The cophylogeny of populations and cultures: reconstructing the evolution of Iranian tribal craft traditions using trees and jungles

Phylogenetic approaches to culture have shed new light on the role played by population dispersals in the spread and diversification of cultural traditions. However, the fact that cultural inheritance is based on separate mechanisms from genetic inheritance means that socially transmitted traditions have the potential to diverge from population histories. Here, we suggest that associations between these two systems can be reconstructed using techniques developed to study cospeciation between hosts and parasites and related problems in biology. Relationships among the latter are patterned by four main processes: co-divergence, intra-host speciation (duplication), intra-host extinction (sorting) and horizontal transfers. We show that patterns of cultural inheritance are structured by analogous processes, and then demonstrate the applicability of the host-parasite model to culture using empirical data on Iranian tribal populations.     

Human niche construction in interdisciplinary focus

Niche construction is an endogenous causal process in evolution, reciprocal to the causal process of natural selection. It works by adding ecological inheritance, comprising the inheritance of natural selection pressures previously modified by niche construction, to genetic inheritance in evolution. Human niche construction modifies selection pressures in environments in ways that affect both human evolution, and the evolution of other species. Human ecological inheritance is exceptionally potent because it includes the social transmission and inheritance of cultural knowledge, and material culture. Human genetic inheritance in combination with human cultural inheritance thus provides a basis for gene-culture coevolution, and multivariate dynamics in cultural evolution. Niche construction theory potentially integrates the biological and social aspects of the human sciences. We elaborate on these processes, and provide brief introductions to each of the papers published in this theme issue.     

A model for the evolutionary diversification of religions

We address the problem of cultural diversification by studying selection on cultural ideas that colonize human hosts and using diversification of religions as a conceptual example. In analogy to studying the evolution of pathogens or symbionts colonizing animal hosts, we use models for host-pathogen dynamics known from theoretical epidemiology. In these models, religious content colonizes individual humans. Rates of transmission of ideas between humans, i.e., transmission of cultural content, and rates of loss of ideas (loss of belief) are determined by the phenotype of the cultural content, and by interactions between hosts carrying different ideas. In particular, based on the notion that cultural non-conformism can be negative frequency-dependent (for example, religion can lead to oppression of lower classes and emergence of non-conformism and dissent once a religious belief has reached dominance), we assume that the rate of loss of belief increases as the number of humans colonized by a particular religious phenotype increases. This generates frequency-dependent selection on cultural content, and we use evolutionary theory to show that this frequency dependence can lead to the emergence of coexisting clusters of different cultural types. The different clusters correspond to different cultural traditions, and hence our model describes the emergence of distinct descendant cultures from a single ancestral culture in the absence of any geographical isolation.     

Not in their genes: Phenotypic flexibility, behavioural traditions and cultural evolution in wild bonnet macaques

Phenotypic flexibility, or the within-genotype, context-dependent, variation in behaviour expressed by single reproductively mature individuals during their lifetimes, often impart a selective advantage to organisms and profoundly influence their survival and reproduction. Another phenomenon apparently not under direct genetic control is behavioural inheritance whereby higher animals are able to acquire information from the behaviour of others by social learning, and, through their own modified behaviour, transmit such information between individuals and across generations. Behavioural information transfer of this nature thus represents another form of inheritance that operates in many animals in tandem with the more basic genetic system. This paper examines the impact that phenotypic flexibility, behavioural inheritance and socially transmitted cultural traditions may have in shaping the structure and dynamics of a primate society — that of the bonnet macaque (Macaca radiata), a primate species endemic to peninsular India. Three principal issues are considered: the role of phenotypic flexibility in shaping social behaviour, the occurrence of individual behavioural traits leading to the establishment of social traditions, and the appearance of cultural evolution amidst such social traditions. Although more prolonged observations are required, these initial findings suggest that phenotypic plasticity, behavioural inheritance and cultural traditions may be much more widespread among primates than have previously been assumed but may have escaped attention due to a preoccupation with genetic inheritance in zoological thinking. This paper is dedicated with admiration and gratitude to the late Dr Raja Ramanna, a man who held all primates in great esteem.     

