Evolutionary determinants of modular societies in colobines

Modular societies are structurally characterized by nuclear one-male units (OMUs, or harems) embedded within larger relatively coherent social bands. Within the order Primates, modular societies are uncommon, found in only a few species, including humans. Asian colobines (Presbytini) principally form either unimale groups that forage independently and are often territorial, or modular associations, which range from tight bands composed of OMUs to loose neighborhoods of OMUs. A phylogenetic reconstruction of modularity in the Presbytini revealed that the single OMU pattern is probably the ancestral state while the modular pattern is derived. The selective forces favoring the evolution of modular societies have thus far been virtually unexplored. Although some ecological explanations cannot be ruled out at the moment due to lack of comparative and quantitative data, preliminary circumstantial evidence does not seem to support them. Instead, a social factor, bachelor threat, is consistent with many observations. This hypothesis argues that where the pressure from nonreproductive bachelor males is unusually high, OMUs aggregate as a means of decreasing the amount of harassment and the risk of takeovers and infanticide. A comparative test found an association between modular societies and bachelor threat, as proxied by sex ratio within social units. The concentration of modular systems in colobines may be due to their unusual ecology, which leads to unusually low intensity of scramble competition. Modular colobines rely more on nonlimiting ubiquitous resources than nonmodular ones and thus can afford to gather in bands. Moreover, by comparing the slopes of regressions between group size and daily travel distance for several groups of one modular and one nonmodular colobine, we found slopes in the nonmodular to be steeper by a factor 30, indicating that ecological constraints associated with scramble competition prevent higher level groupings in nonmodulars. Thus, modular sociality in Asian colobines may have arisen because both social benefits are substantial and ecological costs are relatively lo

Four levers of reciprocity across human societies: concepts, analysis and predictions

This paper surveys five human societal types – mobile foragers, horticulturalists, pre-state agriculturalists, state-based agriculturalists and liberal democracies – from the perspective of three core social problems faced by interacting individuals: coordination problems, social dilemmas and contest problems. We characterise the occurrence of these problems in the different societal types and enquire into the main force keeping societies together given the prevalence of these. To address this, we consider the social problems in light of the theory of repeated games, and delineate the role of intertemporal incentives in sustaining cooperative behaviour through the reciprocity principle. We analyse the population, economic and political structural features of the five societal types, and show that intertemporal incentives have been adapted to the changes in scope and scale of the core social problems as societies have grown in size. In all societies, reciprocity mechanisms appear to solve the social problems by enabling lifetime direct benefits to individuals for cooperation. Our analysis leads us to predict that as societies increase in complexity, they need more of the following four features to enable the scalability and adaptability of the reciprocity principle: nested grouping, decentralised enforcement and local information, centralised enforcement and coercive power, and formal rules

How Humans Evolved Large Brains: Comparative Evidence

The human brain is about three times as large as that of our closest living relatives, the great apes. Overall brain size is a good predictor of cognitive performance in a variety of tests in primates. 1,2 Therefore, hypotheses explaining the evolution of this remarkable difference have attracted much interest. In this review, we give an overview of the current evidence from comparative studies testing these hypotheses. If cognitive benefits are diverse and ubiquitous, it is possible that most of the variation in relative brain size among extant primates is explained by variation in the ability to avoid the fitness costs of increased brain size (allocation trade-offs and increased minimum energy needs). This is indeed what we find, suggesting that an energetic perspective helps to complement approaches to explain variation in brain size that postulate cognitive benefits. The expensive brain framework also provides a coherent scenario for how these factors may have shaped early hominin brain expansion

Sex Differences in Object Manipulation in Wild Immature Chimpanzees (Pan troglodytes schweinfurthii) and Bonobos (Pan paniscus): Preparation for Tool Use?

Sex differences in immatures predict behavioural differences in adulthood in many mammal species. Because most studies have focused on sex differences in social interactions, little is known about possible sex differences in 'preparation' for adult life with regards to tool use skills. We investigated sex and age differences in object manipulation in immature apes. Chimpanzees use a variety of tools across numerous contexts, whereas bonobos use few tools and none in foraging. In both species, a female bias in adult tool use has been reported. We studied object manipulation in immature chimpanzees at Kalinzu (Uganda) and bonobos at Wamba (Democratic Republic of Congo). We tested predictions of the 'preparation for tool use' hypothesis. We confirmed that chimpanzees showed higher rates and more diverse types of object manipulation than bonobos. Against expectation, male chimpanzees showed higher object manipulation rates than females, whereas in bonobos no sex difference was found. However, object manipulation by male chimpanzees was play-dominated, whereas manipulation types of female chimpanzees were more diverse (e.g., bite, break, carry). Manipulation by young immatures of both species was similarly dominated by play, but only in chimpanzees did it become more diverse with age. Moreover, in chimpanzees, object types became more tool-like (i.e., sticks) with age, further suggesting preparation for tool use in adulthood. The male bias in object manipulation in immature chimpanzees, along with the late onset of tool-like object manipulation, indicates that not all (early) object manipulation (i.e., object play) in immatures prepares for subsistence tool use. Instead, given the similarity with gender differences in human children, object play may also function in motor skill practice for male-specific behaviours (e.g., dominance displays). In conclusion, even though immature behaviours almost certainly reflect preparation for adult roles, more detailed future work is needed to disentangle possible functions of object manipulation during development

