Topological spatial representation in wild chacma baboons (Papio ursinus).

Many species orient towards specific locations to reach important resources using different cognitive mechanisms. Some of these, such as path integration, are now well understood, but the cognitive orientation mechanisms that underlie movements in non-human primates remain the subject of debate. To investigate whether movements of chacma baboons are more consistent with Euclidean or topological spatial awareness, we investigated whether baboons made repeated use of the same network of pathways and tested three predictions resulting from the hypothesized use of Euclidean and topological spatial awareness. We recorded ranging behaviour of a group of baboons during 234 full days and 137 partial days in the Soutpansberg Mountains, South Africa. Results show that our baboons travelled through a dense network of repeated routes. In navigating this route network, the baboons did not approach travel goals from all directions, but instead approached them from a small number of the same directions, supporting topological spatial awareness. When leaving travel goals, baboons' initial travel direction was significantly different from the direction to the next travel goal, again supporting topological spatial awareness. Although we found that our baboons travelled with similar linearity in the core area as in the periphery of their home range, this did not provide conclusive evidence for the existence of Euclidean spatial awareness, since the baboons could have accumulated a similar knowledge of the periphery as of the core area. Overall, our findings support the hypothesis that our baboons navigate using a topological map.

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.

Sequential phenotypic constraints on social information use in wild baboons.

Social information allows the rapid dissemination of novel information among individuals. However, an individual's ability to use information is likely to be dependent on phenotypic constraints operating at three successive steps: acquisition, application, and exploitation. We tested this novel framework by quantifying the sequential process of social information use with experimental food patches in wild baboons (Papio ursinus). We identified phenotypic constraints at each step of the information use sequence: peripheral individuals in the proximity network were less likely to acquire and apply social information, while subordinate females were less likely to exploit it successfully. Social bonds and personality also played a limiting role along the sequence. As a result of these constraints, the average individual only acquired and exploited social information on.

Wild chacma baboons (Papio ursinus) remember single foraging episodes.

Understanding animal episodic-like memory is important for tracing the evolution of the human mind. However, our knowledge about the existence and nature of episodic-like memory in non-human primates is minimal. We observed the behaviour of a wild male chacma baboon faced with a trade-off between protecting his stationary group from aggressive extra-group males and foraging among five out-of-sight platforms. These contained high-priority food at a time of natural food shortage. In 10 morning and eight evening trials, the male spontaneously visited the platforms in five and four different sequences, respectively. In addition, he interrupted foraging sequences at virtually any point on eight occasions, returning to the group for up to 2 h. He then visited some or all of the remaining platforms and prevented revisits to already depleted ones, apparently based on his memory for the previous foraging episode about food value, location, and time. Efficient use of memory allowed him to keep minimal time absent from his group while keeping food intake high. These findings support the idea that episodic-like memory offers an all-purpose solution to a wide variety of problems that require flexible, quick, yet precise decisions in situations arising from competition for food and mates in wild primates.

Evidence for varying social strategies across the day in chacma baboons.

Strong social bonds can make an important contribution to individual fitness, but we still have only a limited understanding of the temporal period relevant to the adjustment of social relationships. While there is growing recognition of the importance of strong bonds that persist for years, social relationships can also vary over weeks and months, suggesting that social strategies may be optimized over shorter timescales. Using biological market theory as a framework, we explore whether temporal variation in the benefits of social relationships might be sufficient to generate daily adjustments of social strategies in wild baboons. Data on grooming, one measure of social relationships, were collected from 60 chacma baboons (Papio ursinus) across two troops over a six month period. Our analyses suggest that social strategies can show diurnal variation, with subordinates preferentially grooming more dominant individuals earlier in the day compared with later in the day. These findings indicate that group-living animals may optimize certain elements of their social strategies over relatively short time periods.

Change point analysis of travel routes reveals novel insights into foraging strategies and cognitive maps of wild baboons.

Efficient space use is a critical challenge for animals relying on stationary resources. It is often difficult with purely observational methods to gain unambiguous insight into any ability of primates to manage and process spatial information. Investigating the visible signs of the decision processes underlying space use often leaves open important issues. We applied the change point test [Byrne et al. (2009). Anim Behav 77: 619-631], a statistical tool to objectively determine change points (CPs) in animal travel paths, to investigate to what degree directional changes in our study group's (Papio ursinus) dry season ranging were associated with important resources and prominent landmarks. One-third of directional changes were associated with fruit feeding, 1/3 with traveling, and 1/3 with dry matter feeding, travel feeding and with drinking. When directional changes were associated with traveling, the subsequent directional changes were likely to result in fruit feeding. Fruit feeding mostly occurred at the apex of the day journeys, while drinking, dry matter feeding, and travel feeding often occurred along straight travel segments. The majority of directional changes did not occur in clusters at distinctive locations, but at distances of more than 120 m apart from each other, many of them along prominent landmarks. We conclude that the CPs do not represent nodes or route bends of a network map. Rather, they represent (1) locations where the decision to turn back to their sleeping site was taken, and (2) locations next to important landmarks (changes of slope, car tracks) where slight adjustment of a movement direction was possible. We found no evidence for a Euclidean map and discuss our findings in the light of a network map representation of space.

