Third-party arbitration and forgiving strategies increase cooperation when perception errors are common.

Humans cooperate in groups in which mutual monitoring is common, and this provides the possibility of third-party arbitration. Third-party arbitration stabilizes reciprocity in at least two ways: first, when it is accurate, it reduces the frequency of misunderstandings resulting from perception errors, and second, even when it is inaccurate, it provides a public signal that allows pairs to align their expectations about how to behave after errors occur. Here, we describe experiments that test for these two effects. We find that in an iterated, sequential Prisoner's Dilemma game with errors, players with the highest average payoffs are those who make use of third-party arbitration and who also employ forgiving strategies. The combination of these two behaviours reduces the detrimental effects of errors on reciprocity, resulting in more cooperation.

Cross-cousin marriage among Tsimane forager-horticulturalists during demographic transition and market integration.

Although still prevalent in many human societies, the practice of cousin marriage has precipitously declined in populations undergoing rapid demographic and socioeconomic change. However, it is still unclear whether changes in the structure of the marriage pool or changes in the fitness-relevant consequences of cousin marriage more strongly influence the frequency of cousin marriage. Here, we use genealogical data collected by the Tsimane Health and Life History Project to show that there is a small but measurable decline in the frequency of first cross-cousin marriage since the mid-twentieth century. Such changes are linked to concomitant changes in the pool of potential spouses in recent decades. We find only very modest differences in fitness-relevant demographic measures between first cousin and non-cousin marriages. These differences have been diminishing as the Tsimane have become more market integrated. The factors that influence preferences for cousin marriage appear to be less prevalent now than in the past, but cultural inertia might slow the pace of change in marriage norms. Overall, our findings suggest that cultural changes in marriage practices reflect underlying societal changes that shape the pool of potential spouses.

Women's subsistence strategies predict fertility across cultures, but context matters.

While it is commonly assumed that farmers have higher, and foragers lower, fertility compared to populations practicing other forms of subsistence, robust supportive evidence is lacking. We tested whether subsistence activities-incorporating market integration-are associated with fertility in 10,250 women from 27 small-scale societies and found considerable variation in fertility. This variation did not align with group-level subsistence typologies. Societies labeled as "farmers" did not have higher fertility than others, while "foragers" did not have lower fertility. However, at the individual level, we found strong evidence that fertility was positively associated with farming and moderate evidence of a negative relationship between foraging and fertility. Markers of market integration were strongly negatively correlated with fertility. Despite strong cross-cultural evidence, these relationships were not consistent in all populations, highlighting the importance of the socioecological context, which likely influences the diverse mechanisms driving the relationship between fertility and subsistence.

Women's subsistence networks scaffold cultural transmission among BaYaka foragers in the Congo Basin.

In hunter-gatherer societies, women's subsistence activities are crucial for food provisioning and children's social learning but are understudied relative to men's activities. To understand the structure of women's foraging networks, we present 230 days of focal-follow data in a BaYaka community. To analyze these data, we develop a stochastic blockmodel for repeat observations with uneven sampling. We find that women's subsistence networks are characterized by cooperation between kin, gender homophily, and mixed age-group composition. During early childhood, individuals preferentially coforage with adult kin, but those in middle childhood and adolescence are likely to coforage with nonkin peers, providing opportunities for horizontal learning. By quantifying the probability of coforaging ties across age classes and relatedness levels, our findings provide insights into the scope for social learning during women's subsistence activities in a real-world foraging population and provide ground-truth values for key parameters used in formal models of cumulative culture.

Modelling animal network data in R using STRAND.

There have been recent calls for wider application of generative modelling approaches in applied social network analysis. At present, however, it remains difficult for typical end users-for example, field researchers-to implement generative network models, as there is a dearth of openly available software packages that make application of such models as simple as other, permutation-based approaches. Here, we outline the STRAND R package, which provides a suite of generative models for Bayesian analysis of animal social network data that can be implemented using simple, base R syntax. To facilitate ease of use, we provide a tutorial demonstrating how STRAND can be used to model proportion, count or binary network data using stochastic block models, social relation models or a combination of the two modelling frameworks. STRAND facilitates the application of generative network models to a broad range of data found in the animal social networks literature.

Fitness consequences of cousin marriage: a life-history assessment in two populations.

