Collective action in culturally similar and dissimilar groups: an experiment on parochialism,conditional cooperation,and their linkages

This study examines the effects of ingroup favoritism and outgroup hostility (“parochialism”), as well as of conditionally cooperative strategies, in explaining contributions to experimental public goods games. The experimental conditions vary group composition along two culturally inheritable traits (political party preference and religious affiliation) and one trivial, “minimal” trait (birth season). We contrast ingroup, outgroup, and random group conditions and investigate the relation between the own contribution to the public good and the expectations about other group members' behavior in each one of them. We find evidence for ingroup favoritism but no support for a separate tendency towards outgroup hostility. Further, conditional cooperation and ingroup bias are, to some extent, linked. Subjects had higher expectations of the contributions of ingroup members, and their own behavior was more strongly conditioned on other group members' expected behavior in the ingroup conditions. In ingroup contexts, subjects displayed a form of “strong reciprocity” by giving more than they expected others to at high expectation levels but less at low expectation levels. Once these interactions are taken into account, we do not find a direct effect of ingroup bias anymore. We discuss these results in the light of theories of cultural group selection and conclude that too much emphasis may have been laid on direct intergroup conflict. Our results suggest that differential cooperativeness, rather than parochialism, may characterize the behavior of individuals in cultural ingroups and outgroups.     

台湾太鲁阁(Truku)族群肤纹学研究

本文报道台湾太鲁阁(Truku)的肤纹参数, 样本包括100名男性和100名女性。研究方法以《ADA标准-CDA版本》为依据、项目以《CDA标准》的Ⅱ级模式样本为依据。还有atd、tPD、猿线、指三角、同名指指纹对应的情况等, 见到非随机组合的现象。这是首次对太鲁阁人肤纹作Ⅱ级模式样本的调查, 为医学、遗传学和人类学等提供Ⅱ级模式样本。肤纹是人类的生物学性状, 由遗传和环境所致。指纹花样有每人每指各不相同和终身稳定两大特点, 成为个人身份鉴定的证明。肤纹在群体间有特异性, 在民族间有明显的差异。研究群体或民族的肤纹, 可以发现人类肤纹的遗传标记。     

Persistence of song types in Darwin's finches,Geospiza fortis,over four decades

Learned bird songs evolve via cultural evolution, with song patterns transmitted across generations by imitative learning. In Darwin''s finches of the Galápagos Islands, males learn songs from their fathers, and song types can be maintained across multiple generations. However, little is known about the time frame over which specific song types are preserved, in the face of copy errors and corresponding modifications to song structure. Here we investigate cultural evolution in songs of male Geospiza fortis, at Academy Bay, Santa Cruz Island, comparing songs recorded in 1961 by R. Bowman (20 individuals) to those recorded in 1999 by J. Podos (16 individuals). For each individual, we characterized four timing and six frequency parameters, and assessed inter-individual variation in song structure using multivariate analysis. Several 1961 song types persisted into 1999, some with remarkable fidelity. Variation among song types was extensive during both years, and we detected no changes in 10 vocal parameters across the sampling period. These results illustrate temporal continuity in a culturally acquired trait, and raise questions about mechanisms that promote stability in song structure.     

Bats are able to maintain long-term social relationships despite the high fission-fusion dynamics of their groups

Elephants, dolphins, as well as some carnivores and primates maintain social links despite their frequent splitting and merging in groups of variable composition, a phenomenon known as fission-fusion. Information on the dynamics of social links and interactions among individuals is of high importance to the understanding of the evolution of animal sociality, including that of humans. However, detailed long-term data on such dynamics in wild mammals with fully known demography and kin structures are scarce. Applying a weighted network analysis on 20,500 individual roosting observations over 5 years, we show that in two wild Bechstein's bat colonies with high fission-fusion dynamics, individuals of different age, size, reproductive status and relatedness maintain long-term social relationships. In the larger colony, we detected two stable subunits, each comprising bats from several family lineages. Links between these subunits were mainly maintained by older bats and persisted over all years. Moreover, we show that the full details of the social structure become apparent only when large datasets are used. The stable multi-level social structures in Bechstein's bat colonies resemble that of elephants, dolphins and some primates. Our findings thus may shed new light on the link between social complexity and social cognition in mammals.     