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.

Group decision-making in chacma baboons: leadership, order and communication during movement.

BACKGROUND: Group coordination is one of the greatest challenges facing animals living in groups. Obligatory trade-offs faced by group members can potentially lead to phenomena at the group level such as the emergence of a leader, consistent structure in the organization of individuals when moving, and the use of visual or acoustic communication. This paper describes the study of collective decision-making at the time of departure (i.e. initiation) for movements of two groups of wild chacma baboons (Papio ursinus). One group was composed of 11 individuals, whilst the other consisted of about 100 individuals. RESULTS: Results for both groups showed that adult males initiated more movements even if the leadership was also distributed to adult females and young individuals. Baboons then joined a movement according to a specific order: adult males and adult females were at the front and the back of the group, sub-adults were at the back and juveniles were located in the central part of the progression. In the two groups, vocalisations, especially loud calls, were more frequently emitted just before the initiation of a group movement, but the frequency of these vocalisations did not influence the success of an initiation in any way. CONCLUSION: The emergence of a leadership biased towards male group members might be related to their dominance rank and to the fact that they have the highest nutrient requirements in the group. Loud calls are probably not used as recruitment signals but more as a cue concerning the motivation to move, therefore enhancing coordination between group members.

The dining etiquette of desert baboons: the roles of social bonds, kinship, and dominance in co-feeding networks.

To better understand how individual relationships influence patterns of social foraging in primate groups, we explored networks of co-feeding in wild desert baboons (Papio ursinus). To minimize the risk of aggression and injury associated with contest competition, we expected that individual group members would choose to co-feed with those group-mates that are most likely to show tolerance and a willingness to share food patches. We tested two alternative hypotheses about who those group-mates might be: the "social bonds hypothesis" predicts that preferred foraging partners will be those with whom individuals share strong social bonds, indexed by grooming, whereas the "kinship hypothesis" predicts that preferred foraging partners will be relatives. We also investigated and controlled for the effects of dominance rank, given that competitive ability is known to shape foraging patterns. Social network analyses of over 5,000 foraging events for 14 adults in a single troop revealed that baboon co-feeding was significantly correlated with grooming relationships but not genetic relatedness, and this finding was also true of the female-only co-feeding network. Dominant individuals were also found to be central to the co-feeding network, frequently sharing food patches with multiple group-mates. This polyadic analysis of foraging associations between individuals underlines the importance of dominance and affiliation to patterns of primate social foraging.

Experimental evidence for limited vocal recognition in a wild primate: implications for the social complexity hypothesis.

Although monitoring social information is a key aspect of the social complexity hypothesis, surprisingly little work has compared social knowledge across different species of wild animals. In the present study, I use playback experiments to test for individual recognition in wild male geladas (Theropithecus gelada) to compare with published accounts of social knowledge in chacma baboons (Papio ursinus). Geladas and baboons are closely related primates living in socially complex groups that differ dramatically in group size-geladas routinely associate with more than 10 times the number of conspecifics than do baboons. Using grunts from non-rival males to simulate approaches, I examined the strength of a subject male's response when the 'approach' was from the direction of (i) non-rival males (control), or (ii) rival males (a more salient stimulus if playback grunts are not recognized by the subject). I compared responses separately based on the degree of social overlap between the caller and the subject. Responses indicate that male geladas, unlike baboons, do not use vocalizations to recognize all of the individuals they regularly encounter. This represents, to my knowledge, the first documented evidence of 'missing' social knowledge in a natural primate population. The sharp distinction between baboons and geladas suggests that geladas are either unable or unmotivated to keep track of the individual identity of other males in their multi-level society-even males with whom they have a large degree of social overlap. Thus, these results are consistent with the central assumption of the social complexity hypothesis that social cognition is costly.

Infanticide and reproductive restraint in a polygynous social mammal.

Alpha male chacma baboons experience uncontested access to individual estrus females. Consequently, alpha male paternity certainty is high and underpins significant levels of infanticide by immigrant males that, in turn, has selected for male defense of infants. There is also, however, a high probability that alpha males will be absent during the period when their own offspring are vulnerable, suggesting selection for additional countermeasures. We use data from a long-term study to test the prediction that alpha male chacma baboons cede reproductive opportunities to subordinate males and that this leads to the presence of other fathers that can serve as a buffer against infanticidal attack. We found that subordinate males obtained significantly more conceptive opportunities than predicted by priority of access alone, and that this occurred because alpha males did not consort all receptive periods. There was no evidence that this was due to energetic constraint, large male cohorts, alpha male inexperience, or the competitive strength of queuing subordinates. The number of males who benefited from concession and the length of time that they were resident relative to those who did not benefit in this way greatly reduced the probability that infants of alpha males would face immigrant males without a surrogate father whose own offspring were vulnerable. The absence of such males was associated with observed infanticide as well as, unexpectedly, an increased likelihood of takeover when alpha males with vulnerable infants were present.

How do wild baboons (Papio ursinus) plan their routes? Travel among multiple high-quality food sources with inter-group competition.

