Dawn M. Kitchen

Baboon loud calls advertise male quality: acoustic features and their relation to rank, age, and exhaustion

Free-ranging adult male baboons give loud two-syllable ‘wahoo’ calls during dawn choruses, interactions between groups, when chasing females, and in aggressive interactions with other males. Previous research has shown that the rate and duration of these contest wahoos are correlated with a male’s competitive ability: high-ranking males call more often, call at faster rates, and call for longer bouts than do low-ranking males. Here we report that acoustic features of wahoos also reveal information about male competitive ability. High-ranking males give wahoos with higher fundamental frequencies (F0) and longer ‘hoo’ syllables. Within-subject analyses revealed that, as males fall in rank, the hoo syllables tend to shorten within a period of months. As males age and continue to fall in rank, F0 declines, hoo syllables shorten, and formant dispersion decreases. Independent of age and rank, within bouts of calling F0 declines and hoo syllables become shorter. Because wahoos are often given while males are running or leaping through trees, variation in these acoustic features may function as an indicator of a male’s stamina. The acoustic features of contest wahoos thus potentially allow listeners to assess a male’s competitive ability.

Factors Affecting Reproduction and Mortality Among Baboons in the Okavango Delta, Botswana

We present results of a 10-year study of free-ranging gray-footed chacma baboons (Papio ursinus griseipes) in the Okavango Delta of Botswana. The majority of deaths among adult females and juveniles were due to predation, while infants were more likely to die of infanticide. There were strong seasonal effects on birth and mortality, with the majority of conceptions occurring during the period of highest rainfall. Mortality due to predation and infanticide was highest during the 3-mo period when flooding was at its peak, when the group was more scattered and constrained to move along predictable routes. The reproductive parameters most likely to be associated with superior competitive ability—interbirth interval and infant growth rates—conferred a slight fitness advantage on high-ranking females. However, it was counterbalanced by the effects of infanticide and predation. Infanticide affected high- and low-ranking females more than middle-ranking females, while predation affected females of all ranks relatively equally. As a result, there were few rank-related differences in estimated female lifetime reproductive success.

Factors affecting individual participation in group-level aggression among non-human primates

Summary Group members do not always act cohesively when facing extra-group rivals. When benefits such as group-defence are not monopolizable, it poses an economics problem: who should contribute to public goods and who should freeload? A collective action framework compliments existing theoretical explanations for cooperation, and provides testable hypotheses about group-level behaviour based on individual costs and benefits. Using this approach, we review research on intergroup encounters in non-human primates published over the last 20 years, focusing on participation by different classes of individuals. While food- and matedefence explain much between-sex variation in participation, rank and reproductive access frequently explain within-sex variation. In some species, individuals may use intergroup interactions to survey potential transfer locations and mating options, which might coincidently intimidate rivals. Experimental evidence suggests that when intergroup dominance is based on relative number of fighters, individual participation still varies with sex, rank, companion behaviour and dependent offspring presence. Relatively few studies have examined how factors such as relationships within and between groups or individual temperament mediate aggression. Long-term studies of multiple habituated groups and methodological advances (e.g., playback experiments) will continue to improve our understanding of how complex group-level patterns are predictable when viewed from an individual perspective.

Hedging their bets? Male and female chacma baboons form friendships based on likelihood of paternity

In chacma baboons (Papio hamadryas ursinus), adult males and lactating females form preferential associations, or ‘friendships’, that provide protection against potentially infanticidal attacks. Little is known about the mechanisms by which males and females form friendships, or the function of friendships for males. We examined the relationship between mating activity, paternity and friendships over a 3-year period. A male’s proportion of consort activity with a female within and outside of her period of most likely conception, along with his dominance rank, each contributed importantly in models of paternity outcomes and formation of friendships. In 87% of births, the male that monopolized the greatest proportion of a female’s consort days during her last conceptive cycle was the father of her offspring. In 82% of the births where the female’s primary consort partner was present at parturition, he became her friend. When the primary consort partner was absent at parturition, females formed friendships with males that had monopolized the second or third greatest proportion of their previous consort days. Similarly, 78% of males became friends with females with whom they had obtained their greatest consort success. Such a flexible strategy based on probabilistic assessment of paternity may be adaptive for females, given unpredictable fluctuations in male presence within the group. In a mating system with high dominance-based reproductive skew, males may benefit by investing preferentially in those infants that they are most likely to have fathered.

