Social learning is a critical adaptation for dealing with different forms of variability. Uncertainty is a severe form of variability where the space of possible decisions or probabilities of associated outcomes are unknown. We identified four theoretically-important sources of uncertainty: temporal environmental variability, payoff ambiguity, selection-set size, and effective lifespan. When these combine, it is nearly impossible to fully learn about the environment. We develop an evolutionary agent-based model to test how each form of uncertainty affects the evolution of social learning. Agents perform one of several behaviors, modeled as a multi-armed bandit, to acquire payoffs. All agents learn about behavioral payoffs individually through an adaptive behavior-choice model that uses a softmax decision rule. Use of vertical and oblique payoff-biased social learning evolved to serve as a scaffold for adaptive individual learning—they are not opposite strategies. Different types of uncertainty had varying effects. Temporal environmental variability suppressed social learning, whereas larger selection-set size promoted social learning, even when the environment changed frequently. Payoff ambiguity and lifespan interacted with other uncertainty parameters. This study begins to explain how social learning can predominate despite highly variable real-world environments when effective individual learning helps individuals recover from learning outdated social information.

Hidden cluster problems can manifest when broad ethnic categories are used as proxies for cultural traits, especially when traits are assumed to encode cultural distances between groups. We suggest a granular understanding of cultural trait distributions within and between ethnic categories is fundamental to the interpretation of heritability estimates as well as general behavioral outcomes.

The generalizability crisis is compounded, or even partially caused, by a lack of specificity in psychological theories. Expanding the use of mechanistic models among psychologists is therefore important, but faces numerous hurdles. A cultural evolutionary approach can help guide and evaluate interventions to improve modeling efforts in psychology, such as developing standards and implementing them at the institutional level.

We suggest that the accounts offered by the target articles could be strengthened by acknowledging the role of group selection and cultural niche construction in shaping the evolutionary trajectory of human music. We argue that group level traits and highly variable cultural niches can explain the diversity of human song, but the target articles' accounts are insufficient to explain such diversity.

Disease transmission and behaviour change are both fundamentally social phenomena. Behaviour change can have profound consequences for disease transmission, and epidemic conditions can favour the more rapid adoption of behavioural innovations. We analyse a simple model of coupled behaviour change and infection in a structured population characterised by homophily and outgroup aversion. Outgroup aversion slows the rate of adoption and can lead to lower rates of adoption in the later-adopting group or even behavioural divergence between groups when outgroup aversion exceeds positive ingroup influence. When disease dynamics are coupled to the behaviour-adoption model, a wide variety of outcomes are possible. Homophily can either increase or decrease the final size of the epidemic depending on its relative strength in the two groups and on R0 for the infection. For example, if the first group is homophilous and the second is not, the second group will have a larger epidemic. Homophily and outgroup aversion can also produce dynamics suggestive of a ‘second wave’ in the first group that follows the peak of the epidemic in the second group. Our simple model reveals dynamics that are suggestive of the processes currently observed under pandemic conditions in culturally and/or politically polarised populations such as the USA.

We support the goal to integrate models of culture and cognition. However, we are not convinced that the free energy principle and Thinking Through Other Minds will be useful in achieving it. There are long traditions of modeling both cultural evolution and cognition. Demonstrating that FEP or TTOM can integrate these models will require a bit more math.

Heyes’ book is an important contribution that rightly integrates cognitive development and cultural evolution. However, understanding the cultural evolution of cognitive gadgets requires a deeper appreciation of complexity, feedback, and self-organization than her book exhibits.

We examine a model of network formation in single-layer and multiplex networks in which individuals have positive incentives for social ties, closed triangles, and spillover edges. In particular, we investigate the influence of shocks to the network in which the cost of social ties changes after an initial equilibrium. We highlight the emergence of structural entrenchment: the retention of structural features, such as closed triangles and spillover edges, which are formed under historically different conditions from those currently driving network evolution. This work has broad implications for understanding path dependence in the structure and dynamics of single-layer and multiplex networks.

The target article is plagued by imprecision, making it largely impossible to evaluate the authors' theory in a scientific manner.

