Cryptic Kin Selection: Kin Structure in Vertebrate Populations and Opportunities for Kin‐Directed Cooperation

Animal societies of varying complexity have been the favoured testing ground for inclusive fitness theory, and there is now abundant evidence that kin selection has played a critical role in the evolution of cooperative behaviour. One of the key theoretical and empirical findings underlying this conclusion is that cooperative systems have a degree of kin structure, often the product of delayed dispersal, that facilitates interactions with relatives. However, recent population genetic studies have revealed that many non‐cooperative animals also have kin‐structured populations, providing more cryptic opportunities for kin selection to operate. In this article, I first review the evidence that kin structure is widespread among non‐cooperative vertebrates, and then consider the various contexts in which kin selection may occur in such taxa, including: leks, brood parasitism, crèches, breeding associations, territoriality and population dynamics, foraging and predator deterrence. I describe the evidence that kin‐selected benefits arise from interacting with kin in each of these contexts, notwithstanding the potential costs of kin competition and inbreeding. I conclude that as the tools required to determine population genetic structure are readily available, measurement of kin structure and the potential for kin selection on a routine basis is likely to reveal that this process has been an important driver of evolutionary adaptation in many non‐cooperative as well as cooperative species.     

Signature Systems and Kin Recognition

SYNOPSIS. This paper examines mechanisms of individual identificationin the kin recognition process. In type 1 kin recognition, theindividual (e.g., a parent) learns the identifying cues (the"signature") of particular kin (e.g., offspring) at some stagewhen there are reliable contextual cues as to kinship (e.g.,the parent finds the young in its nest). These kin are subsequentlyrecognized via their signatures in situations in which contextualevidence of kinship is lacking. The primary selection pressurein this case is for a mechanism that can generate a large numberof distinctive signatures, and thus permit recognition in largegroups. I develop a model that predicts the necessary informationcapacity of a signature system, and I test this prediction forthe signature call of the bank swallow (Riparia riparia). Themeasured information capacity of 17 bits corresponds well tothe prediction and, moreover, is substantially greater thanthat of the homologous call of the similar but non-colonialrough-winged swallow (Stelgidopteryx ruficollis). In type 2kin recognition, reliable contextual evidence as to kinshipis absent at all times, and kin must be recognized via a signature-matchingprocess. For example, the individual might compare its own signatureto that of the unknown individual, inferring kinship if theirsignatures are sufficiently similar. I describe a multi-locusgenetic signature mechanism that would permit discriminationof kin from non-kin, and suggest how this model might be tested     

Kin Preference and Partner Choice

This paper presents a comparison of social kinship (patrilineage) and biological kinship (genetic relatedness) in predicting cooperative relationships in two different economic contexts in the fishing and whaling village of Lamalera, Indonesia. A previous analysis (Alvard, Human Nature 14:129-163, 2003) of boat crew affiliation data collected in the village in 1999 found that social kinship (patrilineage) was a better predictor of crew affiliation than was genetic kinship. A replication of this analysis using similar data collected in 2006 finds the same pattern: lineage is a better predictor than genetic kinship of crew affiliation, and the two together explain little additional variance over that explained by lineage alone. However, an analogous test on food-sharing relationships finds the opposite pattern: biological kinship is a better predictor of food-sharing relationships than is social kinship. The difference between these two cooperative contexts is interpreted in terms of kin preferences that shape partner choice, and the relative autonomy with which individuals can seek to satisfy those preferences. Drawing on stable matching theory, it is suggested that unilineal descent may serve as a stable compromise among multiple individuals' incongruent partner preferences, with patriliny favored over matriliny in the crew-formation context because it leads to higher mean degrees of relatedness among male cooperators. In the context of food-sharing, kin preferences can be pursued relatively autonomously, without the necessity of coordinating preferences with those of other households through the institution of lineage.     

Kin discrimination in salmonids

Summary The data presented here suggest that significant selection pressures towards kin discrimination behaviour patterns result from kin-biased territorial defence behaviour patterns. Salmonids employ a phenotype matching recognition mechanism allowing individuals to discriminate unfamiliar kin. Kin discrimination abilities allow individuals to reduce the levels of aggression associated with territorial defence towards related conspecifics and to defend smaller territories near kin versus non-kin. This kin-biased territorial defence behaviour is observed in at least one species under a wide range of territorial quality conditions. Within kin groups, subordinate individuals obtain a greater number of foraging attempts, resulting in kin-biased foraging within the social group. As a result of this kin-bias, individuals within kin groups show significantly higher mean weight increases (increased direct fitness benefits) and reduced variance in these increases (increased indirect benefits). Since all individuals within the kin groups obtained higher, less variable weight increases, we can argue that individuals are increasing their inclusive fitness as a result of these kin-biased behaviour patterns.Based on these results, and on what is known about the life history of a variety of salmonid and non-salmonid species, we can formulate a number of testable predictions. By testing these predictions, we may be better able to understand both the proximate and ultimate causation of kin discrimination abilities in a variety of fishes.     

