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• W2912459920 abstract "What are multilevel societies? Multilevel societies feature different structural orders of grouping. They are best known from primates, but also from some other mammalian taxa. While sub-structuring or cliquishness typifies many mammal societies, the subunits in multilevel societies are generally socially cohesive and in some cases can be clearly spatially delineated. The core entities are usually polygynous reproductive units which are nested within a larger social matrix. The low-level units stay in regular or permanent proximity and coordinate their day-to-day activities. Social interactions are more frequent within the inner layers of a multilevel society and become increasingly ephemeral toward the outer layers. In multilevel societies, the higher levels can number several hundred individuals. Why are they important? Multilevel societies are among the most complex social systems in nature, but their origins and function are still poorly understood. Studying them is of fundamental importance for a holistic understanding of the evolution of sociality. Furthermore, the vast majority of human societies show precisely such a multi-tiered system, and some primates can serve as models to reconstruct their origins. In hunter-gatherer societies, for example, families are part of local bands of around 25 individuals which are relatively flexible in terms of size and composition, and higher-level tribes are formed by several bands that fuse together. Where do multilevel societies occur? Multilevel societies are found in some primate taxa (Figure 1), such as papionin monkeys (gelada, Hamadryas baboon, Guinea baboon), Asian colobine monkeys (snub-nosed monkeys, proboscis monkey and douc langurs) and humans. Multilevel societies may also exist in lesser known and more elusive species such as uacaris, drills and Angolan colobus monkeys, but detailed data on their social organization are not yet available. Prominent examples of non-primate mammals with multilevel societies are elephants, zebras, giraffes and sperm whales. How are multilevel societies structured? The complexity of multilevel societies is probably best epitomized by Hamadryas baboons. At the core of their multilevel societies are ‘one-male units’, semi-permanent associations between a male and one or several females and their offspring. They often also contain one or more follower males who are affiliated with the females but do not sire offspring. A collection of one-male units constitutes a ‘clan’, which also includes solitary males. Several clans form bands, which aggregate into larger troops. While the band is a genuine social group, troops are the result of several bands coming together at sleeping cliffs. Other multilevel societies show variation of this structure. Despite similarities in multilevel societies in different species, the underlying social dynamics can be fundamentally different. Geladas and Hamadryas, for example, differ significantly in the patterning of social interactions, levels of coercion and sex biases in dispersal. For instance, in geladas, units are composed of closely related females, whereas in Hamadryas there are stronger bonds among male relatives in clans. Are multilevel societies the same as fission–fusion societies? The term ‘multilevel’ is sometimes used interchangeably with fission–fusion, but the latter refers to fluid grouping patterns and is not tied to a particular social organization. Fission and fusion can be found in both multilevel societies, such as those of sperm whales and Hamadryas baboons and multimale–multifemale groups as seen in chimpanzees; in the former, fission–fusion occurs along defined societal seams such as those between one-male units, whereas in the latter it generates subunits that can change in size and composition. Why are multilevel societies not more common? Given the sheer size of multilevel societies they are expected to become manifest only when the costs of grouping are attenuated. When resources occur in sufficiently large patches and are sufficiently abundant, an increase in group size does not result in a drop in food intake and more competition. Interestingly, multilevel societies are particularly common among species with a reliable resource (grass in geladas and plains zebras, lichen in snub-nosed monkeys). How did multilevel societies evolve? In the case of primates, different evolutionary pathways giving rise to multilevel societies have been identified. In Hamadryas baboons, phylogenetic reconstructions point to ancestrally mixed-sex groups becoming divided into modules (one-male units). An increase in aggregation size due to localized resources is thought to have coincided with an increasing risk of harassment by unfamiliar individuals, which forced females into small stable one-male units for better protection. In colobines, amalgamation of ancestrally separate units into bands appears more likely. Social pressures such as persistent threat from bachelor males that encroach on one-male units could have created incentives for banding. What benefits do individuals derive from multilevel societies? Different levels fulfil different adaptive functions and have evolved in response to different cost–benefit trade-offs. In humans, reproduction and close social support are the domains of the lowest-level tier, mid-level tiers are a pool from which one can choose partners for cooperative breeding and cooperative hunting while cooperative defence is best served by high-level tiers. In essence, higher levels of sociality can tackle problems that core structures cannot. Lactating zebras benefit from associating with other groups of lactating females to lower predation risk. Multilevel species can thus simultaneously reap the benefits of various grouping levels and thereby avoid the optimization problem that a one-size-fits-all group may face. What are the consequences of living in multilevel societies? Primates living in multilevel societies are characterized by pronounced sexual size dimorphism. Being large may increase a male’s mating success when competition over access to females is frequent and intense. Primate males in multilevel societies also boast extravagant secondary sexual traits, such as the elongated nose of proboscis monkeys. In a crowded and anonymous social environment wherein traditional means of getting to know each other may not work, selection must have placed a premium on such amplified signals of individual identity, rank or attractiveness. The cognitive consequences of living in multilevel societies are largely unexplored. One would expect individuals to possess significant cognitive skill to navigate such a complex social landscape. One could also hypothesize that complex social organization translates into the need for more intricate social knowledge." @default.
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• W2912459920 date "2017-09-01" @default.
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• W2912459920 title "Multilevel societies" @default.
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