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Evolutionary scientists studying social and cultural evolution have proposed a multitude of mechanisms by which cultural change can be effected. In this article we discuss two influential ideas from the theory of biological evolution that can inform this debate: the contrast between the micro- and macro-evolution, and the distinction between the tempo and mode of evolution. We add the empirical depth to these ideas by summarizing recent results from the analyses of data on past societies in Seshat: Global History Databank . Our review of these results suggests that the tempo (rates of change, including their acceleration and deceleration) of cultural macroevolution is characterized by periods of apparent stasis interspersed by rapid change. Furthermore, when we focus on large-scale changes in cultural traits of whole groups, the most important macroevolutionary mode involves inter-polity interactions, including competition and warfare, but also cultural exchange and selective imitation; mechanisms that are key components of cultural multilevel selection (CMLS) theory.

Most human societies are characterized by the presence of different identity groups which cooperate but also compete for resources and power. To deepen our understanding of the underlying social dynamics, we model a society subdivided into groups with constant sizes and dynamically changing powers. Both individuals within groups and groups themselves participate in collective actions. The groups are also engaged in political contests over power which determines how jointly produced resources are divided. Using analytical approximations and agent-based simulations, we show that the model exhibits rich behavior characterized by multiple stable equilibria and, under some conditions, non-equilibrium dynamics. We demonstrate that societies in which individuals act independently are more stable than those in which actions of individuals are completely synchronized. We show that mechanisms preventing politically powerful groups from bending the rules of competition in their favor play a key role in promoting between-group cooperation and reducing inequality between groups. We also show that small groups can be more successful in competition than large groups if the jointly-produced goods are rivalrous and the potential benefit of cooperation is relatively small. Otherwise large groups dominate. Overall our model contributes towards a better understanding of the causes of variation between societies in terms of the economic and political inequality within them., (© 2021. The Author(s).)

Human decision-making is affected by a diversity of factors including material cost-benefit considerations, normative and cultural influences, learning and conformity with peers and external authorities (e.g. cultural, religious, political, organisational). Also important are dynamically changing personal perceptions of the situation and beliefs about actions and expectations of others as well as psychological phenomena such as cognitive dissonance and social projection. To better understand these processes, I develop a unifying modelling framework describing the joint dynamics of actions and attitudes of individuals and their beliefs about the actions and attitudes of their groupmates. I consider which norms get internalised and which factors control beliefs about others. I predict that the long-term average characteristics of groups are largely determined by a balance between material payoffs and the values promoted by the external authority. Variation around these averages largely reflects variation in individual costs and benefits mediated by individual psychological characteristics. The efforts of an external authority to change the group behaviour in a certain direction can, counter-intuitively, have an opposite effect on individual behaviour. I consider how various factors can affect differences between groups and societies in the tightness/looseness of their social norms. I show that the most important factors are social heterogeneity, societal threat, effects of authority, cultural variation in the degree of collectivism/individualism, the population size and the subsistence style. My results can be useful for achieving a better understanding of human social behaviour and historical and current social processes, and in developing more efficient policies aiming to modify social behaviour., (© The Author(s) 2021.)

Charles Darwin's The Descent of Man , published 150 years ago, laid the grounds for scientific studies into human origins and evolution. Three of his insights have been reinforced by modern science. The first is that we share many characteristics (genetic, developmental, physiological, morphological, cognitive, and psychological) with our closest relatives, the anthropoid apes. The second is that humans have a talent for high-level cooperation reinforced by morality and social norms. The third is that we have greatly expanded the social learning capacity that we see already in other primates. Darwin's emphasis on the role of culture deserves special attention because during an increasingly unstable Pleistocene environment, cultural accumulation allowed changes in life history; increased cognition; and the appearance of language, social norms, and institutions., (Copyright © 2021 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works.)

AbstractThe patterns and outcomes of coevolution are expected to depend on intraspecific trait variation. Various evolutionary factors can change this variation in time. As a result, modeling coevolutionary processes solely in terms of mean trait values may not be sufficient; one may need to study the dynamics of the whole trait distribution. Here, we develop a theoretical framework for studying the effects of evolving intraspecific variation in two-species coevolutionary systems. In particular, we build and study mathematical models of competition, exploiter-victim interactions, and mutualism in which the strength of within- and between-species interactions depends on the difference in continuously varying traits between individuals reproducing asexually. We use analytical approximations based on the invasion analysis and supplement them with numerical results. We find that intraspecific variation can be maintained if stabilizing selection is weak in at least one species. When intraspecific variation is maintained under competition or mutualism, coexistence in a stable equilibrium is promoted when between-species interactions mostly happen between individuals similar in trait values. In contrast, in exploiter-victim systems coexistence typically requires strong interactions between dissimilar exploiters and victims. We show that trait distributions can become multimodal. Our approach and results contribute to the understanding of the ecological consequences of intraspecific variation in coevolutionary systems by exploring its effects on population densities and trait distributions.

Injunctive social norms are behaviours that one is expected to follow and expects others to follow in a given social situation; they are maintained by the threat of disapproval or punishment and by the process of internalization. Injunctive norms govern all aspects of our social life but the understanding of their effects on individual and group behaviour is currently rather incomplete. Here I develop a general mathematical approach describing the dynamics of injunctive norms in heterogeneous groups. My approach captures various costs and benefits, both material and normative, associated with norm-related behaviours including punishment and disapproval by others. It also allows for errors in decision-making and explicitly accounts for differences between individuals in their values, beliefs about the population state, and sensitivity to the actions of others. In addition, it enables one to study the consequences of mixing populations with different normative values and the effects of persuasive interventions. I describe how interactions of these factors affect individual and group behaviour. As an illustration, I consider policies developed by practitioners to abolish the norms of footbinding and female genital cutting, to decrease college students' drinking, and to increase pro-environmental behaviours. The theory developed here can be used for achieving a better understanding of historical and current social processes as well as for developing practical policies better accounting for human social behaviour., (© The Author(s) 2020.)