Characterizing symbiont inheritance during host–microbiota evolution: Application to the great apes gut microbiota

Microbiota play a central role in the functioning of multicellular life, yet understanding their inheritance during host evolutionary history remains an important challenge. Symbiotic microorganisms are either acquired from the environment during the life of the host (i.e. environmental acquisition), transmitted across generations with a faithful association with their hosts (i.e. strict vertical transmission), or transmitted with occasional host switches (i.e. vertical transmission with horizontal switches). These different modes of inheritance affect microbes’ diversification, which at the two extremes can be independent from that of their associated host or follow host diversification. The few existing quantitative tools for investigating the inheritance of symbiotic organisms rely on cophylogenetic approaches, which require knowledge of both host and symbiont phylogenies, and are therefore often not well adapted to DNA metabarcoding microbial data. Here, we develop a model‐based framework for identifying vertically transmitted microbial taxa. We consider a model for the evolution of microbial sequences on a fixed host phylogeny that includes vertical transmission and horizontal host switches. This model allows estimating the number of host switches and testing for strict vertical transmission and independent evolution. We test our approach using simulations. Finally, we illustrate our framework on gut microbiota high‐throughput sequencing data of the family Hominidae and identify several microbial taxonomic units, including fibrolytic bacteria involved in carbohydrate digestion, that tend to be vertically transmitted.     

Studying Cultural Evolution at the Tips: Human Cross-cultural Ecology

While human genetic variation is limited due to a bottleneck on the origin of the species ～200 kya, cultural traits can change more rapidly, and may do so in response to the variation in human habitats. Does cultural diversification simulate a natural experiment in evolution much like biodiversity so that cultural divergences and convergences can be interpreted in terms of the differences and similarities of local environments? Or is cultural diversity simply the result of human behavioral flexibility? Although the majority of cultural data comes from the tips of the hominin phylogeny, anthropologists can follow the example of evolutionary ecologists, who often compare the endpoints of phylogenies when that is all that is available. This article compares 97 contemporary indigenous language communities from around the world, and 24 of their cultural traditions, to help determine whether human cultures and their cultural traits are proportionately dispersed, as predicted by the neutral theory of biodiversity, or whether they show non-proportionalities that could be explained with evolutionary reasoning.     

Consanguineous marriages in Italy: A Note

Abstract

A general multifactorial model is given for the inheritance of traits that exhibit a sexual dimorphism. The model allows for polygenic inheritance, cultural transmission, phenotypic assortative mating, and a common environment of rearing. Several cultural mechanisms are described for which transmission from parent to offspring is sex‐dependent and for which many different patterns of sex‐specific correlations can result. A special case of the general model is described in which phenotypic differences between males and females are due only to differences in nontransmissible environmental factors and/or genetic factors that do not contribute to variability within a sex. Application of these models to human spatial visualizing ability, using data reported by others, gives an estimate of 45 per cent for the proportion of the variance that is accounted for by transmissible factors. Neither an X‐linked hypothesis nor a sex‐specific cultural mechanism is required to explain the transmission of spatial ability.     

Learning bias, cultural evolution of language, and the biological evolution of the language faculty

The biases of individual language learners act to determine the learnability and cultural stability of languages: learners come to the language learning task with biases which make certain linguistic systems easier to acquire than others. These biases are repeatedly applied during the process of language transmission, and consequently should effect the types of languages we see in human populations. Understanding the cultural evolutionary consequences of particular learning biases is therefore central to understanding the link between language learning in individuals and language universals, common structural properties shared by all the world’s languages. This paper reviews a range of models and experimental studies which show that weak biases in individual learners can have strong effects on the structure of socially learned systems such as language, suggesting that strong universal tendencies in language structure do not require us to postulate strong underlying biases or constraints on language learning. Furthermore, understanding the relationship between learner biases and language design has implications for theories of the evolution of those learning biases: models of gene-culture coevolution suggest that, in situations where a cultural dynamic mediates between properties of individual learners and properties of language in this way, biological evolution is unlikely to lead to the emergence of strong constraints on learning.     

Assortative mating and grandparental transmission facilitate the persistence of a sign language.

