Altruism and reciprocity among friends and kin in a Tibetan village

To explain the high level of altruism among unrelated friends, some people propose that friends may apply the feeling and emotions that underlie kin-selected nepotism to each other. Alternatively, altruism among friends may be sustained by an evolved psychology that is sensitive to the dynamic of contingent reciprocity. A recent study (Stewart-Williams, S., 2007. Altruism among kin vs. nonkin: effects of cost of help and reciprocal exchange. Evolution and Human Behavior, 28, 193–198) with North American students showed that the amount of help given and received was less likely to be balanced among friends (as among relatives) than among acquaintances, suggesting that exchange among friends and relatives may rely on similar psychological mechanisms. To assess whether the similar pattern in exchange reflects the similar psychology in exchange, I tested how different types of relationship regulated the amount of help given and received, and people’s emotional reaction to non-reciprocation. I asked 45 participants from a Tibetan village in China to imagine how unhappy they would feel if the target person (sibling, cousin, friend or acquaintance) failed to reciprocate their help in various contexts. I found that overall, friends and relatives were more tolerant to non-reciprocation than acquaintances were. For emotional help, friends were more likely to feel unhappy about non-reciprocation than relatives were, but were similar to relatives in responses to other types of help (aid during crisis, labor, and financial help). The study suggests that people may evaluate the importance of reciprocity differently in various situations, and exhibit different levels of sensitivity to the dynamic of reciprocity. Thus it calls for careful distinctions between friends and kin in the everyday lives of individuals.     

Human Beings as Evolved Nepotists

Inclusive fitness theory provides a compelling explanation for the evolution of altruism among kin. However, a completely satisfactory account of non-kin altruism is still lacking. The present study compared the level of altruism found among siblings with that found among friends and mates and sought to reconcile the findings with an evolutionary explanation for human altruism. Participants (163 males and 156 females) completed a questionnaire about help given to a sibling, friend, or mate. Overall, participants gave friends and mates as much or more help than they gave siblings. However, as the cost of help increased, siblings received a progressively larger share of the help, whereas friends and mates received a progressively smaller share, despite the fact that participants were closer emotionally to friends and mates than they were to siblings. These findings help to explain the relative standing of friends and mates as recipients of altruistic aid.

Kin recognition and cannibalism in polyphenic salamanders

We investigated kin discrimination among larvae of Arizona tigersalamanders (Ambystoma tigrinum nebulosum) which occur as "typical"morphs that feed mostly on invertebrate prey and occasionallyon conspecifics, and as "cannibal" morphs that feed primarilyon conspecifics. When housed with smaller larvae that differedin relatedness, both cannibals and typicals preferentially consumedless-related individuals. Cannibals ate typicals much quickerwhen the choice was between nonkin and siblings than when thechoice was between nonkin and cousins, indicating that cannibalscould distinguish different categories of relatives. Cannibalswere less likely to eat a larval sibling that was a cannibalmorph than a sibling that was a typical morph. Occluding animals'nares temporarily eliminated kin discrimination, implying thatolfaction is important in recognition. Larvae from differentsibships varied considerably in their ability to discriminatekin, and the greater the probability that a larva from a givensibship would develop into a cannibal morph, the more likelythe members of that sibship were to discriminate kin. Our resultsenable us to infer the functional significance of kin recognitionin this species and to develop an evolutionary model of themechanisms underlying the joint control of kin recognition andcannibalistic polyphenism.     

Evolution of indirect reciprocity by social information: the role of trust and reputation in evolution of altruism

The complexity of human's cooperative behavior cannot be fully explained by theories of kin selection and group selection. If reciprocal altruism is to provide an explanation for altruistic behavior, it would have to depart from direct reciprocity, which requires dyads of individuals to interact repeatedly. For indirect reciprocity to rationalize cooperation among genetically unrelated or even culturally dissimilar individuals, information about the reputation of individuals must be assessed and propagated in a population. Here, we propose a new framework for the evolution of indirect reciprocity by social information: information selectively retrieved from and propagated through dynamically evolving networks of friends and acquaintances. We show that for indirect reciprocity to be evolutionarily stable, the differential probability of trusting and helping a reputable individual over a disreputable individual, at a point in time, must exceed the cost-to-benefit ratio of the altruistic act. In other words, the benefit received by the trustworthy must out-weigh the cost of helping the untrustworthy.     

