In most social ungulate species, males are larger than females and the sexes live in separate groups outside the breeding season. It is important for our understanding of the evolution of sociality to find out why sexual segregation is so widespread not only in ungulates but also in other mammals. Sexual body size dimorphism was proposed as a central factor in the evolution of sexual segregation in ungulates. We tested three hypotheses put forward to explain sexual segregation : the predation-risk, the forage-selection, and the activity budget hypothesis. We included in our analyses ungulate species ranging from non-dimorphic to extremely dimorphic in body size. We observed oryx, zebra, bighorn sheep and ibex in the field and relied on literature data for 31 additional species. The predation-risk hypothesis predicts that females will use relatively predator-safe habitats, while males are predicted to use habitats with higher predation risk but better food quality. Out of 24 studies on different species of ungulates, females and their offspring chose poorer quality but safer habitat in only eight cases. The forage-selection hypothesis predicts that females would select habitat based on food quality, while males should prefer high forage biomass. In fact, females selected higher quality food in only six out of 18 studies where males and females segregated, in eight studies there was no difference in forage quality and in four studies males were in better quality habitat. The activity budget hypothesis predicts that with increasing dimorphism in body size males and females will increasingly differ in the time spent in different activities. Differences in activity budgets would make it difficult for males and females to stay in mixed-sex groups due to increased costs of synchrony to maintain group cohesion. The predictions of the activity budget hypothesis were confirmed in most cases (22 out of 23 studies). The heavier males were compared to females, the more time females spent foraging compared to males. The bigger the dimorphism in body mass, the more males spent time walking compared to females. Lactating females spent more time foraging than did non-lactating females or males. Whether species were mainly bulk or intermediate feeders did not affect sexual differences in time spent foraging. We conclude that sexual differences in activity budgets are most likely driving sexual segregation and that sexual differences in predation risk or forage selection are additive factors.

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In many mammals, males and females live solitarily or in separate groups outside the breeding season. Sexual segregation is widespread in ungulates, but also occurs in whales, seals, monkeys, macropods, elephants, sh and bird species. What causes segregation by sex is still poorly understood, despite intense research done mainly on different ungulate species. In most species studied, males were clearly larger than females. The evolution of sexual dimorphism in body size has largely been attributed to sexual selection and mating strategies. While the consequences of body-size differences on energy requirements and metabolic rates received most attention, studies on consequences of sexual body-size differences on behavior are lacking. This review emphasizes the importance to study a wide range of social mammals with a different or no degree of sexual segregation and sexual dimorphism in body size; something which has not been done and has greatly limited our ability to test alternative hypotheses. More emphasis has to be put on the study of activity budgets, sociality and habitat choice of non-dimorphic species to explain the evolution of permanent territoriality, of long-term bonds of male-female pairs and of the occurrence of mixed-sex groups (adults), as well as its absence in dimorphic species. We review ve hypotheses proposed to explain sexual segregation, discuss alternative outcomes and predictions for each hypothesis, suggest alternative explanations for the evolution of sexual segregation and mating systems, and indicate new and important directions for research. We conclude that a phylogenetic comparison of behavior of a wide range of ungulates and other mammals will be needed to solve the enigma of sexual segregation.

SUMMARYMany studies have identified various host behavioural and ecological traits that are associated with parasite infection, including host gregariousness. By use of meta-analyses, we investigated to what degree parasite prevalence, intensity and species richness are correlated with group size in gregarious species. We predicted that larger groups would have more parasites and higher parasite species richness. We analysed a total of 70 correlations on parasite prevalence, intensity and species richness across different host group sizes. Parasite intensity and prevalence both increased positively with group size, as expected. No significant relationships were found between host group size and parasite species richness, suggesting that larger groups do not harbour more rare or novel parasite species than smaller groups. We further predicted that the mobility of the host (mobile, sedentary) and the mode of parasite transmission (direct, indirect, mobile) would be important predictors of the effects of group sizes on parasite infection. It was found that group size was positively correlated with the prevalence and intensity of directly and indirectly transmitted parasites. However, a negative relationship was observed between group size and mobile parasite intensity, with larger groups having lower parasite intensities. Further, intensities of parasites did not increase with group size of mobile hosts, suggesting that host mobility may negate parasite infection risk. The implications for the evolution and maintenance of sociality in host species are discussed, and future research directions are highlighted.

Sexual segregation is very common in vertebrates that live in groups. In this article, I will review proximate and ultimate causes of sexual segregation in social species and in particular in ungulates in which the bulk of research on the topic has been carried out. In most social ungulate species, males and females live in separate groups outside the breeding season, sometimes using different home ranges and types of habitat. In most of these species, males are larger than females. Dimorphism in body size can lead to sexual differences in ecology and behavior making it difficult for the two sexes to stay in the same group. It is important for our better understanding of the evolution of sociality, sexual dimorphism and different mating systems to determine why sexual segregation is so widespread not only in ungulates but also in other vertebrates. In this article, I discuss the ecology of the two sexes by reviewing proximate and ultimate causes of sexual segregation. To do this, I compare a range of studies of ruminants and include explanations for social segregation as well as for habitat segregation by gender. This leads into a review and updates current knowledge of the phenomenon. Although I present a number of different hypotheses, I focus in particular on predation risk, forage selection and activity budget and discuss the social-factors hypothesis. I stress that the key in solving the enigma of sexual segregation lies in clearly separating hypotheses that try to explain social segregation and habitat segregation, as well as in including experiments or model systems. To that end, I present a preliminary study on a test of the activity-budget hypothesis in three-spine sticklebacks and explain why I believe that shoaling fish are useful for analysing the underlying processes and mechanisms that lead to sexual segregation in animals. Lastly, I argue that it is unlikely that a single factor can explain social segregation or habitat segregation but that a model integrating different factors and different levels of segregation might succeed in describing proximate and ultimate causes of sexual segregation.