Dispersal of sibling coalitions promotes helping among immigrants in a cooperatively breeding bird

Kin selection is a major force in social evolution, but dispersal is often assumed to reduce its impact by diluting kinship. In most cooperatively breeding vertebrates, in which more than two individuals care for young, juveniles delay dispersal and become helpers in family groups. In long-tailed tits (Aegithalos caudatus), however, offspring disperse to breed and helpers are failed breeders that preferentially aid kin. Helping also occurs among immigrants, but their origins are unknown and cooperation in these cases is poorly understood. Here, we combine long-term demographic and genetic data from our study population to investigate immigration and helping in this species. We first used a novel application of parentage analysis to discriminate between immigrants and unknown philopatric recruits. We then cross-checked sibship reconstruction with pairwise relatedness estimates to show that immigrants disperse in sibling coalitions and helping among them is kin biased. These results indicate that dispersal need not preclude sociality, and dispersal of kin coalitions may help maintain kin-selected cooperation in the absence of delayed dispersal.     

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.     

Origins of altruism diversity ii: runaway coevolution of altruistic strategies via "reciprocal niche construction"

Understanding the evolution of altruism requires knowledge of both its constraints and its drivers. Here we show that, paradoxically, ecological constraints on altruism may ultimately be its strongest driver. We construct a two-trait, coevolutionary adaptive dynamics model of social evolution in a genetically structured population with local resource competition. The intensity of local resource competition, which influences the direction and strength of social selection and which is typically treated as a static parameter, is here allowed to be an evolvable trait. Evolution of survival/fecundity altruism, which requires weak local competition, increases local competition as it evolves, creating negative environmental feedback that ultimately inhibits its further evolutionary advance. Alternatively, evolution of resource-based altruism, which requires strong local competition, weakens local competition as it evolves, also ultimately causing its own evolution to stall. When evolving independently, these altruistic strategies are intrinsically self-limiting. However, the coexistence of these two altruism types transforms the negative ecoevolutionary feedback generated by each strategy on itself into positive feedback on the other, allowing the presence of one trait to drive the evolution of the other. We call this feedback conversion "reciprocal niche construction." In the absence of constraints, this process leads to runaway coevolution of altruism types. We discuss applications to the origins and evolution of eusociality, division of labor, the inordinate ecological success of eusocial species, and the interaction between technology and demography in human evolution. Our theory suggests that the evolution of extreme sociality may often be an autocatalytic process.     

A general definition of the heritable variation that determines the potential of a population to respond to selection

Genetic selection is a major force shaping life on earth. In classical genetic theory, response to selection is the product of the strength of selection and the additive genetic variance in a trait. The additive genetic variance reflects a population's intrinsic potential to respond to selection. The ordinary additive genetic variance, however, ignores the social organization of life. With social interactions among individuals, individual trait values may depend on genes in others, a phenomenon known as indirect genetic effects. Models accounting for indirect genetic effects, however, lack a general definition of heritable variation. Here I propose a general definition of the heritable variation that determines the potential of a population to respond to selection. This generalizes the concept of heritable variance to any inheritance model and level of organization. The result shows that heritable variance determining potential response to selection is the variance among individuals in the heritable quantity that determines the population mean trait value, rather than the usual additive genetic component of phenotypic variance. It follows, therefore, that heritable variance may exceed phenotypic variance among individuals, which is impossible in classical theory. This work also provides a measure of the utilization of heritable variation for response to selection and integrates two well-known models of maternal genetic effects. The result shows that relatedness between the focal individual and the individuals affecting its fitness is a key determinant of the utilization of heritable variance for response to selection.     

Express your personality or go along with the group: what determines the behaviour of shoaling perch?

Behavioural syndromes, defined as correlated behaviours in different contexts, have been studied across species and taxa including humans as part of a personality concept. While most studies have focused on solitary individuals, less is known on how shoaling fish compromise between own personality and group behaviour. Risk-taking behaviour in 1-year-old perch (Perca fluviatilis) was observed to compare individual behaviour when in a group and when alone. An experimental design gave the fish the choice between foraging in an open area in the presence of a piscivore and hiding in the vegetation. We quantified the variation accountable by the effect of individuals being in a group, individuals alone and repeated measurements, using hierarchical mixed effects models. Within-group variances were low, but when individuals were later tested alone, individual differences explained most of the variation. Still, the individual best linear unbiased predictors (BLUPs) of time spent in the open area, extracted from the random effects of the mixed effects model, were positively correlated with the corresponding BLUPs when alone. The results indicate that individual behavioural traits are to some degree expressed also within groups. Most fish showed a shyer behaviour when alone, but bolder individuals changed less between treatments than did shyer ones, suggesting a more influential role of bold fish in the group.     