Egg-dumping lace bugs preferentially oviposit with kin

Egg dumping, or abandoning eggs and young to the care of other conspecifics, results in an extreme form of alloparental care. It is unclear, however, if egg dumpers discriminate among kin and nonkin egg recipients. In the lace bug Gargaphia solani (Heteroptera: Tingidae), some females with eggs (guards) also accept and defend eggs of conspecifics. Other females (egg dumpers) abandon their offspring after oviposition, leaving a single guard as the caregiver. We asked if egg dumpers preferentially dump their eggs among unguarded eggs of kin or nonkin. When given a choice between dumping among eggs of full siblings and eggs of nonsiblings, most eggs (67%) were dumped with full siblings' eggs. Furthermore, egg dumpers were just as likely to oviposit among eggs of kin with whom they had interacted on a shared host plant during juvenile development as they were to oviposit with kin reared on different host plants. Thus, egg dumpers discriminate kin by using cues associated with eggs, and such cues are not likely to be acquired through interaction on a common host plant environment. Copyright 2000 The Association for the Study of Animal Behaviour.     

Caring across Boundaries versus Keeping Boundaries Intact: Links between Moral Values and Interpersonal Orientations

Prior work has established robust diversity in the extent to which different moral values are endorsed. Some people focus on values related to caring and fairness, whereas others assign additional moral weight to ingroup loyalty, respect for authority and established hierarchies, and purity concerns. Five studies explore associations between endorsement of distinct moral values and a suite of interpersonal orientations: Machiavellianism, prosocial resource distribution, Social Dominance Orientation, and reported likelihood of helping and not helping kin and close friends versus acquaintances and neighbors. We found that Machiavellianism (Studies 1, 3, 4, 5) (e.g., amorality, controlling and status-seeking behaviors) and Social Dominance Orientation (Study 4) were negatively associated with caring values, and positively associated with valuation of authority. Those higher in caring values were more likely to choose prosocial resource distributions (Studies 2, 3, 4) and to report reduced likelihood of failing to help kin/close friends or acquaintances (Study 4). Finally, greater likelihood of helping acquaintances was positively associated with all moral values tested except authority values (Study 4). The current work offers a novel approach to characterizing moral values and reveals a striking divergence between two kinds of moral values in particular: caring values and authority values. Caring values were positively linked with prosociality and negatively associated with Machiavellianism, whereas authority values were positively associated with Machiavellianism and Social Dominance Orientation.     

Differential responses of kin and nonkin salmon to patterns of water flow: does recirculation influence aggression?

Studies conducted in laboratory streams have shown that juvenile salmonid fish (parr) can differentiate between their kin and nonkin and may be less aggressive towards their kin. Chemicals produced by salmonids are also known to be used as cues to aid kin recognition. We tested the hypothesis that the ability of Atlantic salmon, Salmo salar, to recognize kin and hence modulate their level of aggression is influenced by the amount of water recirculation, which would be expected to affect the concentration of odour. Levels of aggression were similar between pairs of kin and pairs of nonkin when there was negligible recirculation of water. However, when water was recirculated, pairs of nonkin were on average 1.56 times more aggressive than pairs of kin, owing to an increase in attacks by the dominant fish. The data do not support the idea that odour concentration affects kin recognition and hence reduces aggression among siblings. Instead they indicate that recirculation of water instigates heightened aggression in dominant fish but only towards nonkin subordinates. The study suggests that the advantages for juvenile salmonids of associating with kin vary spatially, being influenced by water flow dynamics. Copyright 2000 The Association for the Study of Animal Behaviour.     

Coalition formation among male Barbary macaques (Macaca sylvanus)

A coalition is formed when one animal intervenes in an ongoing conflict between two parties to support one side. Since support of one party is also an act against the other party, coalitions are triadic interactions involving a supporter, a recipient, and a target. The purpose of this study was to test which of three possible theories explains coalition formation among male Barbary macaques: 1) Males support kin to enhance their indirect fitness (kin selection). 2) Males support nonkin to receive future reciprocal support (reciprocal altruism). 3) Males pursue self-interests and immediately benefit via nonkin support (cooperation). Coalition formation was investigated among 31 semi-free male Barbary macaques in the Salem Monkey Park, Germany during the mating season. The results show: 1) Males intervened more often in dyadic conflicts in which a related opponent was involved and supported related opponents more than unrelated opponents. Close kin supported each other more often than distant kin. 2) Some evidence for reciprocal support was found. However, reciprocity was probably a by-product of targeting the same individuals for dominance. 3) Coalition formation among nonkin is best interpreted as cooperation, based on self-interests. Male Barbary macaques seem to intervene more often to stabilize and less often to improve their rank. Although our data were limited, the results revealed that kin support, reciprocal support, and cooperative support were all involved in coalition formation among male Barbary macaques.     