How do humans and animals travel between multiple destinations on a given foraging trip? This question is of theoretical and practical interest, yet few empirical data exist to date. We examined how a group of wild chacma baboons travelled among multiple, simultaneously fruiting mountain fig trees (Ficus glumosa). In the course of a 16-month study, this highly preferred fruit was available during a 3-week period, from relatively few sites, which were also utilized by four larger baboon groups. We used directness of route and travel speed of 13 days of observation, and approach rates of 31 days of observation to differentiate between purposeful and opportunistic encounters with 50 fig trees. The study group visited a total of 30 fig trees overall, but only 8 trees per day on average. Each morning, they travelled along a highly repetitive route on all days of observation, thereby visiting 2-4 fig trees. They approached these trees rapidly along highly directed paths without intermittently exploiting other food sources that were available in large quantities. Then, they abruptly changed behaviour, switching to lower travel speed and less directed routes as they foraged on a variety of foods. They approached additional fig trees later in the day, but approach rates were similar to those at times of year when fruit of this fig species was unavailable; this suggested that encounters with trees after the behavioural switch were not planned. Comparing visits to purposefully and opportunistically encountered trees, we found no difference in the average time spent feeding or frequency of feeding supplants, suggesting that purposefully and opportunistically visited trees had similar values. We conclude that when foraging for mountain fig fruit the baboons' cognitive maps either contain information on relatively few trees or of only a single route along which several trees are situated, leading to very limited planning abilities.

Comparing responses to novel objects in wild baboons (Papio ursinus) and geladas (Theropithecus gelada).

Behavioral flexibility is considered by some to be one of the hallmarks of advanced cognitive ability. One measure of behavioral flexibility is how subjects respond to novel objects. Despite growing interest in comparative cognition, no comparative research on neophilia in wild primates has been conducted. Here, we compare responses to novel objects in wild chacma baboons (Papio ursinus) and geladas (Theropithecus gelada). Baboons and geladas are closely related taxa, yet they differ in their ecology and degree of social tolerance: (1) baboons are habitat and dietary generalists, whereas geladas have one of the most specialized primate diets (90% grass); (2) baboons exhibit an aversion toward extra-group individuals, whereas geladas typically exhibit an attraction toward them. Using subjects of all age and sex classes, we examined responses to three different objects: a plastic doll, a rubber ball, and a metal can. Overall, baboon subjects exhibited stronger responses to the objects (greater neophilia and exploration) than gelada subjects, yet we found no evidence that the geladas were afraid of the objects. Furthermore, baboons interacted with the objects in the same way they might interact with a potential food item. Responses were unrelated to sex, but immatures showed more object exploration than adults. Results corroborate novel object research conducted in captive populations and suggest that baboons and geladas have differences in behavioral flexibility (at least in this cognitive domain) that have been shaped by ecological (rather than social) differences between the two species.

Dominance and affiliation mediate despotism in a social primate.

Group-living animals routinely have to reach a consensus decision and choose between mutually exclusive actions in order to coordinate their activities and benefit from sociality. Theoretical models predict "democratic" rather than "despotic" decisions to be widespread in social vertebrates, because they result in lower "consensus costs"-the costs of an individual foregoing its optimal action to comply with the decision-for the group as a whole. Yet, quantification of consensus costs is entirely lacking, and empirical observations provide strong support for the occurrence of both democratic and despotic decisions in nature. We conducted a foraging experiment on a wild social primate (chacma baboons, Papio ursinus) in order to gain new insights into despotic group decision making. The results show that group foraging decisions were consistently led by the individual who acquired the greatest benefits from those decisions, namely the dominant male. Subordinate group members followed the leader despite considerable consensus costs. Follower behavior was mediated by social ties to the leader, and where these ties were weaker, group fission was more likely to occur. Our findings highlight the importance of leader incentives and social relationships in group decision-making processes and the emergence of despotism.

Mental maps in chacma baboons (Papio ursinus): using inter-group encounters as a natural experiment.

Encounters between groups of wild chacma baboons (Papio ursinus) can be viewed as a natural experiment to investigate the nature of these primates' mental representations of large-scale space. During a 16-month field study in a high population density habitat we recorded the foraging routes and the most important resources of a group of 25 individuals. Also, we estimated the locations of additional baboon groups relative to the study group. Routes were less linear, travel speed was higher, and inter-resource distances were larger when other groups were present within 500 m of the focal group; thus, the study group avoided others by taking detours. We predicted that evasive manoeuvres would be characteristic of different possible orientation mechanisms, and compared them with our observations. We analysed 34 evasive manoeuvres in detail. In an area that lacked prominent landmarks, detours were small; larger detours occurred when resources were directly visible, or in the vicinity of a hill offering conspicuous landmarks. In areas without prominent landmarks, detours were along familiar routes and waiting bouts of up to 60 min occurred; on one occasion the study group aborted their entire day's journey. We discuss these findings in the light of time and energy costs and suggest that the baboons lack the ability to compute Euclidean relations among locations, but use network maps to find their way to out-of-sight locations.