Fitness consequences of spousal relatedness in 46 small-scale societies

Social norms that regulate reproductive and marital decisions generate impressive cross-cultural variation in the prevalence of kin marriages. In some societies, marriages among kin are the norm and this inbreeding creates intensive kinship networks concentrated within communities. In others, especially forager societies, most marriages are between more genealogically and geographically distant individuals, which generates a larger number of kin and affines of lesser relatedness in more extensive kinship networks spread out over multiple communities. Here, we investigate the fitness consequence of kin marriages across a sample of 46 small-scale societies (12 439 marriages). Results show that some non-forager societies (including horticulturalists, agriculturalists and pastoralists), but not foragers, have intensive kinship societies where fitness outcomes (measured as the number of surviving children in genealogies) peak at commonly high levels of spousal relatedness. By contrast, the extensive kinship systems of foragers have worse fitness outcomes at high levels of spousal relatedness. Overall, societies with greater levels of inbreeding showed a more positive relationship between fitness and spousal relatedness.     

Relatedness and Social Behaviors in <Emphasis Type="Italic">Cercopithecus solatus</Emphasis>

Inclusive fitness and kin selection theories predict that organisms will evolve biased behavior toward kin when the inclusive fitness benefits outweigh the costs of such behaviors. Researchers have long observed that primates bias their behavior toward relatives, particularly maternal kin. We examined the effect of kinship on social behaviors in a semifree-ranging colony of Cercopithecus solatus, a poorly studied forest guenon species. We used microsatellite loci and paternity analyses to determine the degree of relatedness between individuals, as well as kinship. Individuals biased some of their behavior according to relatedness. Specifically, related individuals are more spatially associated and less aggressive toward each other. When we replaced the relatedness coefficients with defined kin categories, Cercopithecus solatus seemed to behave preferentially toward maternal kin versus paternal kin. Even though the setting of the colony and the small sample size limit our conclusions, we discuss the potential implications of our finding for the study of the impact of kin selection in primate social relationships.     

Amazonian horticulturalists live in larger,more related groups than hunter–gatherers

The relatedness of human groups has important ramifications for kin (group) selection to favor more collective action and invites the potential for more exploitation by political leaders. Endogamous marriages among kin create intensive kinship systems with high group relatedness, while exogamous marriages among nonrelatives create extensive kinship with low group relatedness. Here, a sample of 58 societies (7,565 adults living in 353 residential groups) shows that average group relatedness is higher in lowland horticulturalists than in hunter–gatherers. Higher relatedness in horticulturalists is remarkable given that village sizes are larger, harboring over twice the average number of adults than in hunter–gatherer camps. The relatedness differential between subsistence regimes increases for larger group sizes. Large and dense networks of kin may have favored an increased propensity for some forms of in-group cooperation and political inequality that emerged relatively recently in human history, after the advent of farming.     

A theoretical basis for measures of kin selection in subdivided populations: finite populations and localized dispersal

The analysis of kin selection in subdivided populations has been hampered by the lack of well‐defined measures of genealogical relatedness in the presence of localized dispersal. Furthermore, the usual arguments underlying the definition of game‐theoretical measures of inclusive fitness are not exact under localized dispersal. We define such measures to give the first‐order effects of selection on the probability of fixation of an allele. The derived measures of kin selection and relatedness are valid in finite populations and under localized dispersal. For the infinite island model, the resulting measure of kin selection is equivalent to a previously used measure. In other cases its definition is based on definitions of relatedness which are different from the usual ones. To illustrate the approach, we reanalyse a model with localized dispersal. We consider sex ratio evolution under sex‐specific dispersal behaviour, and the results confirm the earlier conclusion that the sex ratio is biased towards the sex with the dispersal rate closer to the optimal dispersal rate in the absence of sex‐specific dispersal behaviour.     