Testing kin altruism in long-tailed macaques (Macaca fascicularis) in a food-sharing experiment

Social grooming and coalition formation have been main foci in studies concerning altruism in monkeys. Results have been inconclusive because the altruistic nature of these behaviors remains unclear. I investigated altruism in a more transparent context via an apparatus in which captive long-tailed macaque females had the choice to exploit a food source themselves or to yield the food to a test partner. I hypothesized that if potential donors behaved altruistically toward kin, they would yield the food sources longer to kin than to nonkin. Of 11 tested potential donors, 8 did not discriminate between daughters or sisters and nonkin. Thus, the experiments do not support the kin altruism hypothesis. Three females let their youngest offspring have more food than matched juvenile nonrelatives. Behavioral observations strongly suggested,however, that it was primarily spiteful behavior toward juvenile nonrelatives that caused the differences between kin and nonkin in these three cases.     

Reduced plant competition among kin can be explained by Jensen's inequality

Plants often compete with closely related individuals due to limited dispersal, leading to two commonly invoked predictions on competitive outcomes. Kin selection, from evolutionary theory, predicts that competition between relatives will likely be weaker. The niche partitioning hypothesis, from ecological theory, predicts that competition between close relatives will likely be stronger. We tested for evidence consistent with either of these predictions by growing an annual legume in kin and nonkin groups in the greenhouse. We grew plant groups in treatments of symbiotic nitrogen fixing bacteria differing in strain identity and composition to determine if differences in the microbial environment can facilitate or obscure plant competition patterns consistent with kin selection or niche partitioning. Nonkin groups had lower fitness than expected, based on fitness estimates of the same genotypes grown among kin. Higher fitness among kin groups was observed in mixtures of N‐fixing bacteria strains compared to single inoculations of bacteria strains present in the soil, which increased fitness differences between kin and nonkin groups. Lower fitness in nonkin groups was likely caused by increased competitive asymmetry in nonkin groups due to genetic differences in plant size combined with saturating relationships with plant size and fitness‐ i.e. Jensen's inequality. Our study suggests that microbial soil symbionts alter competitive dynamics among kin and nonkin. Our study also suggests that kin groups can have higher fitness, as predicted by kin selection theory, through a commonly heritable trait (plant size), without requiring kin recognition mechanisms.     

Reproductive competition between females in the matrilineal Mosuo of southwestern China

The matrilineal Mosuo of southwestern China live in communal households where brothers and sisters of three generations live together (duolocal residence), and men visit their wives, who reside elsewhere, only at night in ‘visiting’ marriages. Here we show that these communally breeding sisters are in reproductive conflict, in the sense that they share the resources needed to reproduce. We analyse determinants of reproductive success in females and males, and show that co-resident female kin are in competition; the more female kin reside in the household, the more reproductive success is reduced. Male reproductive success, however, is not determined by the kin in his natal household; duolocal males are not in reproductive conflict with their siblings. Competition with female cousins can be worse than that between sisters. We also find that female work on the farm (which is the main communal resource) is not equal. We use a ‘tug-of-war’ model of reproductive skew generated by incomplete control, to model the patterns of effort put into competition between sisters and cousins. The model predicts that more dominant (older) sisters will put less effort into reproductive conflict than will less dominant (younger) sisters; but younger sisters will also have lower reproductive success because they are less efficient at gaining access to the shared resource. Both predictions are consistent with our data. Younger sisters work less in the fields than do older sisters, which may represent a form of conflict or may be because their average relatedness to the household is lower than that of their more fertile older sisters.     

Kin discrimination and sex ratios in a parasitoid wasp

Sex ratio theory provides a clear and simple way to test if nonsocial haplodiploid wasps can discriminate between kin and nonkin. Specifically, if females can discriminate siblings from nonrelatives, then they are expected to produce a higher proportion of daughters if they mate with a sibling. This prediction arises because in haplodiploids, inbreeding (sib-mating) causes a mother to be relatively more related to her daughters than her sons. Here we formally model this prediction for when multiple females lay eggs in a patch, and test it with the parasitoid wasp Nasonia vitripennis. Our results show that females do not adjust their sex ratio behaviour dependent upon whether they mate with a sibling or nonrelative, in response to either direct genetic or a range of indirect environmental cues. This suggests that females of N. vitripennis cannot discriminate between kin and nonkin. The implications of our results for the understanding of sex ratio and social evolution are discussed.     

Kin recognition and cannibalism in a subsocial spider

Evolution of cooperation and group living in spiders from subsocial family groups may be constrained by their cannibalistic nature. A tendency to avoid cannibalizing kin may facilitate tolerance among spiders and implies the ability to identify relatives. We investigated whether the subsocial spider Stegodyphus lineatus discriminates kin by recording cannibalism among juveniles in experiments during which amount of food and size difference among spiders in groups were varied. We hypothesized that family groups should be less cannibalistic than groups of mixed‐parental origin. Further, we tested whether food‐stress would influence cannibalism rates differently in kin and nonkin groups and the effect of relatedness on cannibalism within groups of spiders of variable size compared with those of homogenous size. In groups of six spiders, more spiders were cannibalized in nonsib groups than in sib groups under low food conditions. A tendency for nonkin biased cannibalism in starved spider pairs supported that kin recognition in S. lineatus is expressed when food is limited. Size variance of individuals within well‐fed groups of siblings and unrelated spiders had no influence on cannibalism rates. Apparently, both hunger and high density are important promoters of cannibalism. In addition to inclusive fitness benefits, we suggest that an ability to avoid cannibalizing kin will favour the evolution of cooperation and group living in phylogenetically pre‐adapted solitary species.     