Conditions for the persistence (i.e., protection from loss) of a sign language are investigated assuming monogenic recessive inheritance of deafness, assortative mating for deafness or hearing, and cultural transmission of the sign language to deaf individuals from their deaf parents and deaf maternal grandparents. A new method is introduced to deal with the problem of grandparental transmission in which the basic variables are the frequencies of triplets comprising a mother, a father, and their daughter of permissible phenogenotypes. Usual stability analysis is then done on the system of linear recursions in the frequencies of these triplets, derived on the assumption that signers (users of the sign language) are rare. It is shown that assortative mating is the most important factor contributing to persistence, but that grandparental transmission can also have a significant effect when assortment is as strong as observed in England and the United States.     

Modeling effects of local extinctions on culture change and diversity in the paleolithic

The persistence of early stone tool technologies has puzzled archaeologists for decades. Cognitively based explanations, which presume either lack of ability to innovate or extreme conformism, do not account for the totality of the empirical patterns. Following recent research, this study explores the effects of demographic factors on rates of culture change and diversification. We investigate whether the appearance of stability in early Paleolithic technologies could result from frequent extinctions of local subpopulations within a persistent metapopulation. A spatially explicit agent-based model was constructed to test the influence of local extinction rate on three general cultural patterns that archaeologists might observe in the material record: total diversity, differentiation among spatially defined groups, and the rate of cumulative change. The model shows that diversity, differentiation, and the rate of cumulative cultural change would be strongly affected by local extinction rates, in some cases mimicking the results of conformist cultural transmission. The results have implications for understanding spatial and temporal patterning in ancient material culture.     

Descent with modification and the archaeological record

Recent years have seen major advances in our understanding of the way in which cultural transmission takes place and the factors that affect it. The theoretical foundations of those advances have been built by postulating the existence of a variety of different processes and deriving their consequences mathematically or by simulation. The operation of these processes in the real world can be studied through experiment and naturalistic observation. In contrast, archaeologists have an 'inverse problem'. For them the object of study is the residues of different behaviours represented by the archaeological record and the problem is to infer the microscale processes that produced them, a vital task for cultural evolution since this is the only direct record of past cultural patterns. The situation is analogous to that faced by population geneticists scanning large number of genes and looking for evidence of selection as opposed to drift, but more complicated for many reasons, not least the enormous variety of different forces that affect cultural transmission. This paper reviews the progress that has been made in inferring processes from patterns and the role of demography in those processes, together with the problems that have arisen.     

Stone toolmaking and the evolution of human culture and cognition

Although many species display behavioural traditions, human culture is unique in the complexity of its technological, symbolic and social contents. Is this extraordinary complexity a product of cognitive evolution, cultural evolution or some interaction of the two? Answering this question will require a much better understanding of patterns of increasing cultural diversity, complexity and rates of change in human evolution. Palaeolithic stone tools provide a relatively abundant and continuous record of such change, but a systematic method for describing the complexity and diversity of these early technologies has yet to be developed. Here, an initial attempt at such a system is presented. Results suggest that rates of Palaeolithic culture change may have been underestimated and that there is a direct relationship between increasing technological complexity and diversity. Cognitive evolution and the greater latitude for cultural variation afforded by increasingly complex technologies may play complementary roles in explaining this pattern.     

Cultural transmission can inhibit the evolution of altruistic helping

The study of culturally inherited traits has led to the suggestion that the evolution of helping behaviors is more likely with cultural transmission than without. Here we evaluate this idea through a comparative analysis of selection on helping under both genetic and cultural inheritance. We develop two simple models for the evolution of helping through cultural group selection: one in which selection on the trait depends solely on Darwinian fitness effects and one in which selection is driven by nonreproductive factors, specifically imitation of strategies achieving higher payoffs. We show that when cultural variants affect Darwinian fitness, the selection pressure on helping can be markedly increased relative to that under genetic transmission. By contrast, when variants are driven by nonreproductive factors, the selection pressure on helping may be reduced relative to that under genetic inheritance. This occurs because, unlike biological offspring, the spread of cultural variants from one group to another through imitation does not reduce the number of these variants in the source group. As a consequence, there is increased within-group competition associated with traits increasing group productivity, which reduces the benefits of helping. In these cases, selection for harming behavior (decreasing the payoff to neighbors) may occur rather than selection for helping.     