Model-free Estimation of Recent Genetic Relatedness

Genealogical inference from genetic data is essential for a variety of applications in human genetics. In genome-wide and sequencing association studies, for example, accurate inference on both recent genetic relatedness, such as family structure, and more distant genetic relatedness, such as population structure, is necessary for protection against spurious associations. Distinguishing familial relatedness from population structure with genotype data, however, is difficult because both manifest as genetic similarity through the sharing of alleles. Existing approaches for inference on recent genetic relatedness have limitations in the presence of population structure, where they either (1) make strong and simplifying assumptions about population structure, which are often untenable, or (2) require correct specification of and appropriate reference population panels for the ancestries in the sample, which might be unknown or not well defined. Here, we propose PC-Relate, a model-free approach for estimating commonly used measures of recent genetic relatedness, such as kinship coefficients and IBD sharing probabilities, in the presence of unspecified structure. PC-Relate uses principal components calculated from genome-screen data to partition genetic correlations among sampled individuals due to the sharing of recent ancestors and more distant common ancestry into two separate components, without requiring specification of the ancestral populations or reference population panels. In simulation studies with population structure, including admixture, we demonstrate that PC-Relate provides accurate estimates of genetic relatedness and improved relationship classification over widely used approaches. We further demonstrate the utility of PC-Relate in applications to three ancestrally diverse samples that vary in both size and genealogical complexity.     

Measures of Relatedness

Measures of genetic relatedness are essential to models of evolution by kin selection and determinations of inclusive fitness. Under a kin selection paradigm, individuals are expected to distribute actions influencing the fitness of relatives based on the relatedness of these relatives. In addition, it is necessary to have an accurate measure of relatedness to estimate heritability (h²) of phenotypic characters and to predict the efficacy of selection. Relatedness is often defined as the genotypic correlation between individuals. Assessed on the basis of common ancestry, relatedness can only be determined sensu strictu from pedigree analysis. Recent methodological and statistical advances allow the estimation of relatedness from allele frequency data. Many coefficients of relatedness can be found in the literature; I review and evaluate these, with emphasis on situations for which each is appropriate.     

Cooperative pastoral production — the importance of kinship

While there is a general assumption that labour has a positive effect on pastoral production, studies that have quantified this relationship have been characterized by ambiguous results. This is most likely related to the fact that possible cooperative pastoral production has been little explored in the literature, although it is well documented that nomadic pastoralist households share and exchange labour in so-called cooperative herding groups. Consequently, this study aims at investigating possible cooperative labour-related effects on production among Saami reindeer herders in Norway by using kinship relations as a proxy for cooperation. This study found that cooperative labour investment is important for Saami reindeer herders, but that the effect of kinship and labour needs to be understood in relation to each other. When assessing the effect of labour and kinship simultaneously, both labour and genealogical relationship had positive effects on herd size. We also found a positive interaction between kinship and labour suggesting that high levels of relatedness coupled with a large potential labour pool had an increasingly positive effect on herd size.     

Detecting instability in animal social networks: genetic fragmentation is associated with social instability in rhesus macaques

The persistence of biological systems requires evolved mechanisms which promote stability. Cohesive primate social groups are one example of stable biological systems, which persist in spite of regular conflict. We suggest that genetic relatedness and its associated kinship structure are a potential source of stability in primate social groups as kinship structure is an important organizing principle in many animal societies. We investigated the effect of average genetic relatedness per matrilineal family on the stability of matrilineal grooming and agonistic interactions in 48 matrilines from seven captive groups of rhesus macaques. Matrilines with low average genetic relatedness show increased family-level instability such as: more sub-grouping in their matrilineal groom network, more frequent fighting with kin, and higher rates of wounding. Family-level instability in multiple matrilines within a group is further associated with group-level instability such as increased wounding. Stability appears to arise from the presence of clear matrilineal structure in the rhesus macaque group hierarchy, which is derived from cohesion among kin in their affiliative and agonistic interactions with each other. We conclude that genetic relatedness and kinship structure are an important source of group stability in animal societies, particularly when dominance and/or affilative interactions are typically governed by kinship.     

