Reproductive inequality in humans and other mammals.

To address claims of human exceptionalism, we determine where humans fit within the greater mammalian distribution of reproductive inequality. We show that humans exhibit lower reproductive skew (i.e., inequality in the number of surviving offspring) among males and smaller sex differences in reproductive skew than most other mammals, while nevertheless falling within the mammalian range. Additionally, female reproductive skew is higher in polygynous human populations than in polygynous nonhumans mammals on average. This patterning of skew can be attributed in part to the prevalence of monogamy in humans compared to the predominance of polygyny in nonhuman mammals, to the limited degree of polygyny in the human societies that practice it, and to the importance of unequally held rival resources to women's fitness. The muted reproductive inequality observed in humans appears to be linked to several unusual characteristics of our species-including high levels of cooperation among males, high dependence on unequally held rival resources, complementarities between maternal and paternal investment, as well as social and legal institutions that enforce monogamous norms.

Content, cost, and context: A framework for understanding human signaling systems.

Humans frequently perform extravagant and seemingly costly behaviors, such as widely sharing hunted resources, erecting conspicuous monumental structures, and performing dramatic acts of religious devotion. Evolutionary anthropologists and archeologists have used signaling theory to explain the function of such displays, drawing inspiration from behavioral ecology, economics, and the social sciences. While signaling theory is broadly aimed at explaining honest communication, it has come to be strongly associated with the handicap principle, which proposes that such costly extravagance is in fact an adaptation for signal reliability. Most empirical studies of signaling theory have focused on obviously costly acts, and consequently anthropologists have likely overlooked a wide range of signals that also promote reliable communication. Here, we build on recent developments in signaling theory and animal communication, developing an updated framework that highlights the diversity of signal contents, costs, contexts, and reliability mechanisms present within human signaling systems. By broadening the perspective of signaling theory in human systems, we strive to identify promising areas for further empirical and theoretical work.

Collective ritual and social support networks in rural South India.

The scholarship on religion has long argued that collective worship helps foster social cohesion. Despite the pervasiveness of this contention, rigorous quantitative evaluations of it have been surprisingly limited. Here, I draw on network data representing the ties of social support among Hindu residents of a South Indian village to evaluate the association between collective religious ritual and social cohesion. I find that those who partake in collective religious rituals together have a higher probability of having a supportive relationship than those who do not. At the structural level, this corresponds to denser connections among co-participants. At the individual level, participants are more embedded in the local community of co-religionists, but are not disassociating themselves from members of other religious denominations. These patterns hold most strongly for co-participation in the recurrent, low-arousal monthly worships at the temple, and are suggestive for co-participation in the intense and dysphoric ritual acts carried out as part of an annual festival. Together, these findings provide clear empirical evidence of the lasting relationship between collective religious ritual and social cohesion.

Positive and negative effects of a threatened parrotfish on reef ecosystems.

Species that are strong interactors play disproportionately important roles in the dynamics of natural ecosystems. It has been proposed that their presence is necessary for positively shaping the structure and functioning of ecosystems. We evaluated this hypothesis using the case of the world's largest parrotfish (Bolbometopon muricatum), a globally imperiled species. We used direct observation, animal tracking, and computer simulations to examine the diverse routes through which B. muricatum affects the diversity, dispersal, relative abundance, and survival of the corals that comprise the foundation of reef ecosystems. Our results suggest that this species can influence reef building corals in both positive and negative ways. Field observation and simulation outputs indicated that B. muricatum reduced the abundance of macroalgae that can outcompete corals, but they also feed directly on corals, decreasing coral abundance, diversity, and colony size. B. muricatum appeared to facilitate coral advancement by mechanically dispersing coral fragments and opening up bare space for coral settlement, but they also damaged adult corals and remobilized a large volume of potentially stressful carbonate sediment. The impacts this species has on reefs appears to be regulated in part by its abundance-the effects of B. muricatum were more intense in simulation scenarios populated with high densities of these fish. Observations conducted in regions with high and low predator (e.g., sharks) abundance generated results that are consistent with the hypothesis that these predators of B. muricatum may play a role in governing their abundance; thus, predation may modulate the intensity of the effects they have on reef dynamics. Overall our results illustrate that functionally unique and threatened species may not have universally positive impacts on ecosystems and that it may be necessary for environmental managers to consider the diverse effects of such species and the forces that mediate the strength of their influence.

Dynamics of cooperative networks associated with gender among South Indian Tamils.

