Female emancipation in a male dominant, sexually dimorphic primate under natural conditions

In most group-living animals, a dominance hierarchy reduces the costs of competition for limited resources. Dominance ranks may reflect prior attributes, such as body size, related to fighting ability or reflect the history of self-reinforcing effects of winning and losing a conflict (the winner-loser effect), or both. As to prior attributes, in sexually dimorphic species, where males are larger than females, males are assumed to be dominant over females. As to the winner-loser effect, the computational model DomWorld has shown that despite the female’s lower initial fighting ability, females achieve some degree of dominance of females over males. In the model, this degree of female dominance increases with the proportion of males in a group. This increase was supposed to emerge from the higher fraction of fights of males among themselves. These correlations were confirmed in despotic macaques, vervet monkeys, and in humans. Here, we first investigate this hypothesis in DomWorld and next in long-term data of 9,300 observation hours on six wild groups of robust capuchin monkeys (Sapajus libidinosus; S. nigritus, and S. xanthosternos) in three Brazilian sites. We test whether both the proportion of males and degree of female dominance over males are indeed associated with a higher relative frequency of aggression among males and a higher relative frequency of aggression of females to males. We confirm these correlations in DomWorld. Next, we confirm in empirical data of capuchin monkeys that with the proportion of males in the group there is indeed an increase in female dominance over males, and in the relative frequency of both male-male aggression and aggression of females to males and that the female dominance index is significantly positively associated with male male aggression. Our results reveal that adult sex ratio influences the power relation between the sexes beyond predictions from socioecological models.</p

Dispersal and social structure in black capuchin monkeys (Sapajus nigritus) of Carlos Botelho State Park, São Paulo

Padrões de assimetria sexual na dispersão e relações de parentesco são fatores intimamente relacionados e considerados fundamentais para a compreensão da estrutura social dos primatas. Apesar da relevância desses dois assuntos, pouco se sabe sobre como eles afetam o comportamento individual e a estrutura social nos primatas neotropicais. Assim, essa pesquisa teve como objetivos: determinar o padrão de dispersão de uma população selvagem de macacosprego (Sapajus nigritus) por meio de análises genéticas e examinar o efeito do parentesco sobre a estrutura social dos grupos de S. nigritus. Esta pesquisa foi realizada no Parque Estadual Carlos Botelho, localizado no município de São Miguel Arcanjo/SP. Todo material genético foi obtido através de amostras fecais dos indivíduos adultos e subadultos de três grupos sociais. Após a extração do DNA, parte dele foi amplificada através da técnica da Reação em Cadeia da Polimerase. Para as análises genéticas utilizamos marcadores moleculares do tipo microssatélites. Os métodos de amostragem por varredura instantânea e ad libitum foram utilizados para o registro do comportamento dos indivíduos adultos de dois grupos. Os dados genéticos obtidos nesta tese indicaram que ambos os sexos dispersam, pois: 1) o grau de parentesco intragupo entre machos não diferiu do grau de parentesco entre fêmeas, 2) não houve diferenciação genética ao analisar a distribuição das frêquencias alélicas de machos e de fêmeas, e 3) não houve diferença entre machos e fêmeas quanto à probabilidade de terem nascido dentro dos grupos sociais nos quais foram amostrados. Os resultados de associações espaciais e interações sociais indicaram que as fêmeas adultas não formam fortes relações sociais entre si, sendo pouco afiliativas. Os machos adultos estabelecem fracas relações sociais, que podem ser classificadas como tolerantes e caracterizadas pela ausência de catação e pela baixa frequência de interações agonísticas. As relações sociais mais fortes observadas dentro de cada grupo social foram entre machos e fêmeas. Além de machos e fêmeas associarem-se espacialmente e catarem-se mais do que as díades compostas somente por fêmeas ou por machos, eles também interagiram de maneira agonística numa menor frequência. As díades consideradas como aparentadas não demonstraram ser mais afiliativas ou manter maior proximidadade espacial quando comparadas com as díades não aparenteadas. O baixo grau de parentesco/familiaridade parece ser um fator importante como uma explicação geral para as fracas relações sociais de fêmeas no PECB. No entanto, em termos individuais, aparentemente, esse fator exerceu pouca influência sobre as relações sociais entre as fêmeas e entre os machos. Os resultados apresentados nesta tese ampliam o conhecimento sobre relações sociais em primatas neotropicais e os possíveis fatores que as afetam. O padrão de dispersão e a estrutura social são elementos do sistema social de primatas neotropicais altamente flexíveis, variando entre populações da mesma espécie ou mesmo entre grupos de uma mesma população. Além disso, a alegada assimetria sexual na dispersão talvez não seja passível de generalização entre as populações de Sapajus, e talvez dispersão de ambos os sexos nesse gênero possa ser mais comum do que previamente se consideravaSex-biased dispersal patterns and kinship are related factors extremely important to understand the social structure of primates. In spite of their importance, little is known about how these factors affect individual behavior and the social structure of neotropical primates. Therefore, the aims of this research were: to determine the dispersal pattern of a wild population of black capuchin monkeys (Sapajus nigritus) through genetic analyses, and to verify the effect of kinship on the social structure of groups of S. nigritus. This research was conducted at Carlos Botelho State Park, in the municipality of São Miguel Arcanjo/SP. We used DNA from fecal samples of adult and subadult members from three wild social groups. DNA was amplified by Polymerase Chain Reaction and microsatellite molecular markers were used. Behavioral data were collected systematically for two groups, and we used scan sampling and ad libitum methods to record the behavior of adults. Genetic data indicated that both sexes disperse from their natal groups, since: 1) relatedness between males was not statistically different from relatedness between females, 2) there was no difference between males and females in population genetic differentiation, and 3) there was no difference between males and females in the probability of being born in the group from which they were sampled. The results of the spatial association and social interactions analysis indicate that females are little affiliative and do not form strong relationships among themselves. Adult males established weak relationships, being classified as tolerant, and characterized by no grooming interactions and by low frequency of agonism. The strongest social relationships observed within each group were between males and females. They associated and groomed more than female dyads and male dyads. Moreover, there were fewer agonistic interactions between males and females as compared to same sex dyads. Related and unrelated dyads showed similar rates of association and affiliative behavior. The low level of relatedness/familiarity might be an important factor contributing to the weak social relationship among females in the PECB. However, at the individual level, kinship had low influence on male and female social relationships. The results presented here extended our knowledge about social relationships of neotropical primates and about the factors that influence the relationships. The dispersal pattern and the social structure are flexible elements of social systems, and can vary among populations of the same species or among groups of the same population. In addition, male-biased dispersal is not a general characteristic for all populations of Sapajus, and dispersal by both sexes might be more commom than previously though