Cultural relatedness under oblique and horizontal transmission rules

The concept of genetic relatedness is central to the sociobiological theory of kinship altruism. Genes which code for altruistic acts toward closely related genetic kin can overcome selection pressures favoring self-interested behavior. But genetic relatedness declines rapidly as kinship becomes more distant, thereby restricting genetically-based altruism to very close kin. This paper extends these notions to cultural relatedness, arguing that oblique and horizontal transmission can yield high levels of cultural relatedness in larger groups. A mathematical model of cultural transmission is proposed, and equilibria for several special cases are investigated. For all models, the equilibrium level of relatedness is critically dependent on the influence of exogenous sources. In models with equal influence of nonparents, the level of relatedness varies inversely with group size. On the other hand, when nonparental influence is concentrated on a single individual, the level of relatedness does not vary with group size.     

The role of kin recognition in the evolution of conspecific brood parasitism

Conspecific brood parasitism (CBP) is a common strategy in several species of birds. Currently, some studies suggest that relatedness between host and parasite enhances CBP, since indirect fitness benefits could select for acceptance of related eggs by hosts. Conversely, parasites should avoid laying eggs in nests of relatives if this is costly for the host. Based on the latter argument, kinship should not promote brood parasitism. A recent model clarified this relationship, and showed that kinship can promote brood parasitism, assuming kin recognition. However, in that model kin recognition was assumed perfect. Here we present a model that addresses the role of relatedness and kin selection in CBP, when kin recognition is not perfect and hosts do not always detect parasitism. We consider both the indirect fitness of the parasite and the possible responses of the host. Our results indicate that the existence and accuracy of a kin recognition system is crucial to the final outcome. When CBP represents a cost to the host, a parasitic female that has the choice should avoid parasitizing relatives, unless (1) the costs are not too high and (2) hosts can accurately enough recognize eggs laid by relatives, rejecting them less often than eggs laid by nonkin. But if ‘parasitism’ enhances the direct fitness of the host (which is possible in species with precocial young) parasites should choose relatives whenever possible, even if hosts do not recognize kin eggs. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Blood is not always thicker than water:The limited effect of kin selection on human kinship in the traditional Chinese family

To examine the importance of kin selection in shaping human societies, this study analyzed the kinship system practiced in traditional China for two millennia and teased apart its underlying genetic and other, presumably cultural, components.The results demonstrate that, in the traditional patrilineal Chinese family, both genetic relatedness and the cultural factor of generation were important in determining kinship status for male agnates (genetically related relatives). For female agnates, however, only genetic relatedness was important. Another surprising finding was that the influence of gender was not as important as genetic relatedness. The most interesting finding in this study, however, was that kin selection and culture (I.e., seniority in generation and age) played vastly different roles in different lineages in the Chinese family: for collateral (indirect) agnates, genetic relatedness was the most important factor in determining their kinship status, but for lineal (direct) agnates, its importance was overridden by seniority in generation and age, a cultural factor. Several other bio-cultural factors also explained a considerable amount of variance in kinship status. Since kinship profoundly affected, and was often the foundation of, the legal and social systems in dynastic China, kin selection, while its strength may differ remarkably between lineal and collateral relatives, could act as a selective force in Chinese families.     

Strategies for determining kinship in wild populations using genetic data

Knowledge of kin relationships between members of wild animal populations has broad application in ecology and evolution research by allowing the investigation of dispersal dynamics, mating systems, inbreeding avoidance, kin recognition, and kin selection as well as aiding the management of endangered populations. However, the assessment of kinship among members of wild animal populations is difficult in the absence of detailed multigenerational pedigrees. Here, we first review the distinction between genetic relatedness and kinship derived from pedigrees and how this makes the identification of kin using genetic data inherently challenging. We then describe useful approaches to kinship classification, such as parentage analysis and sibship reconstruction, and explain how the combined use of marker systems with biparental and uniparental inheritance, demographic information, likelihood analyses, relatedness coefficients, and estimation of misclassification rates can yield reliable classifications of kinship in groups with complex kin structures. We outline alternative approaches for cases in which explicit knowledge of dyadic kinship is not necessary, but indirect inferences about kinship on a group‐ or population‐wide scale suffice, such as whether more highly related dyads are in closer spatial proximity. Although analysis of highly variable microsatellite loci is still the dominant approach for studies on wild populations, we describe how the long‐awaited use of large‐scale single‐nucleotide polymorphism and sequencing data derived from noninvasive low‐quality samples may eventually lead to highly accurate assessments of varying degrees of kinship in wild populations.     

