Individuals with more or stronger social bonds experience enhanced survival and reproduction in various species, though the mechanisms mediating these effects are unclear. Social thermoregulation is a common behaviour across many species which reduces cold stress exposure, body heat loss, and homeostatic energy costs, allowing greater energetic investment in growth, reproduction, and survival, with larger aggregations providing greater benefits. If more social individuals form larger thermoregulation aggregations due to having more potential partners, this would provide a direct link between sociality and fitness. We conducted the first test of this hypothesis by studying social relationships and winter sleeping huddles in wild Barbary macaques (Macaca sylvanus), wherein individuals with more social partners experience greater probability of winter survival. Precipitation and low temperature increased huddle sizes, supporting previous research that huddle size influences thermoregulation and energetics. Huddling relationships were predicted by social (grooming) relationships. Individuals with more social partners therefore formed larger huddles, suggesting reduced energy expenditure and exposure to environmental stressors than less social individuals, potentially explaining how sociality affects survival in this population. This is the first evidence that social thermoregulation may be a direct proximate mechanism by which increased sociality enhances fitness, which may be widely applicable across taxa.
25The wounding or death of a conspecific has been shown to elicit varied behavioral responses
The Canidae are successful, being a widespread, abundant, speciose, and adaptable family. Several canids in particular have recently experienced rapid expansions in range and abundance, with similar situations mirrored on several continents by different species. Despite extreme behavioral diversity between and within species, monogamy is a common denominator in canid societies. In this review, we ask why canids are monogamous and how monogamy is related to their success. We begin with an overview of canid social monogamy, describing the pair bonding, paternal care, and often alloparental care that is characteristic of the family, and discuss theories on the evolution of mammalian social monogamy. We discuss why and how monogamy is maintained in canids, either voluntarily or enforced, and how ecological conditions influence either the functional advantages of monogamy or ability for enforcement and thus whether social monogamy is maintained. Social monogamy does not necessitate exclusive mating and many canids exhibit extra-pair paternity. We consider the costs and benefits of extra-pair mating for male and female canids and how ecological conditions can shift this cost/benefit balance and thus affect its prevalence. Monogamy may be responsible for many of the unusual canid reproductive characteristics through facilitating alloparental care and monogamy enforcement, and the domestic dogs' departure from monogamy supports our interpretation that it is an adaptation to resource availability. In asking whether monogamy is responsible, at least in part, for their success, we propose the monogamy as pro-cooperative hypothesis, suggesting four characteristics have contributed to canid success: (1) ecological flexibility, (2) high mobility, (3) high reproductive rates, and (4) sociality/cooperation, with the latter two being consequences of monogamy. These four interconnected traits enhance one another and it is their combination, with monogamy at its foundation enabling cooperative sociality and thereby enhanced reproduction and survival, that together comprise the formula of canid success.
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