Kin selection and ethnic group selection

Ethnicity looks something like kinship on a larger scale. The same math can be used to measure genetic similarity within ethnic/racial groups and relatedness within families. For example, members of the same continental race are about as related (r = 0.18–0.26) as half-siblings (r = 0.25). However (contrary to some claims) the theory of kin selection does not apply straightforwardly to ethnicity, because inclusive fitness calculations based on Hamilton's rule break down when there are complicated social interactions within groups, and/or groups are large and long-lasting. A more promising approach is a theory of ethnic group selection, a special case of cultural group selection. An elementary model shows that the genetic assimilation of a socially enforced cultural regime can promote group solidarity and lead to the regulation of recruitment to groups, and to altruism between groups, based on genetic similarity – in short, to ethnic nepotism. Several lines of evidence, from historical population genetics and political psychology, are relevant here.     

Bonobos Protect and Console Friends and Kin

Post-conflict third-party affiliation has been reported to have different functional meanings, one of them being consolation. Here, we tested the main hypotheses that have been put forth to explain the presence of this phenomenon at a functional level in the bonobo: Self-Protection Hypothesis, Victim-Protection Hypothesis, Relationship-Repair or Substitute for Reconciliation Hypothesis, and Consolation Hypothesis. By analyzing the data collected over 10 years, we investigated what factors affected the distribution of both spontaneous third party affiliation (initiated by the bystander) and solicited third party affiliation (initiated by the victim). We considered factors related to the individual features (sex, rank, age) of victim and bystander, their relationship quality (kinship, affiliation), and the effect that third party affiliation had on the victim (such as protection against further attacks and anxiety reduction). Both spontaneous and solicited third party affiliation reduced the probability of further aggression by group members on the victim (Victim-Protection Hypothesis supported). Yet, only spontaneous affiliation reduced victim anxiety (measured via self-scratching), thus suggesting that the spontaneous gesture – more than the protection itself – works in calming the distressed subject. The victim may perceive the motivational autonomy of the bystander, who does not require an invitation to provide post-conflict affiliative contact. Moreover, spontaneous - but not solicited - third party affiliation was affected by the bond between consoler and victim, being the relationship between consoler and aggressor irrelevant to the phenomenon distribution (Consolation Hypothesis supported). Spontaneous affiliation followed the empathic gradient described for humans, being mostly offered to kin, then friends, then acquaintances. Overall, our findings do not only indicate the consolatory function of spontaneous third-party affiliation but they also suggest that consolation in the bonobo may be an empathy-based phenomenon.     

Kin investment in wage-labor economies

Various human groups, from food foragers to inner-city urban Americans, have used widespread sharing of resources through kin networks as a means of buffering themselves against fluctuations in resource availability in their environments. This paper addresses the effects of progressive incorporation into a wage-labor economy on the benefits of traditional kin networks for two social classes in urban South India. Predictions regarding the effects of kin network wealth, education, and size on child and spouse characteristics and methods of financing marriages are tested using various regression techniques. Despite the rapid growth of participation in a wage-labor economy, it is found that kin network characteristics still have an important impact on investment behavior among families in Bangalore in both social classes. Network wealth is found to have a positive effect on child and spouse characteristics, and large networks are found to act as significant drains on family resources. However, the results for education are broadly consistent with an interpretation of increasing family autonomy as parents’ education has a far stronger influence on child and spouse characteristics across categories than network education does. Finally, professional-class parents are found to prefer financing marriages using formal mechanisms such as savings and bank loans while working-class parents preferentially finance marriages using credit from relatives and friends.     

Evolutionary Stability for Interactions among Kin under Quantitative Inheritance

The theory of evolutionarily stable strategies (ESS) predicts the long-term evolutionary outcome of frequency-dependent selection by making a number of simplifying assumptions about the genetic basis of inheritance. I use a symmetrized multilocus model of quantitative inheritance without mutation to analyze the results of interactions between pairs of related individuals and compare the equilibria to those found by ESS analysis. It is assumed that the fitness changes due to interactions can be approximated by the exponential of a quadratic surface. The major results are the following. (1) The evolutionarily stable phenotypes found by ESS analysis are always equilibria of the model studied here. (2) When relatives interact, one of the two conditions for stability of equilibria differs between the two models; this can be accounted for by positing that the inclusive fitness function for quantitative characters is slightly different from the inclusive fitness function for characters determined by a single locus. (3) The inclusion of environmental variance will in general change the equilibrium phenotype, but the equilibria of ESS analysis are changed to the same extent by environmental variance. (4) A class of genetically polymorphic equilibria occur, which in the present model are always unstable. These results expand the range of conditions under which one can validly predict the evolution of pairwise interactions using ESS analysis.     

Kin recognition and the evolution of altruism

In 1964, Hamilton formalized the idea of kin selection to explain the evolution of altruistic behaviours. Since then, numerous examples from a diverse array of taxa have shown that seemingly altruistic actions towards close relatives are a common phenomenon. Although many species use kin recognition to direct altruistic behaviours preferentially towards relatives, this important aspect of social biology is less well understood theoretically. I extend Hamilton's classic work by defining the conditions for the evolution of kin-directed altruism when recognizers are permitted to make acceptance (type I) and rejection (type II) errors in the identification of social partners with respect to kinship. The effect of errors in recognition on the evolution of kin-directed altruism depends on whether the population initially consists of unconditional altruists or non-altruists (i.e. alternative forms of non-recognizers). Factors affecting the level of these error rates themselves, their evolution and their long-term stability are discussed.