Cousin marriage, a spousal union between close kin, occurs at high frequencies in many parts of the world. The rates of cousin marriage in humans are concordant with empirical studies that challenge the traditionally held view that reproduction with kin is generally avoided in animals. Similarly, some theoretical models in animal behaviour show that inbreeding avoidance is more constrained than previously thought. Such studies highlight the importance of quantifying the costs and benefits of reproduction among close kin over the whole life-course. Here, we use genealogical data from two human populations with high frequencies of cousin marriage (the Dogon from Mali, and the Ancien Régime nobility from Europe) to estimate these potential costs and benefits. We compare age-specific fertility and survival curves, as well as the projected growth rates, of subpopulations of each marriage type. Fitness costs of cousin marriage are present in terms of reduced child survival (in both populations), while benefits exist as increased fertility for men (in the Dogon) and for women (in the Ancien Régime nobility). We also find some differences in the projected growth rates of lineages as a function of marriage type. Finally, we discuss the trade-offs that might shape marriage decisions in different ecological conditions.

Reproductive inequality in humans and other mammals.

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

Reliable network inference from unreliable data: A tutorial on latent network modeling using STRAND.

Social network analysis provides an important framework for studying the causes, consequences, and structure of social ties. However, standard self-report measures-for example, as collected through the popular "name-generator" method-do not provide an impartial representation of such ties, be they transfers, interactions, or social relationships. At best, they represent perceptions filtered through the cognitive biases of respondents. Individuals may, for example, report transfers that did not really occur, or forget to mention transfers that really did. The propensity to make such reporting inaccuracies is both an individual-level and item-level characteristic-variable across members of any given group. Past research has highlighted that many network-level properties are highly sensitive to such reporting inaccuracies. However, there remains a dearth of easily deployed statistical tools that account for such biases. To address this issue, we provide a latent network model that allows researchers to jointly estimate parameters measuring both reporting biases and a latent, underlying social network. Building upon past research, we conduct several simulation experiments in which network data are subject to various reporting biases, and find that these reporting biases strongly impact fundamental network properties. These impacts are not adequately remedied using the most frequently deployed approaches for network reconstruction in the social sciences (i.e., treating either the union or the intersection of double-sampled data as the true network), but are appropriately resolved through the use of our latent network models. To make implementation of our models easier for end-users, we provide a fully documented R package, STRAND, and include a tutorial illustrating its functionality when applied to empirical food/money sharing data from a rural Colombian population. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Distinguishing Intergroup and Long-Distance Relationships.

Intergroup and long-distance relationships are both central features of human social life, but because intergroup relationships are emphasized in the literature, long-distance relationships are often overlooked. Here, we make the case that intergroup and long-distance relationships should be studied as distinct, albeit related, features of human sociality. First, we review the functions of both kinds of relationship: while both can be conduits for difficult-to-access resources, intergroup relationships can reduce intergroup conflict whereas long-distance relationships are especially effective at buffering widespread resource shortfalls. Second, to illustrate the importance of distinguishing the two relationship types, we present a case study from rural Bolivia. Combining ethnography and two different experimental techniques, we find that the importance of intergroup relationships-and the salience of group membership itself-varies across populations and across methods. Although ethnography revealed that participants often rely on long-distance relationships for resource access, we were unable to capture participant preferences for these relationships with a forced-choice technique. Taken together, our review and empirical data highlight that (1) intergroup and long-distance relationships can have different functions and can be more or less important in different contexts and (2) validating experimental field data with ethnography is crucial for work on human sociality. We close by outlining future directions for research on long-distance relationships in humans.

Acoustic regularities in infant-directed speech and song across cultures.

When interacting with infants, humans often alter their speech and song in ways thought to support communication. Theories of human child-rearing, informed by data on vocal signalling across species, predict that such alterations should appear globally. Here, we show acoustic differences between infant-directed and adult-directed vocalizations across cultures. We collected 1,615 recordings of infant- and adult-directed speech and song produced by 410 people in 21 urban, rural and small-scale societies. Infant-directedness was reliably classified from acoustic features only, with acoustic profiles of infant-directedness differing across language and music but in consistent fashions. We then studied listener sensitivity to these acoustic features. We played the recordings to 51,065 people from 187 countries, recruited via an English-language website, who guessed whether each vocalization was infant-directed. Their intuitions were more accurate than chance, predictable in part by common sets of acoustic features and robust to the effects of linguistic relatedness between vocalizer and listener. These findings inform hypotheses of the psychological functions and evolution of human communication.

