their extent of behavioral coordination with their consort partners. The results of the LMM indicate that only estrous stage had a significant influence on OT levels. Females had higher OT levels during their periovulatory period than during other stages of their estrous cycle. There were no differences in the OT levels between consorting and non-consorting periovulatory females. However, among consorting females, there was a significant positive relationship between urinary OT levels and the maintenance of close proximity between consort partners. Our results suggest that physiological and behavioral changes associated with the initiation and maintenance of short-term inter-sexual relationships in baboons correspond with changes in peripheral OT.

Exploring foraging decisions in a social primate using discrete-choice models.

There is a growing appreciation of the multiple social and nonsocial factors influencing the foraging behavior of social animals but little understanding of how these factors depend on habitat characteristics or individual traits. This partly reflects the difficulties inherent in using conventional statistical techniques to analyze multifactor, multicontext foraging decisions. Discrete-choice models provide a way to do so, and we demonstrate this by using them to investigate patch preference in a wild population of social foragers (chacma baboons Papio ursinus). Data were collected from 29 adults across two social groups, encompassing 683 foraging decisions over a 6-month period and the results interpreted using an information-theoretic approach. Baboon foraging decisions were influenced by multiple nonsocial and social factors and were often contingent on the characteristics of the habitat or individual. Differences in decision making between habitats were consistent with changes in interference-competition costs but not with changes in social-foraging benefits. Individual differences in decision making were suggestive of a trade-off between dominance rank and social capital. Our findings emphasize that taking a multifactor, multicontext approach is important to fully understand animal decision making. We also demonstrate how discrete-choice models can be used to achieve this.

Body temperature and physical activity correlates of the menstrual cycle in Chacma Baboons (Papio hamadryas ursinus).

We investigated the temporal relationship between abdominal temperature, physical activity, perineal swelling, and urinary progesterone and estradiol concentrations over the menstrual cycle in unrestrained captive baboons. Using a miniature temperature-sensitive data logger surgically implanted in the abdominal cavity and an activity data logger implanted subcutaneously on the trunk, we measured, continuously over 6 months at 10-min intervals, abdominal temperature and physical activity patterns in four female adult baboons Papio hamadryas ursinus (12.9-19.9 kg), in cages in an indoor animal facility (22-25°C). We monitored menstrual bleeding and perineal swelling changes, and measured urinary progesterone and estradiol concentrations, daily for up to 6 months, to ascertain the stage and length of the menstrual cycle. The menstrual cycle was 36 ± 2 days (mean ± SD) long and the baboons exhibited cyclic changes in perineal swellings, abdominal temperature, physical activity, urinary progesterone, and estradiol concentrations over the cycle. Mean 24-hr abdominal temperature during the luteal phase was significantly higher than during the periovulatory phase (ANOVA, F((2, 9)) = 4.7; P = 0.04), but not different to that during the proliferative phase. Physical activity followed a similar pattern, with mean 24-hr physical activity almost twice as high in the luteal than in the periovulatory phase (ANOVA, P = 0.58; F((2, 12)) = 5.8). We have characterized correlates of the menstrual cycle in baboons and shown, for the first time, a rhythm of physical activity and abdominal temperature over the menstrual cycle, with a nadir of temperature and activity at ovulation.

Derived vocalizations of geladas (Theropithecus gelada) and the evolution of vocal complexity in primates.

Primates are intensely social and exhibit extreme variation in social structure, making them particularly well suited for uncovering evolutionary connections between sociality and vocal complexity. Although comparative studies find a correlation between social and vocal complexity, the function of large vocal repertoires in more complex societies remains unclear. We compared the vocal complexity found in primates to both mammals in general and human language in particular and found that non-human primates are not unusual in the complexity of their vocal repertoires. To better understand the function of vocal complexity within primates, we compared two closely related primates (chacma baboons and geladas) that differ in their ecology and social structures. A key difference is that gelada males form long-term bonds with the 2-12 females in their harem-like reproductive unit, while chacma males primarily form temporary consortships with females. We identified homologous and non-homologous calls and related the use of the derived non-homologous calls to specific social situations. We found that the socially complex (but ecologically simple) geladas have larger vocal repertoires. Derived vocalizations of geladas were primarily used by leader males in affiliative interactions with 'their' females. The derived calls were frequently used following fights within the unit suggesting that maintaining cross-sex bonds within a reproductive unit contributed to this instance of evolved vocal complexity. Thus, our comparison highlights the utility of using closely related species to better understand the function of vocal complexity.

Evaluating home range techniques: use of Global Positioning System (GPS) collar data from chacma baboons.

Global Positioning System (GPS) collars have revolutionized the field of spatial ecology, but to date, few primate studies have used them. We fitted a free-ranging, semi-habituated, juvenile male chacma baboon (Papio hamadryas ursinus) with an automatic self-releasing GPS collar and tracked his movements for 359 days. The collar captured 4254 fixes out of 5719 programmed opportunities, a 74.4 % acquisition rate, suggesting that the collar effectively tracked this baboon in a variety of habitat types. Of the data points captured, 73.7 % were three-dimensional fixes, and of these fixes, 66.9 % were highly accurate, having a dilution of precision of less than four. We calculated home range using three protocols with three estimation methods: minimum convex polygon, fixed kernel-density estimation (KDE), and fixed r local convex hull. Using all data points and the 95 % contour, these methods created home range estimations ranging from 10.8 to 23.1 km(2) for this baboon troop. Our results indicate that the KDE output using all data locations most accurately represented our data set, as it created a continuous home range boundary that excluded unused areas and outlying, potentially exploratory data points while including all seven sleeping sites and a movement corridor. However, home range estimations generated from KDE varied from 15.4 to 18.8 km(2) depending on the smoothing parameter used. Our results demonstrated that the ad hoc smoothing parameter selection technique was a better method for our data set than either the least squares cross-validation or biased cross-validation techniques. Our results demonstrate the need for primatologists to develop a standardized reporting method which documents the tool, screening protocol, and smoothing parameter used in the creation of home range estimations in order to make comparisons that are meaningful.