The main objective of our target article was to sketch the empirical case for the importance of selection at the level of groups on cultural variation. Such variation is massive in humans, but modest or absent in other species. Group selection processes acting on this variation is a framework for developing explanations of the unusual level of cooperation between non-relatives found in our species. Our case for cultural group selection (CGS) followed Darwin's classic syllogism regarding natural selection: If variation exists at the level of groups, if this variation is heritable, and if it plays a role in the success or failure of competing groups, then selection will operate at the level of groups. We outlined the relevant domains where such evidence can be sought and characterized the main conclusions of work in those domains. Most commentators agree that CGS plays some role in human evolution, although some were considerably more skeptical. Some contributed additional empirical cases. Some raised issues of the scope of CGS explanations versus competing ones.

We appreciate and endorse Kline's ethological taxonomy and its application. However, the definition of teaching she presents is problematic, as it replaces mentalistic intent with intention on the part of natural selection. We discuss problems with the strict adaptationist view and suggest instead that the five forms of teaching presented in the taxonomy may constitute exaptations rather than adaptations.

Human cooperation is highly unusual. We live in large groups composed mostly of non-relatives. Evolutionists have proposed a number of explanations for this pattern, including cultural group selection and extensions of more general processes such as reciprocity, kin selection, and multi-level selection acting on genes. Evolutionary processes are consilient; they affect several different empirical domains, such as patterns of behavior and the proximal drivers of that behavior. In this target article, we sketch the evidence from five domains that bear on the explanatory adequacy of cultural group selection and competing hypotheses to explain human cooperation. Does cultural transmission constitute an inheritance system that can evolve in a Darwinian fashion? Are the norms that underpin institutions among the cultural traits so transmitted? Do we observe sufficient variation at the level of groups of considerable size for group selection to be a plausible process? Do human groups compete, and do success and failure in competition depend upon cultural variation? Do we observe adaptations for cooperation in humans that most plausibly arose by cultural group selection? If the answer to one of these questions is “no,” then we must look to other hypotheses. We present evidence, including quantitative evidence, that the answer to all of the questions is “yes” and argue that we must take the cultural group selection hypothesis seriously. If culturally transmitted systems of rules (institutions) that limit individual deviance organize cooperation in human societies, then it is not clear that any extant alternative to cultural group selection can be a complete explanation.

We agree that evolutionary perspectives may help us organize many divergent realms of the science of human behavior. Nevertheless, an imperative to unite all social science under an evolutionary framework risks turning off researchers who have their own theoretical perspectives that can be informed by evolutionary theory without being exclusively defined by it. We propose a few considerations for scholars interested in joining the evolutionary and social sciences.

Most commentators supported the thesis of the target article, though there were also those who were less fully persuaded. I will begin with a response to the most critical commentaries. First, I will justify an evolutionary perspective that includes group organization and nongenetic inheritance. Next, I will discuss the concept of emergence. Following that, I will transition to an exploration of ideas and concerns brought up by some of the more supportive commentators. This will include a discussion of different types of groups; the psychology of group-level traits; the uses and limitations of an institutional perspective; the link between transmission, adaptation, and selection; current and future methodologies; and the variety of fields that may benefit from a group-level traits perspective.

Many of the most important properties of human groups – including properties that may give one group an evolutionary advantage over another – are properly defined only at the level of group organization. Yet at present, most work on the evolution of culture has focused solely on the transmission of individual-level traits. I propose a conceptual extension of the theory of cultural evolution, particularly related to the evolutionary competition between cultural groups. The key concept in this extension is the emergent group-level trait. This type of trait is characterized by the structured organization of differentiated individuals and constitutes a unit of selection that is qualitatively different from selection on groups as defined by traditional multilevel selection (MLS) theory. As a corollary, I argue that the traditional focus on cooperation as the defining feature of human societies has missed an essential feature of cooperative groups. Traditional models of cooperation assume that interacting with one cooperator is equivalent to interacting with any other. However, human groups involve differential roles, meaning that receiving aid from one individual is often preferred to receiving aid from another. In this target article, I discuss the emergence and evolution of group-level traits and the implications for the theory of cultural evolution, including ramifications for the evolution of human cooperation, technology, and cultural institutions, and for the equivalency of multilevel selection and inclusive fitness approaches.

Because of the complexity of human emotional responses, invariants must be sought not in the responses themselves, but in their generating mechanisms. Lindquist et al. show that functional locationism is a theoretical dead end; their proposed mechanistic framework is a first step toward better models of emotional behavior. We caution, however, that emotions may still be quasi-natural perceptual types.