Fluctuating populations and kin interaction in mammals

Kin selection in animals favors less aggressive interaction among related individuals. If the genetic relatedness among neighbors changes with population structure and density, behavioral interaction may also change according to the population structure. Charnov and Finerty proposed a hypothesis that kin selection in voles causes population cycles if the relatedness among neighbors decreases as density increases. Field experiments have recently tested this hypothesis. Furthermore, field studies of social interaction in voles have increased in number, so that the effects of kinship on reproductive success can be reviewed. These studies indicate that although kin interaction might be an important factor affecting social behavior and reproductive success in voles, the relationships both between kinship and degree of amicable behavior or reproductive rate, and between relatedness among neighbors and population density, are far less simple than had been supposed.     

Alliance membership and kinship in wild male bottlenose dolphins (Tursiops aduncus) of southeastern Australia

Bottlenose dolphins are one of only a few mammalian taxa where the males are known to cooperate within their social group in order to maintain mating access to single females against other males. Male bonds in bottlenose dolphins have been hypothesized as evolving through kinship and associated inclusive fitness effects. In this study we tested whether individually identified male bottlenose dolphins preferentially associate and form alliances with kin in a small coastal resident population of southeastern Australia using a combination of behavioural data, genetic sexing, sequences of the mitochondrial DNA control region and nuclear microsatellite markers. Males generally associated significantly more often than expected with one to three other males, with whom they jointly herded females for mating. Associations and alliance membership were not associated with either maternal kinship or genetic relatedness. The majority of male pairs within alliances were randomly related, although high relatedness values were found between males of different alliances in the resident population. These findings indicate that mechanisms other than kin selection may be foremost in the development and maintenance of cooperation between male bottlenose dolphins.     

Ground squirrel sociality and the quest for the 'holy grail': does kinship influence behavioral discrimination by juvenile Columbian ground squirrels?

Research on social discrimination by ground-dwelling squirrelshas focused on die ability of squirrels to discriminate kinfrom nonkin. The predominant mechanism underlying that abilityis social familiarization. Although familiarity-based mechanismsmay result in kin discrimination where kin are associated reliablyin space and time, past investigations have largely ignoredadditional levels of social discrimination that may result fromongoing familiarization in the natural context Here we presentdata from a cross-fostering experiment that examines the relativecontributions of rearing association and relatedness to subsequentbehavioral discrimination among juvenile Columbian ground squirrels(Spermophilus columbianus). Rearing association significantlyinfluenced recognitive and agonistic behavior among juvenileswhile relatedness proved insignificant in affecting behavioralinteractions. Thus, direct familiarization in the natal burrowis both sufficient and necessary to account for the transitiveappearance of kin-differential behavior among newly emergedjuveniles documented in previous studies. Reanalyses of datafrom field experiments on S. columbianus social discriminationfail to detect any evidence of kin-biased behavior, and evidencefrom studies purporting to document kinship effects is equivocalat best Taken together with more recent findings, the scantdata suggestive of kin discrimination by Columbian ground squirrelsare best viewed as an artifact of selection promoting the discriminationof familiar from unfamiliar individuals at the level of localizedgroups. Fitness payoffs of such discrimination may accrue viareciprocal altruism, or ‘dear-enemy’ recognition,and would promote the evolution and maintenance of socialitywithout recourse to indirect components of inclusive fitnesscommonly invoked to explain ground squirrel sociality. Finally,our data call into doubt the notion that mechanisms allowingthe direct assessment of kinship should be common among themore social sciurids.     

ml‐relate: a computer program for maximum likelihood estimation of relatedness and relationship

Genetic data are useful for estimating the genealogical relationship or relatedness between individuals of unknown ancestry. We present a computer program, ml ‐relate that calculates maximum likelihood estimates of relatedness and relationship. ml ‐relate is designed for microsatellite data and can accommodate null alleles. It uses simulation to determine which relationships are consistent with genotype data and to compare putative relationships with alternatives. ml ‐relate runs on the Microsoft Windows operating system and is available from http://www.montana.edu/kalinowski .     

