Carson M. Murray

Male dominance rank and reproductive success in chimpanzees, Pan troglodytes schweinfurthii

Competition for fertile females determines male reproductive success in many species. The priority of access model predicts that male dominance rank determines access to females, but this model has been difficult to test in wild populations, particularly in promiscuous mating systems. Tests of the model have produced variable results, probably because of the differing socioecological circumstances of individual species and populations. We tested the predictions of the priority of access model in the chimpanzees of Gombe National Park, Tanzania. Chimpanzees are an interesting species in which to test the model because of their fission-fusion grouping patterns, promiscuous mating system and alternative male mating strategies. We determined paternity for 34 offspring over a 22-year period and found that the priority of access model was generally predictive of male reproductive success. However, we found that younger males had higher success per male than older males, and low-ranking males sired more offspring than predicted. Low-ranking males sired offspring with younger, less desirable females and by engaging in consortships more often than high-ranking fathers. Although alpha males never sired offspring with related females, inbreeding avoidance of high-ranking male relatives did not completely explain the success of low-ranking males. While our work confirms that male rank typically predicts male chimpanzee reproductive success, other factors are also important; mate choice and alternative male strategies can give low-ranking males access to females more often than would be predicted by the model. Furthermore, the success of younger males suggests that they are more successful in sperm competition.

Dominance rank influences female space use in wild chimpanzees, Pan troglodytes: towards an ideal despotic distribution

Studies from many different taxa have demonstrated that dominance rank greatly influences individual space use. While the importance of dominance among female chimpanzees, Pan troglodytes, was debated in the past, mounting evidence now shows that rank is very important. In particular, rank has been shown to influence body mass, foraging strategies, association patterns, and ultimately, reproductive success. In this study, we investigated how rank influenced female space use among chimpanzees, P.t. schweinfurthii, at Gombe National Park, Tanzania. Analysing 10 years of data, we found that new immigrants used areas away from dominant females, and that subordinates had lower site fidelity. We also found that high-ranking females had smaller core areas and that this size difference was pronounced during periods of food scarcity when food competition is highest. These patterns suggest that dominant females outcompete subordinates, forcing them to settle elsewhere, range more widely, and shift their space use across time.

Severe Aggression Among Female Pan troglodytes schweinfurthii at Gombe National Park, Tanzania

Aggression is generally more severe between males than between females because males gain greater payoffs from escalated aggression. Males that successfully defeat rivals may greatly increase their access to fertile females. Because female reproductive success depends on long-term access to resources, competition between females is often sustained but low key because no single interaction leads to a high payoff. Nonetheless, escalated aggression can sometimes immediately improve a female’s reproductive success. Resisting new immigrants can reduce feeding competition, and infanticide of other females’ young can increase a female’s access to resources. East African chimpanzees live in fission-fusion communities in which females occupy overlapping core areas. Growing evidence indicates that reproductive success correlates with core area quality, and that females compete for long-term access to core areas. Here we document 5 new cases of severe female aggression in the context of such competition: 2 attacks by resident females on an immigrant female, a probable intracommunity infanticide, and 2 attacks on a female and her successive newborn infants by females whose core areas overlapped hers. The cases provide further evidence that females are occasionally as aggressive as males. Factors influencing the likelihood and severity of such attacks include rank and size differences and the presence of dependable allies. Counterstrategies to the threat of female aggression include withdrawing from others around the time of parturition and seeking male protection. We also discuss an unusual case of a female taking the newborn infant of another, possibly to protect it from a potentially infanticidal female.

