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Barplot of the mean number of (a) male bystanders and (b) higher-ranking female bystanders during non-copulation and female-initiated copulation scans. Barplots are drawn from the raw individual means. Error bars indicate the standard error of the mean. ‘*’: p < 0.05, ‘n.s.’: p > 0.05.

Barplot of the mean number of (a) male bystanders and (b) higher-ranking female bystanders during non-copulation and female-initiated copulation scans. Barplots are drawn from the raw individual means. Error bars indicate the standard error of the mean. ‘*’: p < 0.05, ‘n.s.’: p > 0.05.

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Article
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In social species, female mating strategies can be constrained by both male and female groupmates through sexual conflict and reproductive competition, respectively. This study tests if females adjust their sexual behaviour according to the presence of male and female bystanders in wild chacma baboons (Papio ursinus) and assesses their relative imp...

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... Whereas chacma baboon and macaque males, to some extent, are also less likely to initiate mating when additional (higher-ranking) males are in the vicinity(Overduin-de Vries et al., 2012;Overduin-de Vries et al., 2013), orang-utan males initiate copulations despite the presence of dominant males. In more social (group-living) species sanctions for sneaky mating may be high for both males and females: punishment of the female (sensuClutton-Brock & Parker, 1995;Baniel et al., 2019) and direct male aggression towards sneakily mating males(Overduin-de Vries et al., 2012) ...
Article
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Sexual coercion is widespread in the animal kingdom. Its direct forms, including harassment and forced copulation, have largely been investigated as an expression of (alternative) male reproductive strategies, rather than the result of a sexual conflict between the sexes. Likewise, the frequent occurrence of forced copulations in orang-utans (Pongo spp.) has been attributed to male strategies and more recently also to concealed female fecundity. So far, however, the immediate contextual variables leading to forced copulations have rarely been examined. We compared two orang-utan populations, Suaq (Pongo abelii, Sumatra) and Tuanan (Pongo pygmaeus, Borneo), both characterized by an individual-based fission–fusion lifestyle, whereas their socioecology differs. We assessed how the occurrence of female-resisted and voluntary copulations was affected by female reproductive state, male morph (unflanged or flanged), measures of male–male competition, male–female relationship and ecological factors. Besides female reproductive state and male morph, predictors of female resistance were related to male–male competition. First, female resistance was more likely towards subordinate males who were displaced from proximity to the female by another male during that association. Second, the presence of additional flanged males increased the probability of female resistance. Third, the latency to both the arrival of another male and to the end of the association after sexual interactions was shorter if there was female resistance. We conclude that sexual coercion in orang-utans is highly dependent on the vicinity of more dominant males and can only be understood in the light of sexual conflict: While males force copulations when at risk of losing access to a female and thus follow a ‘now-or-never’ strategy, female resistance follows a ‘not-you-now’ pattern, which is ultimately consistent with an infanticide avoidance strategy.
... Audience effects are usually tested by comparing control scan conditions when the targeted behavior is not expressed with conditions when the targeted behaviors are expressed (e.g., in Baniel et al. 2019). In this study, I estimated that the effect of motivational state might be too important, and this traditional approach may have been irrelevant. ...
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Animals exhibit an astonishing diversity of communicative systems, with substantial variation in both the nature and the number of signals they produce. Variation in communicative complexity has been conceptually and empirically attributed to social complexity and formalized as the “social-complexity hypothesis for communicative complexity” (SCHCC). Indeed, group-living animals face complex social environments where they engage in a wide range of interactions with different social partners triggering the need for transmission of a broader diversity of messages. In chapter I (Peckre et al. 2019), I review the literature on the current tests of the SCHCC, pointing out and discussing what I identified as the main gaps in the current state of the art. Specifically, three key issues emerged from my analysis. The first issue concerns the operational definition of the main variables, social and communicative complexity. Notably, when defining communicative complexity, most empirical tests of the SCHCC focus on a single modality (e.g., acoustic, visual, olfactory) whereas several good reasons exist for acknowledging the multimodal nature of both, signals and communicative systems in this framework. At the system level, focusing on only one modality may lead to over- or underestimation of the relationship between social and communicative complexity. The second issue relates to the fact that while numerous studies have highlighted a link between social and communicative complexity, their correlative nature does not permit conclusions about the direction of causality. Indeed, alternative hypotheses involving anatomical, phylogenetical, or ecological factors have also been proposed to explain the evolution of more complex forms of communication. Finally, I note that researchers rarely address the actual ways in which social factors directly affect variation in signaling. Indeed, the underlying mechanisms of this link are usually left unexplored, failing