Inequality or skew in reproductive success (RS) is common across many animal species and is of long-standing interest to the study of social evolution. However, the measurement of inequality in RS in natural populations has been challenging because existing quantitative measures are highly sensitive to variation in group/sample size, mean RS, and age-structure. This makes comparisons across multiple groups and/or species vulnerable to statistical artefacts and hinders empirical and theoretical progress. Here, we present a new measure of reproductive skew, the multinomial index, M , that is unaffected by many of the structural biases affecting existing indices. M is analytically related to Nonacs' binomial index, B , and comparably accounts for heterogeneity in age across individuals; in addition, M allows for the possibility of diminishing or even highly nonlinear RS returns to age. Unlike B , however, M is not biased by differences in sample/group size. To demonstrate the value of our index for cross-population comparisons, we conduct a reanalysis of male reproductive skew in 31 primate species. We show that a previously reported negative effect of group size on mating skew was an artefact of structural biases in existing skew measures, which inevitably decline with group size; this bias disappears when using M . Applying phylogenetically controlled, mixed-effects models to the same dataset, we identify key similarities and differences in the inferred within- and between-species predictors of reproductive skew across metrics. Finally, we provide an R package, SkewCalc, to estimate M from empirical data.

Many animal and human societies exhibit hierarchical structures with different degrees of steepness. Some of these societies also show cooperative behavior, where cooperation means working together for a common benefit. However, there is an increasing evidence that rigidly enforced hierarchies lead to a decrease of cooperation in both human and non-human primates. In this work, we address this issue by means of an evolutionary agent-based model that incorporates fights as social interactions governing a dynamic ranking, communal work to produce a public good, and norm internalization, i.e. a process where acting according to a norm becomes a goal in itself. Our model also includes the perception of how much the individual is going to retain from her cooperative behavior in future interactions. The predictions of the model resemble the principal characteristics of human societies. When ranking is unconstrained, we observe a high concentration of agents in low scores, while a few ones climb up the social hierarchy and exploit the rest, with no norm internalization. If ranking is constrained, thus leading to bounded score differences between agents, individual positions in the ranking change more, and the typical structure shows a division of the society in upper and lower classes. In this case, we observe that there is a significant degree of norm internalization, supporting large fractions of the population cooperating in spite of the rank differences. Our main results are robust with respect to the model parameters and to the type of rank constraint. We thus provide a mechanism that can explain how hierarchy arises in initially egalitarian societies while keeping a large degree of cooperation.

There has been much interest in the evolutionary forces responsible for, and underlying the diversity in, female primate reproductive cycles. While there has been limited research on sexual receptivity in primates, this has been one recurring topic of interest. Some primate species are like humans, sexually receptive to mating throughout their entire estrus cycle, while other species are the opposite, receptive for mere hours out of their several-week cycles. Why is there such prominent variation in sexual receptivity length among primate species? Here we examine the evolutionary trade-offs associated with sexual receptivity length using mathematical modeling. We investigate how various factors, including having ovulation signs present versus concealed ovulation, female physiological costs, and group size, each influence the length of females' receptive periods. We find that both continuous receptivity and very short lengths of receptivity are able to evolve. Our model predicts that increasing the impacts of infanticide will increase the length of the female receptive period, emphasizing the possible importance of paternity confusion. Similar effects can also be achieved by increasing the non-genetic benefits provided by males. Overall, our work offers a theoretical framework for understanding the evolution and diversity of mating traits in female primates.

Modern speciation theory has greatly benefited from a variety of simple mathematical models focusing on the conditions and patterns of speciation and diversification in the presence of gene flow. Unfortunately the application of general theoretical concepts and tools to specific ecological systems remains a challenge. Here we apply modeling tools to better understand adaptive divergence of whitefish during the postglacial period in lakes of northern Fennoscandia. These lakes harbor up to three different morphs associated with the three major lake habitats: littoral, pelagic, and profundal. Using large-scale individual-based simulations, we aim to identify factors required for in situ emergence of the pelagic and profundal morphs in lakes initially colonized by the littoral morph. The importance of some of the factors we identify and study - sufficiently large levels of initial genetic variation, size- and habitat-specific mating, sufficiently large carrying capacity of the new niche - is already well recognized. In addition, our model also points to two other factors that have been largely disregarded in theoretical studies: fitness-dependent dispersal and strong predation in the ancestral niche coupled with the lack of it in the new niche(s). We use our theoretical results to speculate about the process of diversification of whitefish in Fennoscandia and to identify potentially profitable directions for future empirical research.

Leadership can be effective in promoting cooperation within a group, but as the saying goes "heavy is the head that wears the crown". A lot of debate still surrounds exactly what motivates individuals to expend the effort necessary to lead their groupmates. Evolutionary game theoretic models represent individual's thought processes by strategy update protocols. The most common of these are random mutation, individual learning, selective imitation, and myopic optimization. Recently we introduced a new strategy update protocol - foresight - which takes into account future payoffs, and how groupmates respond to one's own strategies. Here we apply our approach to a new 2 × 2 game, where one player, a leader, ensures via inspection and punishment that the other player, a subordinate, produces collective good. We compare the levels of inspection and production predicted by Nash Equilibrium, Quantal Response Equilibrium, level-k cognition, fictitious play, reinforcement learning, selective payoff-biased imitation, and foresight. We show that only foresight and selective imitation are effective at promoting contribution by the subordinate and inspection and punishment by the leader. The role of selective imitation in cultural and social evolution is well appreciated. In line with our prior findings, foresight is a viable alternative route to cooperation.