Social behaviour of adult male northern muriquis (Brachyteles hypoxanthus) in heterosexual and all male group, at the Estação Biológica de Caratinga, MG

Muriquis-do-norte (Brachyteles hypoxanthus) são caracterizados pela filopatria entre os machos, a formação de grupos multi-macho/multi-fêmea e um sistema de acasalamento poligâmico, com alto grau de promiscuidade. No entanto, alterações ambientais (ecológicas ou demográficas) podem conduzir a modificações no sistema social (seja na estrutura social, organização social ou sistema de acasalamento) considerado como o típico da espécie. A formação de um grupo composto somente por machos (denominado UM), que se associa constantemente com um grupo misto, ou seja, multi-macho/multi-fêmea (denominado grupo Nadir), e a transferência de machos entre grupos distintos, são padrões comportamentais não descritos para os muriquis-do-norte da Estação Biológica de Caratinga, MG. Assim, esta pesquisa teve como objetivos: 1) analisar as interações sociais entre os machos da UM com os machos adultos do grupo Nadir, 2) comparar o comportamento dos machos UM com o comportamento dos machos adultos do grupo Nadir, e 3) analisar as relações sociais entre os machos adultos do grupo Nadir. Dezessete machos adultos foram alvo desta pesquisa, 8 indivíduos da UM e 9 do grupo Nadir. Os machos adultos do grupo Nadir foram divididos em: imigrantes (machos que se transferiram da UM para o grupo Nadir, 4 indivíduos), e residentes (machos fundaram o grupo Nadir, 5 indivíduos). Interações afiliativas, dinâmica espacial e vizinhos em diferentes categorias de distância espacial foram registrados pelo método animal focal. O método de todas as ocorrências foi empregado para o registro de cópulas e interações agonísticas. Os resultados demonstraram que apesar dos machos UM se associarem com o grupo Nadir, sendo constantemente observados na periferia ou até mesmo dentro do grupo, as interações sociais entre os machos UM e GN não se caracterizaram como relações sociais de indivíduos pertencentes ao mesmo grupo. Entretanto, as interações sociais dos machos UM com os machos GN não apresentaram um padrão homogêneo, sendo as interações com os imigrantes diferenciadas das interações com os residentes. Ao comparar o comportamento dos machos UM e GN, houve diferença significativa somente no padrão de associação espacial. A análise dos dados referente às relações sociais dos machos GN revelou uma assimetria, pois os imigrantes formaram associações mais fortes com os residentes do que entre si e foram os responsáveis pela manutenção de proximidade e interações afiliativas com os residentes. A tolerância entre os machos UM e machos GN, principalmente dos imigrantes, foi explicada pela familiaridade entre eles. Em relação às relações sociais entre os machos GN, inferi um status social diferenciado entre imigrantes e residentes, devido à maior atratividade que os residentes exerceram sobre os imigrantes. A capacidade de se adaptar a condições demográficas específicas, variando sua organização e estrutura social, evidencia a flexibilidade comportamental da espécie.Northern muriquis (Brachyteles hypoxanthus) are characterized by male philopatry, by multi-male/multi-female groups and by a polygamous mating system, with high levels of promiscuity. Nevertheless, environmental changes (ecological or demographic) can lead to changes in the social system (in the social structure, as well as in the social organization, or in the mating system) considered as typical of the species. Behavioural patterns such as the formation of an all male group (called UM), constantly associated with a multi-male/multi-female group (called Nadir group), and the transfer of males between distinct groups, are not described yet for northern muriquis of the Estação Biológica de Caratinga, MG. Therefore, the aims of this research were: 1) to analyse the social interactions between the UM males and the adult males of the Nadir group, 2) to compare the behaviour of UM males and the behaviour of adult males of the Nadir group, and 3) to analyse the social relationships among the adult males of the Nadir group. Seventeen adult males were the subjects of this research, 8 individuals belonged to the UM and 9 belonged to the Nadir group. The adult males of Nadir group were separeted into: immigrant (males that transfer from UM to Nadir group, 4 individuals), and resident (males that founded the Nadir group, 5 individuals). Afilliative interactions, spacial dynamics and neighbours at different categories of spacial distances were registered by focal animal sampling. The all occurrences sampling was used to record episodes of copulation and agonistic interactions. The results indicated that the social interactions between UM e GN males did not characterize social relationships between members of the same group, even though the UM males associated with the Nadir group, being often observed at the border or even inside the group. The interactions between males of the two groups did not obey a unique pattern. The interactions with immigrant males were different from those with resident males. The comparison between the UM and GN male behaviour revealed significant differences only in the pattern of spatial association. The analysis of GN males social relationships revealed an asymmetry, because the immigrants had stronger associations with residents than among themselves and were the responsible for approaching and for mantaining afilliative interactions with residents. GN males, particularly immigrants, were tolerant towards UM males, probably due to the familiarity among them. Regarding the social relationships among GN males, it was possible to distinguish the social status between immigrants and residents, since resident males were highly attractive to the immigrant males. The adaptive capacity to adjust to specific demographyc conditions, trough changes in the social organization and structure, highlights the behavioural flexibility of the species