Adult Male Chimpanzees Inherit Maternal Ranging Patterns

Space use often correlates with reproductive success [1, 2]. Individual site fidelity is ubiquitous across a variety of taxa, including birds, mammals, insects, and reptiles [3-9]. Individuals can benefit from using the same area because doing so affords access to known resources, including food and/or breeding sites. The majority of studies on site fidelity have focused upon strictly territorial species in which individuals range in well-defined, exclusive areas (e.g., [4, 9]). By comparison, the transient groups that define fission-fusion species allow for considerable flexibility in individual space use. Although there is evidence that individual space use can influence reproductive success [2], relatively little is known about individual ranging patterns in fission-fusion species. Here, we investigate three potential correlates of male site fidelity (age, habitat quality, and maternal space use) in wild chimpanzees (Pan troglodytes). We found that when alone, each male preferentially concentrated his space use near the area where his mother ranged when he was dependent. We suggest that solitary ranging allows males to avoid direct competition with conspecifics and that foraging in familiar areas maximizes foraging efficiency. These results highlight the importance of male foraging strategies in a species in which male ranging is typically explained in terms of mating access to females.

Alpha male chimpanzee grooming patterns: Implications for dominance "style"

In social primates, individuals use various tactics to compete for dominance rank. Grooming, displays and contact aggression are common components of a male chimpanzees dominance repertoire. The optimal combination of these behaviors is likely to differ among males with individuals exhibiting a dominance “style” that reflects their tendency to use cooperative and/or agonistic dominance tactics. Here, we examine the grooming behavior of three alpha male chimpanzees at Gombe National Park, Tanzania. We found that (1) these males differed significantly in their tendency to groom with other males; (2) each males grooming patterns remained consistent before, during and after his tenure as alpha, and (3) the three males tended to groom with high‐ middle‐ and low‐ranking partners equally. We suggest that body mass may be one possible determinant of differences in grooming behavior. The largest male exhibited the lowest overall grooming rates, whereas the smallest male spent the most time grooming others. This is probably because large males are more effective at physically intimidating subordinates. To achieve alpha status, a small male may need to compensate for reduced size by investing more time and energy in grooming, thereby ensuring coalitionary support from others. Rates of contact aggression and charging displays conformed to this prediction, suggesting that each male exhibited a different dominance “style.” Am. J. Primatol. 71:136–144, 2009.

Boys will be boys: sex differences in wild infant chimpanzee social interactions

Sex differences in the behaviour of human children are a hotly debated and often controversial topic. However, several recent studies have documented a biological basis to key aspects of child social behaviour. To further explore the evolutionary basis of such differences, we investigated sex differences in sociability in wild chimpanzee, Pan troglodytes schweinfurthii, infants at Gombe National Park, Tanzania. We used a long-term data set on mother-infant behaviour to analyse the diversity of infant chimpanzee social partners from age 30 to 36 months. Male infants (N = 12) interacted with significantly more individuals than female infants did (N = 8), even when maternal sociability was controlled for. Furthermore, male infants interacted with significantly more adult males than female infants did. Our data indicate that the well-documented sex differences in adult chimpanzee social tendencies begin to appear quite early in development. Furthermore, these data suggest that the behavioural sex differences of human children are fundamentally rooted in our biological and evolutionary heritage.

The Appearance and Spread of Ant Fishing among the Kasekela Chimpanzees of Gombe: A Possible Case of Intercommunity Cultural Transmission.

Chimpanzees exhibit cultural variation, yet examples of successful cultural transmission between wild communities are lacking. Here we provide the first account of tool-assisted predation (“ant fishing”) on Camponotus ants by the Kasekela and Mitumba communities of Gombe National Park. We then consider three hypotheses for the appearance and spread of this behavior in Kasekela: (1) changes in prey availability or other environmental factors, (2) innovation, and (3) introduction. Ant fishing was recognized as habitual in the Mitumba community by 1992, soon after their habituation began. Apart from one session in 1978, Camponotus predation (typically with tools) was documented in the Kasekela community beginning only in 1994, despite decades of prior observation. By February 2010, ant fishing was customary in Kasekela and with one exception was practiced exclusively by chimpanzees born after 1981 and immigrant females. We hypothesize that changes in insect prey availability over time and/or the characteristics of one popular ant-fishing site may have influenced the establishment of ant fishing. Though innovation cannot be completely ruled out, the circumstantial evidence suggests that a Mitumba immigrant introduced ant fishing to Kasekela. We submit that this report represents the first documented case of successful transmission of a novel cultural behavior between wild chimpanzee communities.