Helping behaviour is thought to play a major role in the evolution of group-living animals. Yet, it is unclear to what extent human males and human females use the same strategies to secure support. Accordingly, we investigate help-seeking over a 5-year period in relation to gender using data from virtually all adults in two Tamil villages (N = 782). Simulations of network dynamics (i.e. stochastic actor-oriented models) calibrated to these data broadly indicate that women are more inclined than men to create and maintain supportive bonds via multiple mechanisms of cooperation (e.g. reciprocity, kin bias, friend bias, generalized exchange). However, gender-related differences in the simulated dynamics of help-seeking are modest, vary based on structural position (e.g. out-degree), and do not appear to translate to divergence in the observed structure of respondents' egocentric networks. Findings ultimately suggest that men and women in the two villages are similarly social but channel their sociality differently. This article is part of the theme issue 'Cooperation among women: evolutionary and cross-cultural perspectives'.

Social hierarchies and social networks in humans.

Across species, social hierarchies are often governed by dominance relations. In humans, where there are multiple culturally valued axes of distinction, social hierarchies can take a variety of forms and need not rest on dominance relations. Consequently, humans navigate multiple domains of status, i.e. relative standing. Importantly, while these hierarchies may be constructed from dyadic interactions, they are often more fundamentally guided by subjective peer evaluations and group perceptions. Researchers have typically focused on the distinct elements that shape individuals' relative standing, with some emphasizing individual-level attributes and others outlining emergent macro-level structural outcomes. Here, we synthesize work across the social sciences to suggest that the dynamic interplay between individual-level and meso-level properties of the social networks in which individuals are embedded are crucial for understanding the diverse processes of status differentiation across groups. More specifically, we observe that humans not only navigate multiple social hierarchies at any given time but also simultaneously operate within multiple, overlapping social networks. There are important dynamic feedbacks between social hierarchies and the characteristics of social networks, as the types of social relationships, their structural properties, and the relative position of individuals within them both influence and are influenced by status differentiation. This article is part of the theme issue 'The centennial of the pecking order: current state and future prospects for the study of dominance hierarchies'.

Four Puzzles of Reputation-Based Cooperation : Content, Process, Honesty, and Structure.

Research in various disciplines has highlighted that humans are uniquely able to solve the problem of cooperation through the informal mechanisms of reputation and gossip. Reputation coordinates the evaluative judgments of individuals about one another. Direct observation of actions and communication are the essential routes that are used to establish and update reputations. In large groups, where opportunities for direct observation are limited, gossip becomes an important channel to share individual perceptions and evaluations of others that can be used to condition cooperative action. Although reputation and gossip might consequently support large-scale human cooperation, four puzzles need to be resolved to understand the operation of reputation-based mechanisms. First, we need empirical evidence of the processes and content that form reputations and how this may vary cross-culturally. Second, we lack an understanding of how reputation is determined from the muddle of imperfect, biased inputs people receive. Third, coordination between individuals is only possible if reputation sharing and signaling is to a large extent reliable and valid. Communication, however, is not necessarily honest and reliable, so theoretical and empirical work is needed to understand how gossip and reputation can effectively promote cooperation despite the circulation of dishonest gossip. Fourth, reputation is not constructed in a social vacuum; hence we need a better understanding of the way in which the structure of interactions affects the efficiency of gossip for establishing reputations and fostering cooperation.

No good deed goes unpunished: the social costs of prosocial behaviour.

Performing costly helpful behaviours can allow individuals to improve their reputation. Those who gain a good reputation are often preferred as interaction partners and are consequently better able to access support through cooperative relationships with others. However, investing in prosocial displays can sometimes yield social costs: excessively generous individuals risk losing their good reputation, and even being vilified, ostracised or antisocially punished. As a consequence, people frequently try to downplay their prosocial actions or hide them from others. In this review, we explore when and why investments in prosocial behaviour are likely to yield social costs. We propose two key features of interactions that make it more likely that generous individuals will incur social costs when: (a) observers infer that helpful behaviour is motivated by strategic or selfish motives; and (b) observers infer that helpful behaviour is detrimental to them. We describe how the cognition required to consider ulterior motives emerges over development and how these tendencies vary across cultures - and discuss how the potential for helpful actions to result in social costs might place boundaries on prosocial behaviour as well as limiting the contexts in which it might occur. We end by outlining the key avenues and priorities for future research.

When does reputation lie? Dynamic feedbacks between costly signals, social capital and social prominence.