Remotely sensed between‐individual functional trait variation in a temperate forest

1. Trait‐based ecology holds the promise to explain how plant communities work, for example, how functional diversity may support community productivity. However, so far it has been difficult to combine field‐based approaches assessing traits at the level of plant individuals with limited spatial coverage and approaches using remote sensing (RS) with complete spatial coverage but assessing traits at the level of vegetation pixels rather than individuals. By delineating all individual‐tree crowns within a temperate forest site and then assigning RS‐derived trait measures to these trees, we combine the two approaches, allowing us to use general linear models to estimate the influence of taxonomic or environmental variation on between‐ and within‐species variation across contiguous space.
2. We used airborne imaging spectroscopy and laser scanning to collect individual‐tree RS data from a mixed conifer‐angiosperm forest on a mountain slope extending over 5.5 ha and covering large environmental gradients in elevation as well as light and soil conditions. We derived three biochemical (leaf chlorophyll, carotenoids, and water content) and three architectural traits (plant area index, foliage‐height diversity, and canopy height), which had previously been used to characterize plant function, from the RS data. We then quantified the contributions of taxonomic and environmental variation and their interaction to trait variation and partitioned the remaining within‐species trait variation into smaller‐scale spatial and residual variation. We also investigated the correlation between functional trait and phylogenetic distances at the between‐species level. The forest consisted of 13 tree species of which eight occurred in sufficient abundance for quantitative analysis.
3. On average, taxonomic variation between species accounted for more than 15% of trait variation in biochemical traits but only around 5% (still highly significant) in architectural traits. Biochemical trait distances among species also showed a stronger correlation with phylogenetic distances than did architectural trait distances. Light and soil conditions together with elevation explained slightly more variation than taxonomy across all traits, but in particular increased plant area index (light) and reduced canopy height (elevation). Except for foliage‐height diversity, all traits were affected by significant interactions between taxonomic and environmental variation, the different responses of the eight species to the within‐site environmental gradients potentially contributing to the coexistence of the eight abundant species.
4. We conclude that with high‐resolution RS data it is possible to delineate individual‐tree crowns within a forest and thus assess functional traits derived from RS data at individual level. With this precondition fulfilled, it is then possible to apply tools commonly used in field‐based trait ecology to partition trait variation among individuals into taxonomic and potentially even genetic variation, environmental variation, and interactions between the two. The method proposed here presents a promising way of assessing individual‐based trait information with complete spatial coverage and thus allowing analysis of functional diversity at different scales. This information can help to better understand processes shaping community structure, productivity, and stability of forests.

     

Paternal Relatedness Predicts the Strength of Social Bonds among Female Rhesus Macaques

Forming strong, equitable, and enduring social bonds with a few individuals in a group carries adaptive benefits in terms of increased longevity, offspring survival and paternity success in birds and mammals, including humans. These recent insights generated a new interest in the factors creating variation in the strength of social relationships. Whether and how animals discriminate paternal kin from non-kin and bias their social behavior accordingly is being debated. This study explores the relative importance of dominance rank, age, maternal and paternal relatedness in shaping dyadic affiliative relationships in a group of 30 captive rhesus macaque females. The strength of social relationships, measured by the composite sociality index from observational data, was independently predicted in GLMMs by both maternal and paternal relatedness as well as rank similarity. In addition, social bonds between paternal half-sisters were stronger than between distantly related kin suggesting that females biased their affiliative effort towards paternal relatives. Despite identical relatedness coefficients bonds between mothers and their daughters were three times as strong as those between full sisters. Together these results add to the growing body of evidence for paternal kin biases in affiliative behavior and highlight that females prefer descendent over lateral kin.     