FACTORS MEDIATING INTER-GROUP ENCOUNTERS IN SAVANNAH BABOONS (PAPIO CYNOCEPHALUS URSINUS)

Summary Inter-group encounters among baboons range from peaceful to aggressive. During 23 months we observed 110 inter-group interactions involving four groups of chacma baboons in the Okavango Delta, Botswana. Results supported the hypothesis that male behavior functions to prevent extra-group males from gaining access to sexually receptive females. Males were more likely to chase females in their own group when estrous females were present, and their chases targeted estrous females more often than expected. Males also chased members of the opposing group more when estrous females were present. When estrous females were absent, male displays were shorter in duration, involved fewer participants, were less intense, and were more likely to result in peaceful mingling between groups. The alpha male was the individual most actively involved in inter-group chases and displays, but males of all ranks participated, especially when they were in consort with a female. However, males did not cooperate in group defense. While behavior during encounters was affected by the presence of

Male chacma baboons (Papio hamadryas ursinus) discriminate loud call contests between rivals of different relative ranks

Males in multi-male groups of chacma baboons (Papio hamadryas ursinus) in Botswana compete for positions in a linear dominance hierarchy. Previous research suggests that males treat different categories of rivals differently; competitive displays between males of similar rank are more frequent and intense than those between disparately ranked males. Here we test whether males also respond differently to male–male interactions in which they are not directly involved, using playbacks of the loud ‘wahoo’ calls exchanged between competing males in aggressive displays. We played paired sequences of vocal contests between two adjacently ranked and two disparately ranked males to ten subjects, half ranking below the signalers in the call sequences and half above. Subjects who ranked above the two signalers showed stronger responses than lower-ranking subjects. Higher-ranking subjects also responded more strongly to sequences involving disparately ranked, as opposed to adjacently ranked opponents, suggesting that they recognized those individuals’ relative ranks. Strong responses to sequences between disparately ranked opponents might have occurred either because such contests typically involve resources of high fitness value (defense of meat, estrous females or infants vulnerable to infanticide) or because they indicate a sudden change in one contestant’s condition. In contrast, subjects who ranked lower than the signalers responded equally strongly to both types of sequences. These subjects may have been able to distinguish between the two categories of opponents but did not respond differently to them because they had little to lose or gain by a rank reversal between males that already ranked higher than they did.

Subordinate male black howler monkey (Alouatta pigra) responses to loud calls: Experimental evidence for the effects of intra-group male relationships and age

When assessing a groups overall fighting ability, functional group size (i. e. the number of cooperative members willing to confront opponents) may be more important than actual group size. Despite obvious benefits, group members do not always act in a collective manner. For example, participation by subordinate male black howler monkeys (Alouatta pigra) in natural group-level contests is highly variable. Using playback experiments, we examined whether alpha-subordinate relationships, number of intruders, or age affected subordinate contest participation. Long-term census data were used to estimate the nature and duration of intra-group male associations. Some subordinate males had long-term relationships with the alpha that existed prior to residence in the current group or they were the alpha males grown sons. Other subordinate males were in more recent associations with the alpha that formed under seemingly antagonistic circumstances (e. g. the deposed alpha or his grown sons). We found that subordinate males in long-term alpha-subordinate associations had stronger howling and approach responses than males in short-term associations. Younger long-term associates had the strongest reactions to simulated intruders, similarly aged shortterm associates rarely responded, and older males in both association categories responded at intermediate levels. We discuss the variable strategies of subordinate male black howlers and suggest that males who were more likely to participate in group-defence might be gaining direct or indirect fitness benefits by group living.

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.

Comparing responses of four ungulate species to playbacks of baboon alarm calls

A growing body of evidence suggests that a wide range of animals can recognize and respond appropriately to calls produced by other species. Social learning has been implicated as a possible mechanism by which heterospecific call recognition might develop. To examine whether familiarity and/or shared vulnerability with the calling species might influence the ability of sympatric species to distinguish heterospecific alarm calls, we tested whether four ungulate species (impala: Aepyceros melampus; tsessebe: Damaliscus lunatus; zebra: Equus burchelli; wildebeest: Connochaetes taurinus) could distinguish baboon (Papio hamadryas ursinus) alarm calls from other loud baboon calls produced during intra-specific aggressive interactions (‘contest’ calls). Overall, subjects’ responses were stronger following playback of alarm calls than contest calls. Of the species tested, impala showed the strongest responses and the greatest difference in composite response scores, suggesting they were best able to differentiate call types. Compared with the other ungulate species, impala are the most frequent associates of baboons. Moreover, like baboons, they are susceptible to both lion and leopard attacks, whereas leopards rarely take the larger ungulates. Although it seems possible that high rates of association and/or shared vulnerability may influence impala’s greater ability to distinguish among baboon call types, our results point to a stronger influence of familiarity. Ours is the first study to compare such abilities among several community members with variable natural histories, and we discuss future experiments that would more systematically examine development of these skills in young ungulates.

Coprological Surveys of Alouatta pigra at Two Sites in Belize

Fecal samples were collected from black howler monkeys (Alouatta pigra) in north central Belize and analyzed for evidence of endoparasite life stages. At least six types of endoparasites were found in Alouatta pigra fecal samples collected in the Lamanai Archaeological Reserve and the Community Baboon Sanctuary in 1999. These include a digenean trematode, an oxyurid nematode, a strongyle-type nematode, an ascarid presumed to be Ascaris sp., Entamoeba coli and Iodamoeba bütschlii. Higher trematode prevalence was found in adult Alouatta pigra compared to juveniles and higher prevalence of nematode larvae in all animals was found in the wet season compared to the dry season.