A long-standing evolutionary puzzle surrounds female sexual signals visible around the time of ovulation. Even among just primates, why do some species have substantial sexual swellings and/or bright colorations visible around females' genital regions, while other species are like humans, with no signs of ovulation visible? What is the evolutionary purpose behind not just these signs, but also this great variation seen among species? Here, we examine the evolutionary trade-offs associated with visual ovulation signalling using agent-based modelling. Our model predicts how various factors, including male genetic heterogeneity and reproductive inequality, female physiological costs, group size, and the weighting of genetic versus non-genetic benefits coming from males, each influence the strength of ovulation signalling. Our model also predicts that increasing the impacts of infanticide will increase ovulation signalling. We use comparative primate data to show that, as predicted by our model, larger group size and higher risk of infanticide each correlate with having stronger visual ovulation signs. Overall, our work resolves some old controversies and sheds new light on the evolution of visual female sexual signalling., (© 2018 The Author(s).)

Human behavior is strongly affected by culturally transmitted norms and values. Certain norms are internalized (i.e., acting according to a norm becomes an end in itself rather than merely a tool in achieving certain goals or avoiding social sanctions). Humans' capacity to internalize norms likely evolved in our ancestors to simplify solving certain challenges-including social ones. Here we study theoretically the evolutionary origins of the capacity to internalize norms. In our models, individuals can choose to participate in collective actions as well as punish free riders. In making their decisions, individuals attempt to maximize a utility function in which normative values are initially irrelevant but play an increasingly important role if the ability to internalize norms emerges. Using agent-based simulations, we show that norm internalization evolves under a wide range of conditions so that cooperation becomes "instinctive." Norm internalization evolves much more easily and has much larger effects on behavior if groups promote peer punishment of free riders. Promoting only participation in collective actions is not effective. Typically, intermediate levels of norm internalization are most frequent but there are also cases with relatively small frequencies of "oversocialized" individuals willing to make extreme sacrifices for their groups no matter material costs, as well as "undersocialized" individuals completely immune to social norms. Evolving the ability to internalize norms was likely a crucial step on the path to large-scale human cooperation., Competing Interests: The authors declare no conflict of interest.

When group cohesion is essential, groups must have efficient strategies in place for consensus decision-making. Recent theoretical work suggests that shared decision-making is often the most efficient way for dealing with both information uncertainty and individual variation in preferences. However, some animal and most human groups make collective decisions through particular individuals, leaders, that have a disproportionate influence on group decision-making. To address this discrepancy between theory and data, we study a simple, but general, model that explicitly focuses on the dynamics of consensus building in groups composed by individuals who are heterogeneous in preferences, certain personality traits (agreeability and persuasiveness), reputation, and social networks. We show that within-group heterogeneity can significantly delay democratic consensus building as well as give rise to the emergence of informal leaders, i.e. individuals with a disproportionately large impact on group decisions. Our results thus imply strong benefits of leadership particularly when groups experience time pressure and significant conflict of interest between members (due to various between-individual differences). Overall, our models shed light on why leadership and decision-making hierarchies are widespread, especially in human groups.

The sexes share the same autosomal genomes, yet sexual dimorphism is common due to sex-specific gene expression. When present, XX and XY karyotypes trigger alternate regulatory cascades that determine sex-specific gene expression profiles. In mammals, secretion of testosterone (T) by the testes during foetal development is the master switch influencing the gene expression pathways (male vs. female) that will be followed, but many genes have sex-specific expression prior to T secretion. Environmental factors, like endocrine disruptors and mimics, can interfere with sexual development. However, sex-specific ontogeny can be canalized by the production of epigenetic marks (epimarks) generated during early ontogeny that increase sensitivity of XY embryos to T and decrease sensitivity of XX embryos. Here, we integrate and synthesize the evidence indicating that canalizing epimarks are produced during early ontogeny. We will also describe the evidence that such epimarks sometimes carry over across generations and produce mosaicism in which some traits are discordant with the gonad. Such carryover epimarks are sexually antagonistic because they benefit the individual in which they were formed (via canalization) but harm opposite-sex offspring when they fail to erase across generations and produce gonad-trait discordances. SA-epimarks have the potential to: i) magnify phenotypic variation for many sexually selected traits, ii) generate overlap along many dimensions of the masculinity/femininity spectrum, and iii) influence medically important gonad-trait discordances like cryptorchidism, hypospadias and idiopathic hirsutism., (© 2015 John Wiley & Sons Ltd.)

Leadership is an active area of research in both the biological and social sciences. This review provides a transdisciplinary synthesis of biological and social-science views of leadership from an evolutionary perspective, and examines patterns of leadership in a set of small-scale human and non-human mammalian societies. We review empirical and theoretical work on leadership in four domains: movement, food acquisition, within-group conflict mediation, and between-group interactions. We categorize patterns of variation in leadership in five dimensions: distribution (across individuals), emergence (achieved versus inherited), power, relative payoff to leadership, and generality (across domains). We find that human leadership exhibits commonalities with and differences from the broader mammalian pattern, raising interesting theoretical and empirical issues., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

I review the theoretical and experimental literature on the collective action problem in groups whose members differ in various characteristics affecting individual costs, benefits and preferences in collective actions. I focus on evolutionary models that predict how individual efforts and fitnesses, group efforts and the amount of produced collective goods depend on the group's size and heterogeneity, as well as on the benefit and cost functions and parameters. I consider collective actions that aim to overcome the challenges from nature or win competition with neighbouring groups of co-specifics. I show that the largest contributors towards production of collective goods will typically be group members with the highest stake in it or for whom the effort is least costly, or those who have the largest capability or initial endowment. Under some conditions, such group members end up with smaller net pay-offs than the rest of the group. That is, they effectively behave as altruists. With weak nonlinearity in benefit and cost functions, the group effort typically decreases with group size and increases with within-group heterogeneity. With strong nonlinearity in benefit and cost functions, these patterns are reversed. I discuss the implications of theoretical results for animal behaviour, human origins and psychology., (© 2015 The Author(s).)

