Evolution of culture-dependent discriminate sociality: a gene-culture coevolutionary model

Animals behave cooperatively towards certain conspecifics while being indifferent or even hostile to others. The distinction is made primarily according to kinship as predicted by the kin selection theory. With regards to humans, however, this is not always the case; in particular, humans sometimes exhibit a discriminate sociality on the basis of culturally transmitted traits, such as personal ornaments, languages, rituals, etc. This paper explores the possibility that the human faculty of cultural transmission and resultant cultural variation among individuals may have facilitated the evolution of discriminate sociality in humans. To this end, a gene-culture coevolutionary model is developed focusing on competition over control of resource as a context in which discriminate sociality may have evolved. Specifically, two types of culture-dependent discriminate sociality are considered: ingroup favouritism, with ingroup and outgroup being distinguished by the presence or absence of a cultural trait; and prestige hierarchies, with the prestige being conferred on the bearer of a cultural trait. The model specifies the conditions under which emergence and evolutionary stability of the two types of discriminate sociality are promoted by the presence of cultural variation among individuals.     

An evolutionary theory of large‐scale human warfare: Group‐structured cultural selection

When humans wage war, it is not unusual for battlefields to be strewn with dead warriors. These warriors typically were men in their reproductive prime who, had they not died in battle, might have gone on to father more children. Typically, they are also genetically unrelated to one another. We know of no other animal species in which reproductively capable, genetically unrelated individuals risk their lives in this manner. Because the immense private costs borne by individual warriors create benefits that are shared widely by others in their group, warfare is a stark evolutionary puzzle that is difficult to explain. Although several scholars have posited models of the evolution of human warfare, 1 - 6 these models do not adequately explain how humans solve the problem of collective action in warfare at the evolutionarily novel scale of hundreds of genetically unrelated individuals. We propose that group‐structured cultural selection explains this phenomenon.     

An ancient evolutionary origin of genes associated with human genetic diseases

Several thousand genes in the human genome have been linked to a heritable genetic disease. The majority of these appear to be nonessential genes (i.e., are not embryonically lethal when inactivated), and one could therefore speculate that they are late additions in the evolutionary lineage toward humans. Contrary to this expectation, we find that they are in fact significantly overrepresented among the genes that have emerged during the early evolution of the metazoa. Using a phylostratigraphic approach, we have studied the evolutionary emergence of such genes at 19 phylogenetic levels. The majority of disease genes was already present in the eukaryotic ancestor, and the second largest number has arisen around the time of evolution of multicellularity. Conversely, genes specific to the mammalian lineage are highly underrepresented. Hence, genes involved in genetic diseases are not simply a random subset of all genes in the genome but are biased toward ancient genes.     

The evolution of intergroup tolerance in nonhuman primates and humans

Primate individuals use a variety of strategies in intergroup encounters, from aggression to tolerance; however, recent focus on the evolution of either warfare or peace has come at the cost of characterizing this variability. We identify evolutionary advantages that may incentivize tolerance toward extra‐group individuals in humans and nonhuman primates, including enhanced benefits in the domains of transfer, mating, and food acquisition. We highlight the role these factors play in the flexibility of gorilla, chimpanzee, bonobo, and human behavior. Given humans have an especially broad range of intergroup behavior, we explore how the human foraging ecology, especially large spatial and temporal fluctuations in resource availability, may have selected for a greater reliance on tolerant between‐community relationships—relationships reinforced by status acquisition and cultural institutions. We conclude by urging careful, theoretically motivated study of behavioral flexibility in intergroup encounters in humans and the nonhuman great apes.     

Cultural and reproductive success and the causes of war: A Yanomamö perspective

Inter-group competition including warfare is posited to be a key force in human evolution (Alexander, 1990; Choi & Bowles, 2007; Wrangham, 1999). Chagnon's research on the Yanomamö is seminal to understanding warfare in the types of societies characteristic of human evolutionary history. Chagnon's empirical analyses of the hypothesis that competition for status or cultural success is linked to reproduction (Irons, 1979) and warfare attracted considerable controversy. Potential causal factors include “blood revenge”, mate competition, resource shortages or inequality, and peace-making institutions (Boehm, 1984; Keeley's (1997); Meggitt, 1977; Wiessner and Pupu, 2012; Wrangham et al., 2006). Here we highlight Chagnon's contributions to the study of human warfare.     