The Big Man Mechanism: how prestige fosters cooperation and creates prosocial leaders

Anthropological evidence from diverse societies suggests that prestige-based leadership may provide a foundation for cooperation in many contexts. Here, inspired by such ethnographic observations and building on a foundation of existing research on the evolution of prestige, we develop a set of formal models to explore when an evolved prestige psychology might drive the cultural evolution of n-person cooperation, and how such a cultural evolutionary process might create novel selection pressures for genes that make prestigious individuals more prosocial. Our results reveal (i) how prestige can foster the cultural emergence of cooperation by generating correlated behavioural phenotypes, both between leaders and followers, and among followers; (ii) why, in the wake of cultural evolution, natural selection favours genes that make prestigious leaders more prosocial, but only when groups are relatively small; and (iii), why the effectiveness of status differences in generating cooperation in large groups depends on cultural transmission (and not primarily on deference or coercion). Our theoretical framework, and the specific predictions made by these models, sketch out an interdisciplinary research programme that cross-cuts anthropology, biology, psychology and economics. Some of our predictions find support from laboratory work in behavioural economics and are consistent with several real-world patterns.     

Differences in prey personality mediate trophic cascades

Functional trait approaches in ecology chiefly assume the mean trait value of a population adequately predicts the outcome of species interactions. Yet this assumption ignores substantial trait variation among individuals within a population, which can have a profound effect on community structure and function. We explored individual trait variation through the lens of animal personality to test whether among‐individual variation in prey behavior mediates trophic interactions. We quantified the structure of personalities within a population of generalist grasshoppers and examined, through a number of field and laboratory‐based experiments, how personality types could impact tri‐trophic interactions in a food chain. Unlike other studies of this nature, we used spatial habitat domains to evaluate how personality types mechanistically map to behaviors relevant in predator–prey dynamics and found shy and bold individuals differed in both their habitat use and foraging strategy under predation risk by a sit‐and‐wait spider predator. In the field‐based mesocosm portion of our study, we found experimental populations of personality types differed in their trophic impact, demonstrating that prey personality can mediate trophic cascades. We found no differences in respiration rates or body size between personality types used in the mesocosm experiment, indicating relative differences in trophic impact were not due to variation in prey physiology but rather variation in behavioral strategies. Our work demonstrates how embracing the complexity of individual trait variation can offer mechanistically richer understanding of the processes underlying trophic interactions.     

Personality and problem-solving performance explain competitive ability in the wild

Competitive ability is a major determinant of fitness, but why individuals vary so much in their competitiveness remains only partially understood. One increasingly prevalent view is that realized competitive ability varies because it represents alternative strategies that arise because of the costs associated with competitiveness. Here we use a population of great tits (Parus major) to explore whether individual differences in competitive ability when foraging can be explained by two traits that have previously been linked to alternative behavioural strategies: the personality trait 'exploration behaviour' and a simple cognitive trait, 'innovative problem-solving performance'. We assayed these traits under standardized conditions in captivity and then measured competitive ability at feeders with restricted access in the wild. Competitive ability was repeatable within individual males across days and correlated positively with exploration behaviour, representing the first such demonstration of a link between a personality trait and both competitive ability and food intake in the wild. Competitive ability was also simultaneously negatively correlated with problem-solving performance; individuals who were poor competitors were good at problem-solving. Rather than being the result of variation in 'individual quality', our results support the hypothesis that individual variation in competitive ability can be explained by alternative behavioural strategies.     

Quantifying individual heterogeneity and its influence on life‐history trajectories: different methods for different questions and contexts

Heterogeneity among individuals influences the life‐history trajectories we observe at the population level because viability selection, selective immigration and emigration processes, and ontogeny change the proportion of individuals with specific trait values with increasing age. Here, we review the two main approaches that have been proposed to account for these processes in life‐history trajectories, contrasting how they quantify ontogeny and selection, and proposing ways to overcome some of their limitations. Nearly all existing approaches to model individual heterogeneity assume either a single normal distribution or a priori known groups of individuals. Ontogenetic processes, however, can vary across individuals through variation in life‐history tactics. We show the usefulness of describing ontogenetic processes by modelling trajectories with a mixture model that focuses on heterogeneity in life‐history tactics. Additionally, most methods examine individual heterogeneity in a single trait, ignoring potential correlations among multiple traits caused by latent common sources of individual heterogeneity. We illustrate the value of using a joint modelling approach to assess the presence of a shared latent correlation and its influence on life‐history trajectories. We contrast the strengths and limitations of different methods for different research questions, and we exemplify the differences among methods using empirical data from long‐term studies of ungulates.     