Sex Differences in Wild Chimpanzee Behavior Emerge during Infancy

The role of biological and social influences on sex differences in human child development is a persistent topic of discussion and debate. Given their many similarities to humans, chimpanzees are an important study species for understanding the biological and evolutionary roots of sex differences in human development. In this study, we present the most detailed analyses of wild chimpanzee infant development to date, encompassing data from 40 infants from the long-term study of chimpanzees at Gombe National Park, Tanzania. Our goal was to characterize age-related changes, from birth to five years of age, in the percent of observation time spent performing behaviors that represent important benchmarks in nutritional, motor, and social development, and to determine whether and in which behaviors sex differences occur. Sex differences were found for indicators of social behavior, motor development and spatial independence with males being more physically precocious and peaking in play earlier than females. These results demonstrate early sex differentiation that may reflect adult reproductive strategies. Our findings also resemble those found in humans, which suggests that biologically-based sex differences may have been present in the common ancestor and operated independently from the influences of modern sex-biased parental behavior and gender socialization.

Validation of a field technique and characterization of fecal glucocorticoid metabolite analysis in wild chimpanzees (Pan troglodytes).

Monitoring adrenocortical activity in wild primate populations is critical, given the well‐documented relationship between stress, health, and reproduction. Although many primate studies have quantified fecal glucocorticoid metabolite (FGM) concentrations, it is imperative that researchers validate their method for each species. Here, we describe and validate a technique for field extraction and storage of FGMs in wild chimpanzees (Pan troglodytes). Our method circumvents many of the logistical challenges associated with field studies while yielding similar results to a commonly used laboratory method. We further validate that our method accurately reflects stress physiology using an adrenocorticotropic hormone challenge in a captive chimpanzee and an FGM peak at parturition in a wild subject. Finally, we quantify circadian patterns for FGMs for the first time in this species. Understanding these patterns may allow researchers to directly link specific events with the stress response. Am. J. Primatol. 75:57‐64, 2013.

Early social exposure in wild chimpanzees: mothers with sons are more gregarious than mothers with daughters.

Significance Early social experience is critical to developing appropriate adult behavior. Mothers are particularly important social partners but may also facilitate or restrict access to others. Studies investigating human parental facilitation are limited and confounded by culture. Comparative studies therefore provide invaluable insight into how parents provide social opportunities for offspring. We investigated maternal subgrouping patterns by infant sex in one of our closest relatives, chimpanzees, and found that mothers with sons were more gregarious. Infants themselves may influence patterns later in infancy, but differences were apparent early in life, when mothers and infants are in almost constant contact. Furthermore, mothers with sons associated more with adult males, potential behavioral models, early in life. These differences foreshadow sex-typical adult social behavior. In many mammals, early social experience is critical to developing species-appropriate adult behaviors. Although mother–infant interactions play an undeniably significant role in social development, other individuals in the social milieu may also influence infant outcomes. Additionally, the social skills necessary for adult success may differ between the sexes. In chimpanzees (Pan troglodytes), adult males are more gregarious than females and rely on a suite of competitive and cooperative relationships to obtain access to females. In fission–fusion species, including humans and chimpanzees, subgroup composition is labile and individuals can vary the number of individuals with whom they associate. Thus, mothers in these species have a variety of social options. In this study, we investigated whether wild chimpanzee maternal subgrouping patterns differed based on infant sex. Our results show that mothers of sons were more gregarious than mothers of daughters; differences were especially pronounced during the first 6 mo of life, when infant behavior is unlikely to influence maternal subgrouping. Furthermore, mothers with sons spent significantly more time in parties containing males during the first 6 mo. These early differences foreshadow the well-documented sex differences in adult social behavior, and maternal gregariousness may provide sons with important observational learning experiences and social exposure early in life. The presence of these patterns in chimpanzees raises questions concerning the evolutionary history of differential social exposure and its role in shaping sex-typical behavior in humans.