Models of collective action infrequently account for differences across individuals beyond a limited set of strategies, ignoring variation in endowment (e.g. physical condition, wealth, knowledge, personality, support), individual costs of effort, or expected gains from cooperation. However, behavioural research indicates these inter-individual differences can have significant effects on the dynamics of collective action. The papers contributed to this theme issue evaluate how individual differences affect the propensity to cooperate, and how they can catalyse others' likelihood of cooperation (e.g. via leadership). Many of the papers emphasize the relationship between individual decisions and socio-ecological context, particularly the effect of group size. All together, the papers in this theme issue provide a more complete picture of collective action, by embracing the reality of inter-individual variation and its multiple roles in the success or failure of collective action., (© 2015 The Author(s).)

Humans are unique both in their cognitive abilities and in the extent of cooperation in large groups of unrelated individuals. How our species evolved high intelligence in spite of various costs of having a large brain is perplexing. Equally puzzling is how our ancestors managed to overcome the collective action problem and evolve strong innate preferences for cooperative behaviour. Here, I theoretically study the evolution of social-cognitive competencies as driven by selection emerging from the need to produce public goods in games against nature or in direct competition with other groups. I use collaborative ability in collective actions as a proxy for social-cognitive competencies. My results suggest that collaborative ability is more likely to evolve first by between-group conflicts and then later be utilized and improved in games against nature. If collaborative abilities remain low, the species is predicted to become genetically dimorphic with a small proportion of individuals contributing to public goods and the rest free-riding. Evolution of collaborative ability creates conditions for the subsequent evolution of collaborative communication and cultural learning.

Parallel adaptation is common and may often occur from shared genetic variation, but the genomic consequences of this process remain poorly understood. We first use individual-based simulations to demonstrate that comparisons between populations adapted in parallel to similar environments from shared variation reveal a characteristic genomic signature around a selected locus: a low-divergence valley centred at the locus and flanked by twin peaks of high divergence. This signature is initiated by the hitchhiking of haplotype tracts differing between derived populations in the broader neighbourhood of the selected locus (driving the high-divergence twin peaks) and shared haplotype tracts in the tight neighbourhood of the locus (driving the low-divergence valley). This initial hitchhiking signature is reinforced over time because the selected locus acts as a barrier to gene flow from the source to the derived populations, thus promoting divergence by drift in its close neighbourhood. We next empirically confirm the peak-valley-peak signature by combining targeted and RAD sequence data at three candidate adaptation genes in multiple marine (source) and freshwater (derived) populations of threespine stickleback. Finally, we use a genome-wide screen for the peak-valley-peak signature to discover additional genome regions involved in parallel marine-freshwater divergence. Our findings offer a new explanation for heterogeneous genomic divergence and thus challenge the standard view that peaks in population divergence harbour divergently selected loci and that low-divergence regions result from balancing selection or localized introgression. We anticipate that genome scans for peak-valley-peak divergence signatures will promote the discovery of adaptation genes in other organisms., (© 2014 John Wiley & Sons Ltd.)

Conflict with conspecifics from neighbouring groups over territory, mating opportunities and other resources is observed in many social organisms, including humans. Here we investigate the evolutionary origins of social instincts, as shaped by selection resulting from between-group conflict in the presence of a collective action problem. We focus on the effects of the differences between individuals on the evolutionary dynamics. Our theoretical models predict that high-rank individuals, who are able to usurp a disproportional share of resources in within-group interactions, will act seemingly altruistically in between-group conflict, expending more effort and often having lower reproductive success than their low-rank group-mates. Similar behaviour is expected for individuals with higher motivation, higher strengths or lower costs, or for individuals in a leadership position. Our theory also provides an evolutionary foundation for classical equity theory, and it has implications for the origin of coercive leadership and for reproductive skew theory.

Theory building is an integral part of biological research, in general, and of speciation research, in particular. Here, I review the modeling work on speciation done in the last 10 years or so, assessing the progress made and identifying areas where additional effort is required. Specific topics considered include evolutionary dynamics of genetic incompatibilities, spatial and temporal patterns of speciation, links to neutral theory of biodiversity, effects of multidimensionality of phenotype, sympatric and parapatric speciation, adaptive radiation, speciation by sexual conflict, and models tailored for specific biological systems. Particularly challenging questions for future theoretical research identified here are 1) incorporating gene regulatory networks in models describing accumulation of genetic incompatibilities; 2) integrating models of community ecology with those developed in speciation theory; 3) building models providing better insights on the dynamics of parapatric speciation; 4) modeling speciation in multidimensional ecological niches with mating preferences based on multidimensional mating cues and sexual characters; 5) linking microevolutionary processes with macroevolutionary patterns as observed in adaptive radiations and paleontological record; 6) modeling speciation in specific systems studied by empirical biologists; and 7) modeling human origins. The insights from dynamic models of speciation should be useful in developing statistical tools that would enable empiricists to infer the history of past evolutionary divergence and speciation from genomic data., (© The American Genetic Association. 2014. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

How did human societies evolve from small groups, integrated by face-to-face cooperation, to huge anonymous societies of today, typically organized as states? Why is there so much variation in the ability of different human populations to construct viable states? Existing theories are usually formulated as verbal models and, as a result, do not yield sharply defined, quantitative predictions that could be unambiguously tested with data. Here we develop a cultural evolutionary model that predicts where and when the largest-scale complex societies arose in human history. The central premise of the model, which we test, is that costly institutions that enabled large human groups to function without splitting up evolved as a result of intense competition between societies-primarily warfare. Warfare intensity, in turn, depended on the spread of historically attested military technologies (e.g., chariots and cavalry) and on geographic factors (e.g., rugged landscape). The model was simulated within a realistic landscape of the Afroeurasian landmass and its predictions were tested against a large dataset documenting the spatiotemporal distribution of historical large-scale societies in Afroeurasia between 1,500 BCE and 1,500 CE. The model-predicted pattern of spread of large-scale societies was very similar to the observed one. Overall, the model explained 65% of variance in the data. An alternative model, omitting the effect of diffusing military technologies, explained only 16% of variance. Our results support theories that emphasize the role of institutions in state-building and suggest a possible explanation why a long history of statehood is positively correlated with political stability, institutional quality, and income per capita.