DieTryin: An R package for data collection, automated data entry, and post-processing of network-structured economic games, social networks, and other roster-based dyadic data.

Researchers studying social networks and inter-personal sentiments in bounded or small-scale communities face a trade-off between the use of roster-based and free-recall/name-generator-based survey tools. Roster-based methods scale poorly with sample size, and can more easily lead to respondent fatigue; however, they generally yield higher quality data that are less susceptible to recall bias and that require less post-processing. Name-generator-based methods, in contrast, scale well with sample size and are less likely to lead to respondent fatigue. However, they may be more sensitive to recall bias, and they entail a large amount of highly error-prone post-processing after data collection in order to link elicited names to unique identifiers. Here, we introduce an R package, DieTryin, that allows for roster-based dyadic data to be collected and entered as rapidly as name-generator-based data; DieTryin can be used to run network-structured economic games, as well as collect and process standard social network data and round-robin Likert-scale peer ratings. DieTryin automates photograph standardization, survey tool compilation, and data entry. We present a complete methodological workflow using DieTryin to teach end-users its full functionality.

Games and enculturation: A cross-cultural analysis of cooperative goal structures in Austronesian games.

While most animals play, only humans play games. As animal play serves to teach offspring important life-skills in a safe scenario, human games might, in similar ways, teach important culturally relevant skills. Humans in all cultures play games; however, it is not clear whether variation in the characteristics of games across cultural groups is related to group-level attributes. Here we investigate specifically whether the cooperativeness of games covaries with socio-ecological differences across cultural groups. We hypothesize that cultural groups that engage in frequent inter-group conflict, cooperative sustenance acquisition, or that have less stratified social structures, might more frequently play cooperative games as compared to groups that do not share these characteristics. To test these hypotheses, we gathered data from the ethnographic record on 25 ethnolinguistic groups in the Austronesian language family. We show that cultural groups with higher levels of inter-group conflict and cooperative land-based hunting play cooperative games more frequently than other groups. Additionally, cultural groups with higher levels of intra-group conflict play competitive games more frequently than other groups. These findings indicate that games are not randomly distributed among cultures, but rather relate to the socio-ecological settings of the cultural groups that practice them. We argue that games serve as training grounds for group-specific norms and values and thereby have an important function in enculturation during childhood. Moreover, games might server an important role in the maintenance of cultural diversity.

Functional 3-Dimensional Retinal Organoids: Technological Progress and Existing Challenges.

Stem cell scientists have developed methods for the self-formation of artificial organs, often referred to as organoids. Organoids can be used as model systems for research in multiple biological disciplines. Yoshiki Sasai's innovation for deriving mammalian retinal tissue from in vitro stem cells has had a large impact on the study of the biology of vision. New developments in retinal organoid technology provide avenues for in vitro models of human retinal diseases, studies of pathological mechanisms, and development of therapies for retinal degeneration, including electronic retinal implants and gene therapy. Moreover, these innovations have played key roles in establishing models for large-scale drug screening, studying the stages of retinal development, and providing a human model for personalized therapeutic approaches, like cell transplants to replace degenerated retinal cells. Here, we first discuss the importance of human retinal organoids to the biomedical sciences. Then, we review various functional features of retinal organoids that have been developed. Finally, we highlight the current limitations of retinal organoid technologies.

Erratum: Cofeeding tolerance in chimpanzees depends on group composition: A longitudinal study across four communities.

[This corrects the article DOI: 10.1016/j.isci.2021.102175.].

Cofeeding tolerance in chimpanzees depends on group composition: a longitudinal study across four communities.

Social tolerance is generally treated as a stable, species-specific characteristic. Recent research, however, has questioned this position and emphasized the importance of intraspecific variation. We investigate the temporal stability of social tolerance in four groups of sanctuary-housed chimpanzees over eight years using a commonly employed measure: experimental cofeeding tolerance. We then draw on longitudinal data on the demographic composition of each group to identify the factors associated with cofeeding tolerance. We find appreciable levels of variation in cofeeding tolerance across both groups and years that correspond closely to changes in group-level demographic composition. For example, cofeeding tolerance is lower when there are many females with young infants. These results suggest that social tolerance may be a "responding trait" of chimpanzee sociality, reflecting individual-level behavioral responses to social changes. Additional, experimental research is needed to better model the causal drivers of social tolerance within and among species.