Detecting kin selection at work using inclusive fitness

A recent model shows that altruism can evolve with limited migration and variable group sizes, and the authors claim that kin selection cannot provide a sufficient explanation of their results. It is demonstrated, using a recent reformulation of Hamilton's original arguments, that the model falls squarely within the scope of inclusive fitness theory, which furthermore shows how to calculate inclusive fitness and the relevant relatedness. A distinction is drawn between inclusive fitness, which is a method of analysing social behaviour; and kin selection, a process that operates through genetic similarity brought about by common ancestry, but not by assortation by genotype or by direct assessment of genetic similarity. The recent model is analysed, and it turns out that kin selection provides a sufficient explanation to considerable quantitative accuracy, contrary to the authors' claims. A parallel analysis is possible and would be illuminating for all models of social behaviour in which individuals' effects on each other's offspring numbers combine additively.     

Relationships among companies of militia in the colony of New York in 1760 estimated by an analysis of their surnames

Abstract

Surname, analysis, a technique to estimate genetic relatedness, is applied here to differences within and between eight militia companies (N=782) mustered for New York Colony in 1760. Universal service laws of the time imply that militias are unbiased community samples. Colonial American populations are of interest due to their heterogeneous migrant origins, balanced by effects of large kin groups. The surname of each militiaman was checked for identity to that of all others within and between the eight companies. Assuming surnames derive from common ancestry, fractions of name identity index relatedness. Within‐company relatedness is high, below only that found in genetic isolates, probably due to enlistments together of kinsmen. Between‐company relatedness is the lowest yet recorded, apparently because of the diverse origins of these populations.     

Does morality have a biological basis? An empirical test of the factors governing moral sentiments relating to incest.

Kin-recognition systems have been hypothesized to exist in humans, and adaptively to regulate altruism and incest avoidance among close genetic kin. This latter function allows the architecture of the kin recognition system to be mapped by quantitatively matching individual variation in opposition to incest to individual variation in developmental parameters, such as family structure and co-residence patterns. Methodological difficulties that appear when subjects are asked to disclose incestuous inclinations can be circumvented by measuring their opposition to incest in third parties, i.e. morality. This method allows a direct test of Westermarck's original hypothesis that childhood co-residence with an opposite-sex individual predicts the strength of moral sentiments regarding third-party sibling incest. Results support Westermarck's hypothesis and the model of kin recognition that it implies. Co-residence duration objectively predicts genetic relatedness, making it a reliable cue to kinship. Co-residence duration predicts the strength of opposition to incest, even after controlling for relatedness and even when co-residing individuals are genetically unrelated. This undercuts kin-recognition models requiring matching to self (through, for example, major histocompatibility complex or phenotypic markers). Subjects' beliefs about relatedness had no effect after controlling for co-residence, indicating that systems regulating kin-relevant behaviours are non-conscious, and calibrated by co-residence, not belief.     

Population and family studies on Kuru

Abstract

A model of group fissioning, applied by Mange (1964) to a Hutterite population and by Chagnon (1975) to an Amazonian Indian population, is applied to the “Nebraska” Amish community in Central Pennsylvania. Genealogical data were collected on 228 living, married individuals and their known ancestors in two new churches resulting from the fission of a church in 1978. The acrimonious fissioning of the church produced increased levels of intrachurch genealogical relatedness in the two new churches formed, as compared to the level of relatedness in the original church, with a greater increase for male/male relatedness than for female/female relatedness. This general increase in relatedness, documented for three human populations and also for fissioning groups of rhesus macaques, conforms to predictions from kin selection theory.