Social cliques in male northern muriquis Brachyteles hypoxanthus

Analyses of spatial relationships and social interactions provide insights into the social structure of animal societies and the ways in which social preferences among and between dyads affect higher order social relationships. In this paper we describe the patterns of spatial associations and social interactions among adult male northern muriquis in order to evaluate the dynamics of their social networks above the dyadic levels. Systematic observations were made on the 17 adult males present in a multi-male/multi-female group from April 2004 through February 2005, and in July 2005. Analyses of their spatial relationships identified two distinct male cliques; some adult males (called “N” males) were more connected to the females and immatures than other adult males (“MU” males), which were more connected to one another. Affiliative interactions were significantly higher among dyads belonging to the same clique than to different cliques. Although frequencies of dyadic agonistic interactions were similarly low among individuals within and between cliques, MU males appeared to be subordinate to N males. Nonetheless, there were no significant differences in the copulation rates estimated for MU males and N males. Mutual benefits of cooperation between MU and N cliques in intergroup encounters might explain their ongoing associations in the same mixed-sex group [Current Zoology 58 (2): 342-352, 2012]

Flexible and conservative features of social systems in tufted capuchin monkeys: comparing the socioecology of Sapajus libidinosus and Sapajus nigritus

Socioecological models assume that primates adapt their social behavior to ecological conditions, and predict that food availability and distribution, predation risk and risk of infanticide by males affect patterns of social organization, social structure and mating system of primates. However, adaptability and variation of social behavior may be constrained by conservative adaptations and by phylogenetic inertia. The comparative study of closely related species can help to identify the relative contribution of ecological and of genetic determinants to primate social systems. We compared ecological features and social behavior of two species of the genus Sapajus, S. nigritus in Carlos Botelho State Park, an area of Atlantic Forest in Sao Paulo state, and S. libidinosus in Fazenda Boa Vista, a semi-arid habitat in Piaui state, Brazil. S. libidinosus perceived higher predation risk and fed on clumped, high quality, and usurpable resources (fruits) all year round, whereas S. nigritus perceived lower predation risk and relied on evenly distributed, low-quality food sources (leaves) during periods of fruit shortage. As predicted by socioecology models, S. libidinosus females were philopatric and established linear and stable dominance hierarchies, coalitions, and grooming relationships. S. nigritus females competed less often, and could transfer between groups, which might explain the lack of coalitions and grooming bonds among them. Both populations presented similar group size and composition and the same polygynous mating system. The species differed from each other in accordance with differences in the characteristics of their main food sources, as predicted by socioecological models, suggesting that phylogenetic inertia does not constrain social relationships established among female Sapajus. The similarity in mating systems indicates that this element of the social system is not affected by ecological variables and thus, is a more conservative behavioral feature of the genus Sapajus. Am. J. Primatol. 74:315331, 2012. (c) 2011 Wiley Periodicals, Inc.FAPESPFAPESP [06/51577-2, 06/56059-0, 08/55684-3]CNPq [420038/2005-1]CNPqNational Geographic SocietyNational Geographic SocietyLeakey FoundationLeakey FoundationIBAMAIBAMACNPqCNP