The multinomial index: a robust measure of reproductive skew.

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.

Preferences and constraints: the value of economic games for studying human behaviour.

As economic games have spread from experimental economics to other social sciences, so too have critiques of their usefulness for drawing inferences about the 'real world'. What these criticisms often miss is that games can be used to reveal individuals' private preferences in ways that observational and interview data cannot; furthermore, economic games can be designed such that they do provide insights into real-world behaviour. Here, we draw on our collective experience using economic games in field contexts to illustrate how researchers can strategically alter the framing or design of economic games to draw inferences about private-world or real-world preferences. A detailed case study from coastal Colombia provides an example of the subtleties of game design and how games can be combined fruitfully with self-report data. We close with a list of concrete recommendations for how to modify economic games to better match particular research questions and research contexts.

Unpacking mating success and testing Bateman's principles in a human population.

Human marriage systems, characterized by long-term partnerships and extended windows of parental care, differ from the mating systems of pulsed or seasonally breeding non-human animals in which Bateman's principles were originally tested. These features, paradigmatic of but not unique to humans, complicate the accurate measurement of mating success in evaluating Bateman's three principles. Here, we unpack the concept of mating success into distinct components: number of partners, number of years partnered, the timing of partnerships, and the quality of partners. Drawing on longitudinal records of marriage and reproduction collected in a natural-fertility East African population over a 20-year period, we test and compare various models of the relationship between mating success and reproductive success (RS), and show that an accurate assessment of male and female reproductive behaviour requires consideration of all major components of mating success. Furthermore, we demonstrate that while Bateman's third principle holds when mating success is defined in terms of years married, women's fitness increases whereas men's fitness decreases from an increase in the number of marriage partners, holding constant the total effective duration of marriages. We discuss these findings in terms of the distinct, sex-specific pathways through which RS can be optimized, and comment on the contribution of this approach to the broader study of sexual selection.

Kinship ties across the lifespan in human communities.

A hypothesis for the evolution of long post-reproductive lifespans in the human lineage involves asymmetries in relatedness between young immigrant females and the older females in their new groups. In these circumstances, inter-generational reproductive conflicts between younger and older females are predicted to resolve in favour of the younger females, who realize fewer inclusive fitness benefits from ceding reproduction to others. This conceptual model anticipates that immigrants to a community initially have few kin ties to others in the group, gradually showing greater relatedness to group members as they have descendants who remain with them in the group. We examine this prediction in a cross-cultural sample of communities, which vary in their sex-biased dispersal patterns and other aspects of social organization. Drawing on genealogical and demographic data, the analysis provides general but not comprehensive support for the prediction that average relatedness of immigrants to other group members increases as they age. In rare cases, natal members of the community also exhibit age-related increases in relatedness. We also find large variation in the proportion of female group members who are immigrants, beyond simple traditional considerations of patrilocality or matrilocality, which raises questions about the circumstances under which this hypothesis of female competition are met. We consider possible explanations for these heterogenous results, and we address methodological considerations that merit increased attention for research on kinship and reproductive conflict in human societies. This article is part of the theme issue 'The evolution of female-biased kinship in humans and other mammals'.

Nahua mushroom gatherers use area-restricted search strategies that conform to marginal value theorem predictions.

We develop a method of analysis for testing the marginal value theorem (MVT) in natural settings that does not require an independent definition or mapping of patches. We draw on recent theoretical work on area-restricted search (ARS) that links turning-angle and step-size changes to geographically localized encounter-rates. These models allow us to estimate "giving-up times" using encounter-annotated GPS tracking data. Applied to a case study of Nahua mushroom foragers, these models identify distinct forms of intrapatch and interpatch search behavior, with intrapatch search transitioning to interpatch search after a predictable interval of time since the last encounter with a harvested mushroom. Our empirical estimate of giving-up time coincides with the theoretically optimal giving-up time derived under the MVT in the same environment. The MVT is currently underused in studies of human foraging and settlement patterns, due in large part to the difficulty of identifying discrete resource patches and quantifying their characteristics. Our methods mitigate the need to make such discrete maps of patches and thus have the potential to broaden the scope for empirical evaluations of the MVT and related theory in humans. Beyond studies of naturalistic foraging in humans and other animals, our approach has implications for optimization of search behavior in a range of applied fields where search dynamics must be adapted to shifting patterns of environmental heterogeneity affecting prey density and patchiness.