Cognitive differences between orang-utan species: a test of the cultural intelligence hypothesis

We also express our gratitude to the following foundations who have financially supported this project: A. H. Schultz Foundation, Paul Schiller Foundation and the Claraz Foundation in Switzerland and Waldemar von Frenckells Foundation, Ella and Georgh Ehrnrooths Foundation, Otto A. Malms Donationsfond, Nordenskiöld Samfundet and Oskar Öflunds Foundation in Finland.Cultural species can-or even prefer to-learn their skills from conspecifics. According to the cultural intelligence hypothesis, selection on underlying mechanisms not only improves this social learning ability but also the asocial (individual) learning ability. Thus, species with systematically richer opportunities to socially acquire knowledge and skills should over time evolve to become more intelligent. We experimentally compared the problem-solving ability of Sumatran orang-utans (Pongo abelii), which are sociable in the wild, with that of the closely related, but more solitary Bornean orang-utans (P. pygmaeus), under the homogeneous environmental conditions provided by zoos. Our results revealed that Sumatrans showed superior innate problem-solving skills to Borneans, and also showed greater inhibition and a more cautious and less rough exploration style. This pattern is consistent with the cultural intelligence hypothesis, which predicts that the more sociable of two sister species experienced stronger selection on cognitive mechanisms underlying learning.Publisher PDFPeer reviewe

Hibernation constrains brain size evolution in mammals

The expensive brain hypothesis predicts that the lowest stable level of steady energy input acts as a strong constraint on a species’ brain size, and thus, that periodic troughs in net energy intake should select for reduced brain size relative to body mass. Here, we test this prediction for the extreme case of hibernation. Hibernators drastically reduce food intake for up to several months and are therefore expected to have smaller relative brain sizes than nonhibernating species. Using a comparative phylogenetic approach on brain size estimates of 1104 mammalian species, and controlling for possible confounding variables, we indeed found that the presence of hibernation in mammals is correlated with decreased relative brain size. This result adds to recent comparative work across mammals and amphibians supporting the idea that environmental seasonality (where in extremis hibernation is necessary for survival) imposes an energetic challenge and thus acts as an evolutionary constraint on relative brain size

Four levers of reciprocity across human societies: concepts, analysis and predictions

This paper surveys five human societal types – mobile foragers, horticulturalists, pre-state agriculturalists, state-based agriculturalists and liberal democracies – from the perspective of three core social problems faced by interacting individuals: coordination problems, social dilemmas and contest problems. We characterise the occurrence of these problems in the different societal types and enquire into the main force keeping societies together given the prevalence of these. To address this, we consider the social problems in light of the theory of repeated games, and delineate the role of intertemporal incentives in sustaining cooperative behaviour through the reciprocity principle. We analyse the population, economic and political structural features of the five societal types, and show that intertemporal incentives have been adapted to the changes in scope and scale of the core social problems as societies have grown in size. In all societies, reciprocity mechanisms appear to solve the social problems by enabling lifetime direct benefits to individuals for cooperation. Our analysis leads us to predict that as societies increase in complexity, they need more of the following four features to enable the scalability and adaptability of the reciprocity principle: nested grouping, decentralised enforcement and local information, centralised enforcement and coercive power, and formal rules

How institutions shaped the last major evolutionary transition to large-scale human societies

What drove the transition from small-scale human societies centred on kinship and personal exchange, to large-scale societies comprising cooperation and division of labour among untold numbers of unrelated individuals? We propose that the unique human capacity to negotiate institutional rules that coordinate social actions was a key driver of this transition. By creating institutions, humans have been able to move from the default ‘Hobbesian’ rules of the ‘game of life’, determined by physical/environmental constraints, into self-created rules of social organization where cooperation can be individually advantageous even in large groups of unrelated individuals. Examples include rules of food sharing in hunter–gatherers, rules for the usage of irrigation systems in agriculturalists, property rights and systems for sharing reputation between mediaeval traders. Successful institutions create rules of interaction that are self-enforcing, providing direct benefits both to individuals that follow them, and to individuals that sanction rule breakers. Forming institutions requires shared intentionality, language and other cognitive abilities largely absent in other primates. We explain how cooperative breeding likely selected for these abilities early in the Homo lineage. This allowed anatomically modern humans to create institutions that transformed the self-reliance of our primate ancestors into the division of labour of large-scale human social organization

Cooperation in large‐scale human societies — What, if anything, makes it unique, and how did it evolve?

To resolve the major controversy about why prosocial behaviors persist in large-scale human societies, we propose that two questions need to be answered. First, how do social interactions in small-scale and large-scale societies differ? By reviewing the exchange and collective-action dilemmas in both small-scale and large-scale societies, we show they are not different. Second, are individual decision-making mechanisms driven by self-interest? We extract from the literature three types of individual decision-making mechanism, which differ in their social influence and sensitivity to self-interest, to conclude that humans interacting with non-relatives are largely driven by self-interest. We then ask: what was the key mechanism that allowed prosocial behaviors to continue as societies grew? We show the key role played by new social interaction mechanisms—change in the rules of exchange and collective-action dilemmas—devised by the interacting individuals, which allow for self-interested individuals to remain prosocial as societies grow