Performing a dramatic act of religious devotion, creating an art exhibit, or releasing a new product are all examples of public acts that signal quality and contribute to building a reputation. Signalling theory predicts that these public displays can reliably reveal quality. However, data from ethnographic work in South India suggests that more prominent individuals gain more from reputation-building religious acts than more marginalized individuals. To understand this phenomenon, we extend signalling theory to include variation in people's social prominence or social capital, first with an analytical model and then with an agent-based model. We consider two ways in which social prominence/capital may alter signalling: (i) it impacts observers' priors, and (ii) it alters the signallers' pay-offs. These two mechanisms can result in both a 'reputational shield,' where low quality individuals are able to 'pass' as high quality thanks to their greater social prominence/capital, and a 'reputational poverty trap,' where high quality individuals are unable to improve their standing owing to a lack of social prominence/capital. These findings bridge the signalling theory tradition prominent in behavioural ecology, anthropology and economics with the work on status hierarchies in sociology, and shed light on the complex ways in which individuals make inferences about others. This article is part of the theme issue 'The language of cooperation: reputation and honest signalling'.

Networks of reliable reputations and cooperation: a review.

Reputation has been shown to provide an informal solution to the problem of cooperation in human societies. After reviewing models that connect reputations and cooperation, we address how reputation results from information exchange embedded in a social network that changes endogenously itself. Theoretical studies highlight that network topologies have different effects on the extent of cooperation, since they can foster or hinder the flow of reputational information. Subsequently, we review models and empirical studies that intend to grasp the coevolution of reputations, cooperation and social networks. We identify open questions in the literature concerning how networks affect the accuracy of reputations, the honesty of shared information and the spread of reputational information. Certain network topologies may facilitate biased beliefs and intergroup competition or in-group identity formation that could lead to high cooperation within but conflicts between different subgroups of a network. Our review covers theoretical, experimental and field studies across various disciplines that target these questions and could explain how the dynamics of interactions and reputations help or prevent the establishment and sustainability of cooperation in small- and large-scale societies. This article is part of the theme issue 'The language of cooperation: reputation and honest signalling'.

Community detection with node attributes in multilayer networks.

Community detection in networks is commonly performed using information about interactions between nodes. Recent advances have been made to incorporate multiple types of interactions, thus generalizing standard methods to multilayer networks. Often, though, one can access additional information regarding individual nodes, attributes, or covariates. A relevant question is thus how to properly incorporate this extra information in such frameworks. Here we develop a method that incorporates both the topology of interactions and node attributes to extract communities in multilayer networks. We propose a principled probabilistic method that does not assume any a priori correlation structure between attributes and communities but rather infers this from data. This leads to an efficient algorithmic implementation that exploits the sparsity of the dataset and can be used to perform several inference tasks; we provide an open-source implementation of the code online. We demonstrate our method on both synthetic and real-world data and compare performance with methods that do not use any attribute information. We find that including node information helps in predicting missing links or attributes. It also leads to more interpretable community structures and allows the quantification of the impact of the node attributes given in input.

Church attendance and alloparenting: an analysis of fertility, social support and child development among English mothers.

Many aspects of religious rituals suggest they provide adaptive benefits. Studies across societies consistently find that investments in ritual behaviour return high levels of cooperation. Another line of research finds that alloparental support to mothers increases maternal fertility and improves child outcomes. Although plausible, whether religious cooperation extends to alloparenting and/or affects child development remains unclear. Using 10 years of data collected from the Avon Longitudinal Study of Parents and Children (ALSPAC), we test the predictions that church attendance is positively associated with social support and fertility (n = 8207 to n = 8209), and that social support is positively associated with fertility and child development (n = 1766 to n = 6561). Results show that: (i) relative to not attending, church attendance is positively related to a woman's social network support and aid from co-religionists, (ii) aid from co-religionists is associated with increased family size, while (iii) fertility declines with extra-religious social network support. Moreover, while extra-religious social network support decreased over time, co-religionist aid remained constant. These findings suggest that religious and secular networks differ in their longevity and have divergent influences on a woman's fertility. We find some suggestive evidence that support to mothers and aid from co-religionists is positively associated with a child's cognitive ability at later stages of development. Findings provide mixed support for the premise that ritual, such as church attendance, is part of a strategy that returns high levels of support, fertility and improved child outcomes. Identifying the diversity and scope of cooperative breeding strategies across global religions presents an intriguing new horizon in the evolutionary study of religious systems. This article is part of the theme issue 'Ritual renaissance: new insights into the most human of behaviours'.