Mode and tempo in the evolution of socio-political organization: reconciling 'Darwinian' and 'Spencerian' evolutionary approaches in anthropology

Traditional investigations of the evolution of human social and political institutions trace their ancestry back to nineteenth century social scientists such as Herbert Spencer, and have concentrated on the increase in socio-political complexity over time. More recent studies of cultural evolution have been explicitly informed by Darwinian evolutionary theory and focus on the transmission of cultural traits between individuals. These two approaches to investigating cultural change are often seen as incompatible. However, we argue that many of the defining features and assumptions of 'Spencerian' cultural evolutionary theory represent testable hypotheses that can and should be tackled within a broader 'Darwinian' framework. In this paper we apply phylogenetic comparative techniques to data from Austronesian-speaking societies of Island South-East Asia and the Pacific to test hypotheses about the mode and tempo of human socio-political evolution. We find support for three ideas often associated with Spencerian cultural evolutionary theory: (i) political organization has evolved through a regular sequence of forms, (ii) increases in hierarchical political complexity have been more common than decreases, and (iii) political organization has co-evolved with the wider presence of hereditary social stratification.     

Neutralism and selectionism: a network-based reconciliation

Neutralism and selectionism are extremes of an explanatory spectrum for understanding patterns of molecular evolution and the emergence of evolutionary innovation. Although recent genome-scale data from protein-coding genes argue against neutralism, molecular engineering and protein evolution data argue that neutral mutations and mutational robustness are important for evolutionary innovation. Here I propose a reconciliation in which neutral mutations prepare the ground for later evolutionary adaptation. Key to this perspective is an explicit understanding of molecular phenotypes that has only become accessible in recent years.     

Naming a structured world: a cultural route to duality of patterning

The lexicons of human languages organize their units at two distinct levels. At a first combinatorial level, meaningless forms (typically referred to as phonemes) are combined into meaningful units (typically referred to as morphemes). Thanks to this, many morphemes can be obtained by relatively simple combinations of a small number of phonemes. At a second compositional level of the lexicon, morphemes are composed into larger lexical units, the meaning of which is related to the individual meanings of the composing morphemes. This duality of patterning is not a necessity for lexicons and the question remains wide open regarding how a population of individuals is able to bootstrap such a structure and the evolutionary advantages of its emergence. Here we address this question in the framework of a multi-agents model, where a population of individuals plays simple naming games in a conceptual environment modeled as a graph. We demonstrate that errors in communication as well as a blending repair strategy, which crucially exploits a shared conceptual representation of the environment, are sufficient conditions for the emergence of duality of patterning, that can thus be explained in a pure cultural way. Compositional lexicons turn out to be faster to lead to successful communication than purely combinatorial lexicons, suggesting that meaning played a crucial role in the evolution of language.     

Evolution of equal division among unequal partners

One of the hallmarks of human fairness is its insensitivity to power: although strong individuals are often in a position to coerce weak individuals, fairness requires them to share the benefits of cooperation equally. The existence of such egalitarianism is poorly explained by current evolutionary models. We present a model based on cooperation and partner choice that can account for the emergence of a psychological disposition toward fairness, whatever the balance of power between the cooperative partners. We model the evolution of the division of a benefit in an interaction similar to an ultimatum game, in a population made up of individuals of variable strength. The model shows that strong individuals will not receive any advantage from their strength, instead having to share the benefits of cooperation equally with weak individuals at the evolutionary equilibrium, a result that is robust to variations in population size and the proportion of weak individuals. We discuss how this model suggests an explanation for why egalitarian behaviors toward everyone, including the weak, should be more likely to evolve in humans than in any other species.     

The cultural contagion of conflict

Anecdotal evidence abounds that conflicts between two individuals can spread across networks to involve a multitude of others. We advance a cultural transmission model of intergroup conflict where conflict contagion is seen as a consequence of universal human traits (ingroup preference, outgroup hostility; i.e. parochial altruism) which give their strongest expression in particular cultural contexts. Qualitative interviews conducted in the Middle East, USA and Canada suggest that parochial altruism processes vary across cultural groups and are most likely to occur in collectivistic cultural contexts that have high ingroup loyalty. Implications for future neuroscience and computational research needed to understand the emergence of intergroup conflict are discussed.     