Cultural niche construction and the evolution of small family size

A model of cultural niche construction with two culturally transmitted traits is examined. The frequency of individuals with a certain general predisposition, which is transmitted vertically, plays a role as the cultural background, or the cultural niche, of the population. The cultural background determines the rate of oblique, relative to vertical, transmission of another cultural trait that affects fertility of individuals. It is assumed that individuals with fewer offspring are more likely to achieve social roles that influence the succeeding generation and are therefore overrepresented as transmitters in the process of oblique transmission. Our model suggests that even a slight overrepresentation of those with fewer offspring can drive the evolution of small family size, provided that the rate of oblique transmission depends strongly on the cultural background. In addition, our model may help to explain the time lag between the decrease in death rates and the subsequent decrease in birth rates during the demographic transition of industrializing societies.     

ESTIMATING THE FORM OF NATURAL SELECTION ON A QUANTITATIVE TRAIT

The fitness function f relates fitness of individuals to the quantitative trait under natural selection. The function is useful in predicting fitness differences among individuals and in revealing whether an optimum is present within the range of phenotypes in the population. It may also be thought of as describing the ecological environment in terms of the trait. Quadratic regression will approximate the fitness function from data (e.g., Lande and Arnold, 1983), but the method does not reliably indicate features of f such as the presence of modes (stabilizing selection) or dips (disruptive selection). I employ an alternative procedure requiring no a priori model for the function. The method is useful in two ways: it provides a nonparametric estimate of f, of interest by itself, and it can be used to suggest an appropriate parametric model. I also discuss measures of selection intensity based on the fitness function. Analysis of six data sets yields a variety of forms of f and provides new insights for some familiar cases. Low amounts of variation and a low density of data points near the tails of many phenotype distributions emerge as limitations to gaining information on fitness functions. An experimental approach in which the distribution of a quantitative trait is broadened through manipulation would minimize these problems.     

Geographic patterns in the distribution of social systems in terrestrial arthropods

The role of ecology in the evolution and maintenance of arthropod sociality has received increasing research attention in recent years. In some organisms, such as halictine bees, polistine wasps, and social spiders, researchers are investigating the environmental factors that may contribute to high levels of variation in the degree of sociality exhibited both among and within species. Within lineages that include only eusocial members, such as ants and termites, studies focus more on identifying extrinsic factors that may contribute to the dramatic variation in colony size, number of queens, and division of labour that is evident across these species. In this review, I propose a comparative approach that seeks to identify environmental factors that may have a common influence across such divergent social arthropod groups. I suggest that seeking common biogeographic patterns in the distribution of social systems or key social traits may help us to identify ecological factors that play a common role in shaping the evolution of sociality across different organisms. I first review previous studies of social gradients that form along latitudinal and altitudinal axes. Within families and within species, many organisms show an increasing degree of sociality at lower latitudes and altitudes. In a smaller number of cases, organisms form larger groups or found nests cooperatively at higher latitudes and altitudes. I then describe several environmental factors that vary consistently along such gradients, including climate variables and abundance of predators, and outline their proposed role in the social systems of terrestrial arthropods. Finally, I map distributions of a social trait against several climatic factors in five case studies to demonstrate how future comparative studies could inform empirical research.     

Non‐consumptive effects of predation: does perceived risk strengthen the genetic integration of behaviour and morphology in stickleback?

Predators can shape genetic correlations in prey by altering prey perception of risk. We manipulated perceived risk to test whether such non‐consumptive effects tightened behavioural trait correlations in wild‐caught stickleback from high‐ compared to low‐risk environments due to genetic variation in plasticity. We expected tighter genetic correlations within perceived risk treatments than across them, and tighter genetic correlations in high‐risk than in low‐risk treatments. We identified genetic variation in plasticity, with genetic correlations between boldness, sociality, and antipredator morphology, as expected, being tighter within treatments than across them, for both of two populations. By contrast, genetic correlations did not tighten with exposure to risk. Tighter phenotypic correlations in wild stickleback may thus arise because predators induce correlational selection on environmental components of these traits, or because predators tighten residual correlations by causing environmental heterogeneity that is controlled in the laboratory. Our study places phenotypic integration firmly into an ecological context.