Many social behaviors are triggered by social partners. For example, cells in a multicellular organism often become soma via extrinsically regulated differentiation, while individuals in a eusocial colony often become helpers via extrinsic caste determination. One explanation for social triggering is that it informs when it is beneficial to express the behavior. Alternatively, social triggering can represent manipulation where social partners partially or completely control the focal individual's behavior. For instance, caste determination in primitively eusocial taxa is typically accomplished via differential feeding or dominance hierarchies, suggesting some manipulation. However, selection would favor resistance if manipulation is detrimental to manipulated parties, and the outcome of the manipulation conflict remains intricate. We analyze the coevolution of manipulation and resistance in a simple but general setting. We show that, despite possible resistance, manipulated behavior can be established under less stringent conditions than spontaneous (i.e., nonmanipulated) behavior because of resistance costs. The existence of this advantage might explain why primitive eusocial behavior tends to be triggered socially and coercively. We provide a simple condition for the advantage of manipulated behavior that may help infer whether a socially triggered behavior is manipulated. We illustrate our analysis with a hypothetical example of maternal manipulation relevant to primitive eusociality.

We recently synthesized and reinterpreted published studies to advance an epigenetic model for the development of homosexuality (HS). The model is based on epigenetic marks laid down in response to the XX vs. XY karyotype in embryonic stem cells. These marks boost sensitivity to testosterone in XY fetuses and lower it in XX fetuses, thereby canalizing sexual development. Our model predicts that a subset of these canalizing epigenetic marks stochastically carry over across generations and lead to mosaicism for sexual development in opposite-sex offspring--the homosexual phenotype being one such outcome. Here, we begin by outlining why HS has been under-appreciated as a commonplace phenomenon in nature, and how this trend is currently being reversed in the field of neurobiology. We next briefly describe our epigenetic model of HS, develop a set of predictions, and describe how epigenetic profiles of human stem cells can provide for a strong test of the model., (© 2013 The Authors. Bioessays published by WILEY Periodicals, Inc.)

Non-random mating provides multiple evolutionary benefits and can result in speciation. Biological organisms are characterised by a myriad of different traits, many of which can serve as mating cues. We consider multiple mechanisms of non-random mating simultaneously within a unified modelling framework in an attempt to understand better which are more likely to evolve in natural populations going through the process of local adaptation and ecological speciation. We show that certain traits that are under direct natural selection are more likely to be co-opted as mating cues, leading to the appearance of magic traits (i.e. phenotypic traits involved in both local adaptation and mating decisions). Multiple mechanisms of non-random mating can interact so that trait co-evolution enables the evolution of non-random mating mechanisms that would not evolve alone. The presence of magic traits may suggest that ecological selection was acting during the origin of new species., (© 2013 John Wiley & Sons Ltd/CNRS.)

Male and female homosexuality have substantial prevalence in humans. Pedigree and twin studies indicate that homosexuality has substantial heritability in both sexes, yet concordance between identical twins is low and molecular studies have failed to find associated DNA makers. This paradoxical pattern calls for an explanation. We use published data on fetal androgen signaling and gene regulation via nongenetic changes in DNA packaging (epigenetics) to develop a new model for homosexuality. It is well established that fetal androgen signaling strongly influences sexual development. We show that an unappreciated feature of this process is reduced androgen sensitivity in XX fetuses and enhanced sensitivity in XY fetuses, and that this difference is most feasibly mused by numerous sex-specific epigenetic modifications ("epi-marks") originating in embryonic stem cells. These epi-marks buffer XX fetuses from masculinization due to excess fetal androgen exposure and similarly buffer XY fetuses from androgen underexposure. Extant data indicates that individual epi-marks influence some but not other sexually dimorphic traits, vary in strength across individuals, and are produced during ontogeny and erased between generations. Those that escape erasure will steer development of the sexual phenotypes they influence in a gonad-discordant direction in opposite sex offspring, mosaically feminizing XY offspring and masculinizing XX offspring. Such sex-specific epi-marks are sexually antagonistic (SA-epi-marks) because they canalize sexual development in the parent that produced them, but contribute to gonad-trait discordances in opposite-sex offspring when unerased. In this model, homosexuality occurs when stronger-than-average SA-epi-marks (influencing sexual preference) from an opposite-sex parent escape erasure and are then paired with a weaker-than-average de novo sex-specific epi-marks produced in opposite-sex offspring. Our model predicts that homosexuality is part of a wider phenomenon in which recently evolved androgen-influenced traits commonly display gonad-trait discordances at substantial frequency, and that the molecular feature underlying most homosexuality is not DNA polymorphism(s), but epi-marks that evolved to canalize sexual dimorphic development that sometimes carryover across generations and contribute to gonad-trait discordances in opposite-sex descendants.

The evolutionary emergence of the egalitarian syndrome is one of the most intriguing unsolved puzzles related to the origins of modern humans. Standard explanations and models for cooperation and altruism--reciprocity, kin and group selection, and punishment--are not directly applicable to the emergence of egalitarian behavior in hierarchically organized groups that characterized the social life of our ancestors. Here I study an evolutionary model of group-living individuals competing for resources and reproductive success. In the model, the differences in fighting abilities lead to the emergence of hierarchies where stronger individuals take away resources from weaker individuals and, as a result, have higher reproductive success. First, I show that the logic of within-group competition implies under rather general conditions that each individual benefits if the transfer of the resource from a weaker group member to a stronger one is prevented. This effect is especially strong in small groups. Then I demonstrate that this effect can result in the evolution of a particular, genetically controlled psychology causing individuals to interfere in a bully-victim conflict on the side of the victim. A necessary condition is a high efficiency of coalitions in conflicts against the bullies. The egalitarian drive leads to a dramatic reduction in within-group inequality. Simultaneously it creates the conditions for the emergence of inequity aversion, empathy, compassion, and egalitarian moral values via the internalization of behavioral rules imposed by natural selection. It also promotes widespread cooperation via coalition formation.