A History of Research on Warfare in Anthropology

In this brief history of warfare research, I identify four major periods: Foundation Period, Classical Period, Golden Age, and Recent Period. The myth of the peaceful savage arose in the Classical Period rather than in the later periods, as argued by Lawrence Keeley. The myth has its origins in an early evolutionary approach as well as in cultural relativism. It continues to play a major role in warfare research with some anthropologists arguing for the peaceable nature of man, while others argue that man has engaged in warfare from the beginning of prehistory, [ war, theories of warfare, history of warfare research, myth of peaceful savage ]     

Cultural transmission and the evolution of human behaviour: a general approach based on the Price equation

Transmitted culture can be viewed as an inheritance system somewhat independent of genes that is subject to processes of descent with modification in its own right. Although many authors have conceptualized cultural change as a Darwinian process, there is no generally agreed formal framework for defining key concepts such as natural selection, fitness, relatedness and altruism for the cultural case. Here, we present and explore such a framework using the Price equation. Assuming an isolated, independently measurable culturally transmitted trait, we show that cultural natural selection maximizes cultural fitness, a distinct quantity from genetic fitness, and also that cultural relatedness and cultural altruism are not reducible to or necessarily related to their genetic counterparts. We show that antagonistic coevolution will occur between genes and culture whenever cultural fitness is not perfectly aligned with genetic fitness, as genetic selection will shape psychological mechanisms to avoid susceptibility to cultural traits that bear a genetic fitness cost. We discuss the difficulties with conceptualizing cultural change using the framework of evolutionary theory, the degree to which cultural evolution is autonomous from genetic evolution, and the extent to which cultural change should be seen as a Darwinian process. We argue that the nonselection components of evolutionary change are much more important for culture than for genes, and that this and other important differences from the genetic case mean that different approaches and emphases are needed for cultural than genetic processes.     

Larger body size on islands affects silvereye Zosterops lateralis song and call frequency

Differences in song repertoires and characteristics of island and mainland populations of the same avian species are usually explained by dispersal, cultural evolution and/or habitat differences. The influence of morphology is often overlooked, even though island populations are frequently morphologically distinct from mainland populations, and morphology could affect vocalizations. I compared morphological features, songs, contact calls and alarm calls of six isolated island populations of silvereye Zosterops lateralis with those of two mainland populations to examine whether differences between mainland and island vocalizations were consistent across vocalization types, and whether these differences could be linked to morphological differences. Vocalizations were lower in frequency on islands. Island individuals were larger (both in mass and body structure), and body mass was an important predictor of frequency in contact and alarm calls. I argue that this strong association results from the island rule (islands promote larger body sizes) and cascading effects of morphology on vocalization frequency in this species.     

The Hybrid Origin of “Modern” Humans

Recent genomic research has shown that hybridization between substantially diverged lineages is the rule, not the exception, in human evolution. However, the importance of hybridization in shaping the genotype and phenotype of Homo sapiens remains debated. Here we argue that current evidence for hybridization in human evolution suggests not only that it was important, but that it was an essential creative force in the emergence of our variable, adaptable species. We then extend this argument to a reappraisal of the archaeological record, proposing that the exchange of cultural information between divergent groups may have facilitated the emergence of cultural innovation. We discuss the implications of this Divergence and Hybridization Model for considering the taxonomy of our lineage.     

A major evolutionary transition to more than two sexes?

Two recently discovered cases of genetic caste determination in social insects might provide the first example of a major evolutionary transition from two to more than two sexes. I argue here that the system can be interpreted as comprising primarily individuals requiring gametes from three parental types and having four sexes from the perspective of demographic extinction. Additionally, I show how this mating system can be seen as a major evolutionary transition. For these populations, it is apparent that the mechanism for a three- or four-sex system does not lie within the myriad of possible arrangements of chromosomes within individuals, but at the next level of evolutionary complexity, with the arrangement of chromosomes among individuals within a social system.     