to uncover the specific attribute of communication that would be co-evolving with specific aspects of sociality. I, therefore, make a plea for expanding tests of the SCHCC in 1) scope (systematic approach across modalities) and 2) depth (characterization of the observed relationships) as I believe it may significantly advance our understanding of the intricate links between animal sociality and communication. To address point 1), I offer in chapter II a comprehensive approach of the cross-modal communicative systems of two closely related true lemur species having similar morphology, living in similar habitats, but differing in their social systems. I studied wild Eulemur rufifrons and E. mongoz in Madagascar, respectively in Kirindy and Ankatsabe forests for 12 months. I describe a new analytical framework to assess the complexity of signaling systems across modalities. Applying a multimodal approach may help to uncover the different selective pressures acting on the communicative system and to understand better adaptive functions that might be unclear from the study of its separate components independently. E. rufifrons, the species having the more complex social system, also had overall a more complex communicative system than the one of the E. mongoz. Both careful choices of the species to compare to limit the effect of possible additional selective pressures and exploration of the social function of the non-homologous signals allow concluding that this increased complexity of the communicative system in E. rufifrons is most likely associated with social selective pressures. I developed this new analytical framework, partly based on using a cross-modal network approach, with the perspective of facilitating cross-taxonomic comparisons. Moreover, this approach may be combined with new multi-dimensional approaches of social complexity and contribute to a more holistic approach to the tests of the SCHCC. By this, we should be able to derive new testable hypotheses that would contribute to better understand the course of events that have led to the evolution of communication diversity in its distinct dimensions. In chapter III, I address point 2) by investigating the impacts of sociality on the expression of a multimodal signal, the anogenital scent-marking behavior in wild red-fronted lemurs. I specifically investigated intragroup audience effects on anogenital scent-marking behaviors in a wild population of red-fronted lemurs and particularly whether males and females differed in this aspect and if these differences may reveal functional differences associated with anogenital scent-marking across sexes. I found an intragroup audience effect in males but not in females. Males deposited less often anogenital marks when more males were present within a three meters range compared to five- or ten-meters ranges. Males may prefer to reduce the risk of physical contact by avoiding to scent-mark near other males, and/or give priority to other males to scent-mark. With these results, I provide important insights into the functional significance of anogenital scent-marking in red-fronted lemurs and support the idea of greater intragroup social pressures associated with anogenital scent-marking in males than in females in egalitarian species. Studying the flexibility of complex signal usage (e.g., occurrence or structural modifications) across social contexts (audiences) should permit the identification of different individual social characteristics that may elicit or constrain complex signal expression. These social characteristics may later constitute social pressures acting for or against the evolution of these complex signaling behaviors. In chapters IV and V, I also address ethical questions related to this project and the way I tried to adapt and best address my responsibilities for animal welfare. In chapter IV, I expose some technical details and ethical concerns experienced during the choice of my field sites. While in chapter V (Buil and Peckre et al. 2019) I present a remote releasable collar system developed in collaboration with the Neurobiology Laboratory (German Primate Center, Göttingen, Germany) intending to provide a tool to significantly reduce the number of captures in studies using bio-logging for medium-sized mammal species. Overall, by emphasizing the importance of the multimodal nature of communicative systems and the social context in which signals are exchanged, I hope to stimulate the development of new tests of the SCHCC based on this expanded framework. I additionally argue for the importance of looking across research fields since striking parallels may be observed between animal behavior and linguistic research when addressing the origins of communication complexity, be it in the form of human language or animal signaling.
Article
For mammalian females, which bear the energetic costs of gestation and lactation, competition over food resources is generally expected to have a more important impact on reproductive success than competition over mates. However, there are some situations in which mammalian females compete over access to mates, limited supplies of sperm or caretaking for their offspring. Competition over both access to food and access to mates have been documented in female baboons (Papio spp.). Here, we examined the relative importance of competition over food and mates for wild female olive baboons, Papio anubis, in shaping the patterns of aggression among females. Lactating and pregnant females were more aggressive than females in other reproductive states, and sexually receptive females initiated and received relatively little aggression. There was no evidence that females competed over caretakers for their offspring or reduced future competition through reproductive suppression. Our data suggest that competition over food plays a more important role than competition over mates for female olive baboons. The body of current evidence suggests that evolution has finely tuned female baboons’ responses to the competitive pressures that they face in their local environments.