A crucial step in recent theories of human origins is the emergence of strong pair-bonding between males and females accompanied by a dramatic reduction in the male-to-male conflict over mating and an increased investment in offspring. How such a transition from promiscuity to pair-bonding could be achieved is puzzling. Many species would, indeed, be much better off evolutionarily if the effort spent on male competition over mating was redirected to increasing female fertility or survivorship of offspring. Males, however, are locked in a "social dilemma," where shifting one's effort from "appropriation" to "production" would give an advantage to free-riding competitors and therefore, should not happen. Here, I first consider simple models for four prominent scenarios of the human transition to pair-bonding: communal care, mate guarding, food for mating, and mate provisioning. I show that the transition is not feasible under biologically relevant conditions in any of these models. Then, I show that the transition can happen if one accounts for male heterogeneity, assortative pair formation, and evolution of female choice and faithfulness. This process is started when low-ranked males begin using an alternative strategy of female provisioning. At the end, except for the top-ranked individuals, males invest exclusively in provisioning females who have evolved very high fidelity to their mates. My results point to the crucial importance of female choice and emphasize the need for incorporating between-individual variation in theoretical and empirical studies of social dilemmas and behaviors.

The exact nature of the relationship among species range sizes, speciation, and extinction events is not well understood. The factors that promote larger ranges, such as broad niche widths and high dispersal abilities, could increase the likelihood of encountering new habitats but also prevent local adaptation due to high gene flow. Similarly, low dispersal abilities or narrower niche widths could cause populations to be isolated, but such populations may lack advantageous mutations due to low population sizes. Here we present a large-scale, spatially explicit, individual-based model addressing the relationships between species ranges, speciation, and extinction. We followed the evolutionary dynamics of hundreds of thousands of diploid individuals for 200,000 generations. Individuals adapted to multiple resources and formed ecological species in a multidimensional trait space. These species varied in niche widths, and we observed the coexistence of generalists and specialists on a few resources. Our model shows that species ranges correlate with dispersal abilities but do not change with the strength of fitness trade-offs; however, high dispersal abilities and low resource utilization costs, which favored broad niche widths, have a strong negative effect on speciation rates. An unexpected result of our model is the strong effect of underlying resource distributions on speciation: in highly fragmented landscapes, speciation rates are reduced., (© 2011 by The University of Chicago.)

More than half a century has now elapsed since coalition or alliance formation theory (CAFT) was first developed. During that time, researchers have amassed a vast amount of detailed and high-quality data on coalitions or alliances among primates and other animals. But models have not kept pace, and more relevant theory is needed. In particular, even though CAFT is primarily an exercise in polyadic game theory, game theorists have devoted relatively little attention to questions that motivate field research, and much remains largely unexplored. The state of the art is both a challenge and an opportunity. In this review we describe a variety of game-theoretic and related modelling approaches that have much untapped potential to address the questions that field biologists ask., (Copyright © 2010 Elsevier Ltd. All rights reserved.)

Recent empirical studies indicate that grandparents favour some categories of grandchildren over others. Here, we expand the previous theoretical foundation for this finding and show that grandchild-harming phenotypes are predicted to evolve by 'sexually antagonistic zygotic drive (SA-zygotic drive) of the sex chromosomes'. We use the logic of Hamilton's rule to develop a new 'no-cost-to-self nepotism rule' that greatly simplifies the determination of the invasion criteria for mutations that cause grandparents to harm grandchildren. We use this theory to generate predictions that distinguish SA-zygotic drive from theory based solely on paternity assurance. The major diagnostic prediction is that grandmothers, and to a lesser degree grandfathers, will evolve grandson-harming phenotypes that reduce the level of sib competition experienced by their more closely related granddaughters, especially in their sons' families. This prediction is supported by data from recent studies showing (i) grandmothers invest more in granddaughters than grandsons, and counterintuitively, (ii) paternal grandmothers reduce the survival of their grandsons. We conclude that SA-zygotic drive is plausibly operating in humans via sexually antagonistic grandparental care.

Emerging diseases represent a growing worldwide problem accompanying global environmental changes. There is tremendous interest in identifying the factors controlling the appearance and spread of these diseases. Here, we discuss emerging fungal plant diseases, and argue that they often result from host shift speciation (a particular case of ecological speciation). We consider the factors controlling local adaptation and ecological speciation, and show that certain life-history traits of many fungal plant pathogens are conducive for rapid ecological speciation, thus favoring the emergence of novel pathogen species adapted to new hosts. We argue that placing the problem of emerging fungal diseases of plants within the context of ecological speciation can significantly improve our understanding of the biological mechanisms governing the emergence of such diseases., (Copyright 2010 Elsevier Ltd. All rights reserved.)

A crucial step in several major evolutionary transitions is the division of labor between components of the emerging higher-level evolutionary unit. Examples include the separation of germ and soma in simple multicellular organisms, appearance of multiple cell types and organs in more complex organisms, and emergence of casts in eusocial insects. How the division of labor was achieved in the face of selfishness of lower-level units is controversial. I present a simple mathematical model describing the evolutionary emergence of the division of labor via developmental plasticity starting with a colony of undifferentiated cells and ending with completely differentiated multicellular organisms. I explore how the plausibility and the dynamics of the division of labor depend on its fitness advantage, mutation rate, costs of developmental plasticity, and the colony size. The model shows that the transition to differentiated multicellularity, which has happened many times in the history of life, can be achieved relatively easily. My approach is expandable in a number of directions including the emergence of multiple cell types, complex organs, or casts of eusocial insects.