Bridging Taxonomic and Disciplinary Divides in Infectious Disease

Pathogens traverse disciplinary and taxonomic boundaries, yet infectious disease research occurs in many separate disciplines including plant pathology, veterinary and human medicine, and ecological and evolutionary sciences. These disciplines have different traditions, goals, and terminology, creating gaps in communication. Bridging these disciplinary and taxonomic gaps promises novel insights and important synergistic advances in control of infectious disease. An approach integrated across the plant-animal divide would advance our understanding of disease by quantifying critical processes including transmission, community interactions, pathogen evolution, and complexity at multiple spatial and temporal scales. These advances require more substantial investment in basic disease research.     

Cultural panthropology

Culture, in the most basic sense of “tradition,” has been shown to exist in many species. There is more to the phenomenon of culture in humans, however, than the mere existence of traditions. Thus, rather than expecting that culture can be assigned to living or ancestral species in an all‐or‐none fashion, reconstruction of the evolution of this uniquely complex phenomenon is likely to depend on successfully teasing apart its components, which may have evolved in a somewhat mosaic fashion. In this paper, we dissect ten different aspects of human culture and offer evidence that most of them are manifested in chimpanzees, even if in limited ways, permitting inferences about the cultural profile of our common ancestor. The aspects of culture examined include large‐scale patterns of behavioral variation across populations, the mechanisms available for social transmission, and cultural contents. The contrasts thus drawn for humans and chimpanzees offer a framework for cultural comparisons between other taxa from the past and present.     

The effect of cultural transmission on continuous variation.

Cultural transmission may depend on the non-genetic transfer of information from parent to offspring. The consequences of such cultural transmission for continuous variation are investigated theoretically for randomly mating populations. Cultural inheritance may act on genetical and environmental differences between individuals. The consequences for cultural inheritance of polygenic variation and variation due to chance environmental factors are considered. An equilibrium may occur in which the population variance and the covariances between relatives can be expressed as functions of estimable parameters of genetical and environmental variation. Whatever the ultimate origin of culturally inherited differences they are expected to lead to environmental differences between families ("E2" variation). In addition, if cultural transmission maintains differences due ultimately to segregation at many gene loci we may find genotype-environmental covariation is generated.     

Inclusive heritability: combining genetic and non‐genetic information to study animal behavior and culture

Phenotypic variance results from variation in biological information possessed by individuals. Quantitative geneticists often strive to partition out all environmental variance to measure heritability. Behavioral biologists and ecologists however, require methods to integrate genetic and environmental components of inherited phenotypic variance in order to estimate the evolutionary potential of traits, which encompasses any form of information that is inherited. To help develop this integration, we build on the tools of quantitative genetics and offer the concept of ‘inclusive heritability’ which identifies and unifies the various mechanisms of information transmission across generations. A controversial component of non‐genetic information is animal culture, which is the part of phenotypic variance inherited through social learning. Culture has the unique property of being transmitted horizontally and obliquely, as well as vertically. Accounting for cultural variation would allow us to examine a broader range of evolutionary mechanisms. Culture may, for instance, produce behavioral isolating mechanisms leading to speciation. To advance the study of animal culture, we offer a definition of culture that is rooted in quantitative genetics. We also offer four testable criteria to determine whether a trait is culturally inherited. These criteria may constitute a conceptual tool to study animal culture. We briefly discuss methods to partition out cultural variance. Several authors have recently called for ‘modernizing the modern synthesis’ by including non‐genetic factors such as epigenetics and phenotypic plasticity in order to more fully explain phenotypic evolution. Here, we further propose to broaden the concept of inheritance by incorporating the cultural component of behavior. Applying the concept of inclusive heritability may advance the integration of multiple forms of inheritance into the study of evolution.     

The extended evolutionary synthesis and the role of soft inheritance in evolution

In recent years, a number of researchers have advocated extending the modern synthesis in evolutionary biology. One of the core arguments made in favour of an extension comes from work on soft inheritance systems, including transgenerational epigenetic effects, cultural transmission and niche construction. In this study, we outline this claim and then take issue with it. We argue that the focus on soft inheritance has led to a conflation of proximate and ultimate causation, which has in turn obscured key questions about biological organization and calibration across the life span to maximize average lifetime inclusive fitness. We illustrate this by presenting hypotheses that we believe incorporate the core phenomena of soft inheritance and will aid in understanding them.