Simulating the Evolution of the Human Family: Cooperative Breeding Increases in Harsh Environments

Verbal and mathematical models that consider the costs and benefits of behavioral strategies have been useful in explaining animal behavior and are often used as the basis of evolutionary explanations of human behavior. In most cases, however, these models do not account for the effects that group structure and cultural traditions within a human population have on the costs and benefits of its members'' decisions. Nor do they consider the likelihood that cultural as well as genetic traits will be subject to natural selection. In this paper, we present an agent-based model that incorporates some key aspects of human social structure and life history. We investigate the evolution of a population under conditions of different environmental harshness and in which selection can occur at the level of the group as well as the level of the individual. We focus on the evolution of a socially learned characteristic related to individuals'' willingness to contribute to raising the offspring of others within their family group. We find that environmental harshness increases the frequency of individuals who make such contributions. However, under the conditions we stipulate, we also find that environmental variability can allow groups to survive with lower frequencies of helpers. The model presented here is inevitably a simplified representation of a human population, but it provides a basis for future modeling work toward evolutionary explanations of human behavior that consider the influence of both genetic and cultural transmission of behavior.     

Culture and sociobiology

Sociobiological concepts are easily misapplied to human behavior because the latter is culturally as well as biologically organized. Because biological and cultural evolution are two linked but conceptually distinct processes, sociobiology is more readily applied to the evolution of cultural capacity than to contemporary cultural behavior. The extent to which the latter is consistent with sociobiological expectation must be determined empirically, although there are theoretical grounds for predicting a limited degree of concordance . [sociobiology, culture, evolution, reductionism, biosocial anthropology]     

Evolution: like any other science it is predictable

Evolutionary biology rejoices in the diversity of life, but this comes at a cost: other than working in the common framework of neo-Darwinian evolution, specialists in, for example, diatoms and mammals have little to say to each other. Accordingly, their research tends to track the particularities and peculiarities of a given group and seldom enquires whether there are any wider or deeper sets of explanations. Here, I present evidence in support of the heterodox idea that evolution might look to a general theory that does more than serve as a tautology (‘evolution explains evolution’). Specifically, I argue that far from its myriad of products being fortuitous and accidental, evolution is remarkably predictable. Thus, I urge a move away from the continuing obsession with Darwinian mechanisms, which are entirely uncontroversial. Rather, I emphasize why we should seek explanations for ubiquitous evolutionary convergence, as well as the emergence of complex integrated systems. At present, evolutionary theory seems to be akin to nineteenth-century physics, blissfully unaware of the imminent arrival of quantum mechanics and general relativity. Physics had its Newton, biology its Darwin: evolutionary biology now awaits its Einstein.     

Dynamics of alliance formation and the egalitarian revolution

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.     

Pathogen threat and intergroup prejudice using the minimal group paradigm: Evidence from a registered report

Infection by parasites, bacteria, and other microorganisms has been a powerful selection pressure faced by humans and other species. Consequently, avoiding pathogens has played an important role in human evolution and continues to play a role in contemporary social psychological processes. The current research tested the hypothesis that pathogen avoidance promotes intergroup prejudice. Whereas previous tests relied on existing cultural groups, which can conflate outgroup status with pre-existing group stereotypes about disease or geographic variability in pathogen prevalence, the current experiments assessed intergroup bias using a minimal group paradigm. Based on preliminary evidence (Study 1, N = 207, undergraduate students) that experimentally priming pathogen avoidance motivation promoted negativity toward a minimal outgroup, we conducted a Registered Report (Study 2, N = 1339 online participants) to replicate and extend those findings. Our primary hypothesis was that an experimental manipulation of pathogen threat (relative to two control conditions) would produce greater intergroup prejudice (negativity toward the nominal outgroup relative to the nominal ingroup). This hypothesis was not supported in the larger, registered second study. Exploratory analyses provided some evidence for interactions between experimental priming of pathogen threat and individual differences in pathogen disgust, but the interactive pattern differed across the two experiments. Findings call into question the hypothesis that, in the absence of cultural stereotypes, situationally activated pathogen avoidance promotes intergroup prejudice.