Cooperation beyond consanguinity: post-marital residence, delineations of kin and social support among South Indian Tamils.

Evolutionary ecologists have shown that relatives are important providers of support across many species. Among humans, cultural reckonings of kinship are more than just relatedness, as they interact with systems of descent, inheritance, marriage and residence. These cultural aspects of kinship may be particularly important when a person is determining which kin, if any, to call upon for help. Here, we explore the relationship between kinship and cooperation by drawing upon social support network data from two villages in South India. While these Tamil villages have a nominally male-biased kinship system (being patrilocal and patrilineal), matrilateral kin play essential social roles and many women reside in their natal villages, letting us tease apart the relative importance of genetic relatedness, kinship and residence in accessing social support. We find that people often name both their consanguineal and affinal kin as providing them with support, and we see some weakening of support with lesser relatedness. Matrilateral and patrilateral relatives are roughly equally likely to be named, and the greatest distinction instead is in their availability, which is highly contingent on post-marital residence patterns. People residing in their natal village have many more consanguineal relatives present than those who have relocated. Still, relocation has only a small effect on an individual's network size, as non-natal residents are more reliant on the few kin that they have present, most of whom are affines. In sum, marriage patterns have an important impact on kin availability, but the flexibility offered by the broadening of the concept of kin helps people develop the cooperative relationships that they rely upon, even in the absence of genetic relatives. This article is part of the theme issue 'The evolution of female-biased kinship in humans and other mammals'.

Dynamics of beneficial epidemics.

Pathogens can spread epidemically through populations. Beneficial contagions, such as viruses that enhance host survival or technological innovations that improve quality of life, also have the potential to spread epidemically. How do the dynamics of beneficial biological and social epidemics differ from those of detrimental epidemics? We investigate this question using a breadth-first modeling approach involving three distinct theoretical models. First, in the context of population genetics, we show that a horizontally-transmissible element that increases fitness, such as viral DNA, spreads superexponentially through a population, more quickly than a beneficial mutation. Second, in the context of behavioral epidemiology, we show that infections that cause increased connectivity lead to superexponential fixation in the population. Third, in the context of dynamic social networks, we find that preferences for increased global infection accelerate spread and produce superexponential fixation, but preferences for local assortativity halt epidemics by disconnecting the infected from the susceptible. We conclude that the dynamics of beneficial biological and social epidemics are characterized by the rapid spread of beneficial elements, which is facilitated in biological systems by horizontal transmission and in social systems by active spreading behavior of infected individuals.

The social significance of subtle signals.

Acts of prosociality, such as donating to charity, are often analysed in a similar way to acts of conspicuous advertising; both involve costly signals revealing hidden qualities that increase the signaller's prestige. However, experimental work suggests that grand gestures, even if prosocial, may damage one's reputation for trustworthiness and cooperativeness if they are perceived as prestige enhancing: individuals may gain some types of cooperative benefits only when they perform prosocial acts in particular ways. Here, we contrast subtle, less obviously costly, interpersonal forms of prosocial behaviour with high-cost displays to a large audience, drawing on the example of food sharing in subsistence economies. This contrast highlights how highly visible prosocial displays may be effective for attracting new partners, while subtle signals may be crucial for ensuring trust and commitment with long-term partners. Subtle dyadic signals may be key to understanding the long-term maintenance of interpersonal networks that function to reduce unanticipated risks.

Community detection, link prediction, and layer interdependence in multilayer networks.

Complex systems are often characterized by distinct types of interactions between the same entities. These can be described as a multilayer network where each layer represents one type of interaction. These layers may be interdependent in complicated ways, revealing different kinds of structure in the network. In this work we present a generative model, and an efficient expectation-maximization algorithm, which allows us to perform inference tasks such as community detection and link prediction in this setting. Our model assumes overlapping communities that are common between the layers, while allowing these communities to affect each layer in a different way, including arbitrary mixtures of assortative, disassortative, or directed structure. It also gives us a mathematically principled way to define the interdependence between layers, by measuring how much information about one layer helps us predict links in another layer. In particular, this allows us to bundle layers together to compress redundant information and identify small groups of layers which suffice to predict the remaining layers accurately. We illustrate these findings by analyzing synthetic data and two real multilayer networks, one representing social support relationships among villagers in South India and the other representing shared genetic substring material between genes of the malaria parasite.