The two kinds of sex chromosomes in the heterogametic parent are transmitted to offspring with different sexes, causing opposite-sex siblings to be completely unrelated for genes located on these chromosomes. Just as the nest-parasitic cuckoo chick is selected to harm its unrelated nest-mates in order to garner more shared resources, sibling competition causes the sex chromosomes to be selected to harm siblings that do not carry them. Here we quantify and contrast this selection on the X and Y, or Z and W, sex chromosomes. We also develop a hypothesis for how this selection can contribute to the decay of the non-recombining sex chromosome.

We build a spatial individual-based multilocus model of homoploid hybrid speciation tailored for a tentative case of hybrid origin of Heliconius heurippa from H. melpomene and H. cydno in South America. Our model attempts to account for empirical patterns and data on genetic incompatibility, mating preferences and selection by predation (both based on coloration patterns), habitat preference, and local adaptation for all three Heliconius species. Using this model, we study the likelihood of recombinational speciation and identify the effects of various ecological and genetic parameters on the dynamics, patterns, and consequences of hybrid ecological speciation. Overall, our model supports the possibility of hybrid origin of H. heurippa under certain conditions. The most plausible scenario would include hybridization between H. melpomene and H. cydno in an area geographically isolated from the rest of both parental species with subsequent long-lasting geographic isolation of the new hybrid species, followed by changes in the species ranges, the secondary contact, and disappearance of H. melpomene-type ecomorph in the hybrid species. However, much more work (both empirical and theoretical) is necessary to be able to make more definite conclusions on the importance of homoploid hybrid speciation in animals.

Biologists have long been fascinated by the exceptionally high diversity displayed by some evolutionary groups. Adaptive radiation in such clades is not only spectacular, but is also an extremely complex process influenced by a variety of ecological, genetic, and developmental factors and strongly dependent on historical contingencies. Using modeling approaches, we identify 10 general patterns concerning the temporal, spatial, and genetic/morphological properties of adaptive radiation. Some of these are strongly supported by empirical work, whereas for others, empirical support is more tentative. In almost all cases, more data are needed. Future progress in our understanding of adaptive radiation will be most successful if theoretical and empirical approaches are integrated, as has happened in other areas of evolutionary biology.

Genomic conflict is perplexing because it causes the fitness of a species to decline rather than improve. Many diverse forms of genomic conflict have been identified, but this extant tally may be incomplete. Here, we show that the unusual characteristics of the sex chromosomes can, in principle, lead to a previously unappreciated form of sexual genomic conflict. The phenomenon occurs because there is selection in the heterogametic sex for sex-linked mutations that harm the sex of offspring that does not carry them, whenever there is competition among siblings. This harmful phenotype can be expressed as an antagonistic green-beard effect that is mediated by epigenetic parental effects, parental investment, and/or interactions among siblings. We call this form of genomic conflict sexually antagonistic "zygotic drive", because it is functionally equivalent to meiotic drive, except that it operates during the zygotic and postzygotic stages of the life cycle rather than the meiotic and gametic stages. A combination of mathematical modeling and a survey of empirical studies is used to show that sexually antagonistic zygotic drive is feasible, likely to be widespread in nature, and that it can promote a genetic "arms race" between the homo- and heteromorphic sex chromosomes. This new category of genomic conflict has the potential to strongly influence other fundamental evolutionary processes, such as speciation and the degeneration of the Y and W sex chromosomes. It also fosters a new genetic hypothesis for the evolution of enigmatic fitness-reducing traits like the high frequency of spontaneous abortion, sterility, and homosexuality observed in humans., Competing Interests: The authors have declared that no competing interests exist.

We study victim-exploiter coevolution in a spatially heterogeneous island model. In each species, fitness consequences of between-species interactions are controlled by a single haploid diallelic locus. Our emphasis is on the conditions for the maintenance of genetic variation, the dynamic patterns observed, the extent of local adaptation and genetic differentiation between different demes, and on how different parameters (such as the strength and heterogeneity in selection, migration rates, and the number of sites) affect the dynamic and static behavior of the system. We show that under spatially homogeneous selection the maintenance of genetic variation is possible through asynchronous nonlinear dynamics where the allele frequencies in a majority of demes quickly synchronize but the rest do not. Spatially heterogeneous selection can maintain genetic variation even if migration rates are maximal. This happens in an oscillatory way. Genetic variation is most likely to be maintained at high levels if the heterogeneity in selection is large. If there are some restrictions on migration, genetic variation can be maintained at a stable equilibrium. This behavior is most likely at intermediate migration rates. In this case, the system can exhibit high spatial subdivision as measured by F(ST) values but relatively low local adaptation.

Background: Arguably the most influential force in human history is the formation of social coalitions and alliances (i.e., long-lasting coalitions) and their impact on individual power. Understanding the dynamics of alliance formation and its consequences for biological, social, and cultural evolution is a formidable theoretical challenge. In most great ape species, coalitions occur at individual and group levels and among both kin and non-kin. Nonetheless, ape societies remain essentially hierarchical, and coalitions rarely weaken social inequality. In contrast, human hunter-gatherers show a remarkable tendency to egalitarianism, and human coalitions and alliances occur not only among individuals and groups, but also among groups of groups. These observations suggest that the evolutionary dynamics of human coalitions can only be understood in the context of social networks and cognitive evolution., Methodology/principal Findings: Here, we develop a stochastic model describing the emergence of networks of allies resulting from within-group competition for status or mates between individuals utilizing dyadic information. The model shows that alliances often emerge in a phase transition-like fashion if the group size, awareness, aggressiveness, and persuasiveness of individuals are large and the decay rate of individual affinities is small. With cultural inheritance of social networks, a single leveling alliance including all group members can emerge in several generations., Conclusions/significance: We propose a simple and flexible theoretical approach for studying the dynamics of alliance emergence applicable where game-theoretic methods are not practical. Our approach is both scalable and expandable. It is scalable in that it can be generalized to larger groups, or groups of groups. It is expandable in that it allows for inclusion of additional factors such as behavioral, genetic, social, and cultural features. Our results suggest that a rapid transition from a hierarchical society of great apes to an egalitarian society of hunter-gatherers (often referred to as "egalitarian revolution") could indeed follow an increase in human cognitive abilities. The establishment of stable group-wide egalitarian alliances creates conditions promoting the origin of cultural norms favoring the group interests over those of individuals.

We study how correlations in the random fitness assignment may affect the structure of fitness landscapes, in three classes of fitness models. The first is a phenotype space in which individuals are characterized by a large number n of continuously varying traits. In a simple model of random fitness assignment, viable phenotypes are likely to form a giant connected cluster percolating throughout the phenotype space provided the viability probability is larger than 1/2(n). The second model explicitly describes genotype-to-phenotype and phenotype-to-fitness maps, allows for neutrality at both phenotype and fitness levels, and results in a fitness landscape with tunable correlation length. Here, phenotypic neutrality and correlation between fitnesses can reduce the percolation threshold, and correlations at the point of phase transition between local and global are most conducive to the formation of the giant cluster. In the third class of models, particular combinations of alleles or values of phenotypic characters are "incompatible" in the sense that the resulting genotypes or phenotypes have zero fitness. This setting can be viewed as a generalization of the canonical Bateson-Dobzhansky-Muller model of speciation and is related to K-SAT problems, prominent in computer science. We analyze the conditions for the existence of viable genotypes, their number, as well as the structure and the number of connected clusters of viable genotypes. We show that analysis based on expected values can easily lead to wrong conclusions, especially when fitness correlations are strong. We focus on pairwise incompatibilities between diallelic loci, but we also address multiple alleles, complex incompatibilities, and continuous phenotype spaces. In the case of diallelic loci, the number of clusters is stochastically bounded and each cluster contains a very large sub-cube. Finally, we demonstrate that the discrete NK model shares some signature properties of models with high correlations.

A recent study of a pair of sympatric species of cichlids in Lake Apoyo in Nicaragua is viewed as providing probably one of the most convincing examples of sympatric speciation to date. Here, we describe and study a stochastic, individual-based, explicit genetic model tailored for this cichlid system. Our results show that relatively rapid (<20,000 generations) colonization of a new ecological niche and (sympatric or parapatric) speciation via local adaptation and divergence in habitat and mating preferences are theoretically plausible if: (i) the number of loci underlying the traits controlling local adaptation, and habitat and mating preferences is small; (ii) the strength of selection for local adaptation is intermediate; (iii) the carrying capacity of the population is intermediate; and (iv) the effects of the loci influencing nonrandom mating are strong. We discuss patterns and timescales of ecological speciation identified by our model, and we highlight important parameters and features that need to be studied empirically to provide information that can be used to improve the biological realism and power of mathematical models of ecological speciation.

A recent study of a pair of sympatric species of palms on the Lord Howe Island is viewed as providing probably one of the most convincing examples of sympatric speciation to date. Here we describe and study a stochastic, individual-based, explicit genetic model tailored for this palms system. Overall, our results show that relatively rapid (<50,000 generations) colonization of a new ecological niche, and sympatric or parapatric speciation via local adaptation and divergence in flowering periods are theoretically plausible if (i) the number of loci controlling the ecological and flowering period traits is small; (ii) the strength of selection for local adaptation is intermediate; and (iii) an acceleration of flowering by a direct environmental effect associated with the new ecological niche is present. We discuss patterns and time-scales of ecological speciation identified by our model, and we highlight important parameters and features that need to be studied empirically in order to provide information that can be used to improve the biological realism and power of mathematical models of ecological speciation.

Sexual conflict has been suggested as a general cause of genetic diversification in reproductive characters, and as a possible cause of speciation. We use individual-based simulations to study the dynamics of sexual conflict in an isolated diploid population with no spatial structure. To explore the effects of genetic details, we consider two different types of interlocus interaction between female and male traits, and three different types of intra-locus interaction. In the simulations, sexual conflict resulted in at least the following five regimes: (1) continuous coevolutionary chase, (2) evolution toward an equilibrium, (3) cyclic coevolution, (4) extensive genetic differentiation in female traits/genes only, and (5) extensive genetic differentiation in both male and female traits/genes. Genetic differentiation was hardly observed when the traits involved in reproduction were determined additively and interacted in a trait-by-trait way. When the traits interacted in a component-by-component way, genetic differentiation was frequently observed under relatively broad conditions. The likelihood of genetic differentiation largely depended on the number of loci and the type of within-locus dominance. With multiple loci per trait, genetic differentiation was often observed but sympatric speciation was typically hindered by recombination. Sympatric speciation was possible but only under restrictive conditions. Our simulations also highlight the importance of stochastic effects in the dynamics of sexual conflict.

Homosexuality is a common occurrence in humans and other species, yet its genetic and evolutionary basis is poorly understood. Here, we formulate and study a series of simple mathematical models for the purpose of predicting empirical patterns that can be used to determine the form of selection that leads to polymorphism of genes influencing homosexuality. Specifically, we develop theory to make contrasting predictions about the genetic characteristics of genes influencing homosexuality including: (i) chromosomal location, (ii) dominance among segregating alleles and (iii) effect sizes that distinguish between the two major models for their polymorphism: the overdominance and sexual antagonism models. We conclude that the measurement of the genetic characteristics of quantitative trait loci (QTLs) found in genomic screens for genes influencing homosexuality can be highly informative in resolving the form of natural selection maintaining their polymorphism.