Offspring retention in the Siberian jay {Perisoreus infaustus): the prolonged brood care hypothesis

We examined aggressive behavior in Siberian jay groups containingboth retained offspring and immigrant juveniles during winterfeeding and during breeding. Selective tolerance of retainedoffspring by parental birds in winter suggests that cooperationevolved through kin selection. Parents exhibited a self-restraintin aggression towards retained offspring at food in winter.Comparatively, nonkin immigrants were aggressively preventedfrom sharing food by the local pair. Parental tolerance in wintercould bring inclusive fitness gains through the direct kin componentif retained offspring experience relaxed competition and enhancedsurvival. Parental tolerance would then favor the evolutionof delayed dispersal. There is no evidence that delayed dispersalamong Siberian jays should have evolved because of indirectfitness benefits to retained offspring from helping to raiseyounger siblings. Offspring retained by parents did not participatein incubation, feeding or nest defense.     

Promiscuity drives self-referent kin recognition

Kin selection theory has been one of the most significant advances in our understanding of social behavior . However, the discovery of widespread promiscuity has challenged the evolutionary importance of kin selection because it reduces the benefit associated with helping nestmates . This challenge would be resolved if promiscuous species evolved a self-referent kin-recognition mechanism that enables individuals to differentiate kin and nonkin . Here, we take advantage of an asymmetry in the level of promiscuity among males of alternative life histories in the bluegill sunfish (Lepomis macrochirus). We show that, as a consequence of this asymmetry, offspring of "parental" males have a high level of relatedness to nestmates, whereas offspring of "cuckolder" males have a low level of relatedness to nestmates. We find that offspring of parentals do not use a direct recognition mechanism to discriminate among nestmates, whereas offspring of cuckolders use kin recognition by self-referent phenotype matching to differentiate between kin and nonkin. Furthermore, we estimate that the cost of utilizing such self-referent kin recognition is equivalent to a relatedness (R) of at least 0.06. These results provide compelling evidence for adaptive use of kin recognition by self-referent phenotype matching and confirm the importance of kinship in social behavior.     

Social behavior of infant and mother Japanese Macaques (Macaca fuscata): Effects of kinship,partner sex,and infant sex

The behavioral interactions of 22 infant and mother Japanese macaques with other group members were studied. Focal-animal observations were made from the time of each infant’s birth until 1 year of age. Infants and mothers both displayed exceedingly strong preferences for associating with matrilineal kin and, specifically, for female kin. The degree of genetic relatedness was positively correlated with levels of spatial proximity, contact, grooming, aggression, and play. Overall frequencies of interactions with nonkin were very low, and partner sex was not an important factor in interactions with nonkin. There were no significant differences between male and female infants in interactions with kin versus nonkin. There was only one significant difference between male and female infants in interactions with males versus females: female infants showed stronger preferences for initiating proximity with females over males than did male infants. Because mothers provide the focal point for infant interactions during the first year of life, we compared the behavior of infants and mothers. Mothers were the recipients of more social interactions than were infants, mothers engaged in more grooming than did infants, and infants engaged in more social play than did mothers. These findings are only partially consistent with kin-selection theory, and the inadequacies of studying matrilineal kin discrimination to test kin selection are reviewed. The near-absence of infant sex differences in associations with social partners suggests that although maternal kin other than the mother are important to infant socialization, they probably do not contribute to the development of behavioral sex differences until after the first year of life.     

Correlated pay-offs are key to cooperation

The general belief that cooperation and altruism in social groups result primarily from kin selection has recently been challenged, not least because results from cooperatively breeding insects and vertebrates have shown that groups may be composed mainly of non-relatives. This allows testing predictions of reciprocity theory without the confounding effect of relatedness. Here, we review complementary and alternative evolutionary mechanisms to kin selection theory and provide empirical examples of cooperative behaviour among unrelated individuals in a wide range of taxa. In particular, we focus on the different forms of reciprocity and on their underlying decision rules, asking about evolutionary stability, the conditions selecting for reciprocity and the factors constraining reciprocal cooperation. We find that neither the cognitive requirements of reciprocal cooperation nor the often sequential nature of interactions are insuperable stumbling blocks for the evolution of reciprocity. We argue that simple decision rules such as ‘help anyone if helped by someone’ should get more attention in future research, because empirical studies show that animals apply such rules, and theoretical models find that they can create stable levels of cooperation under a wide range of conditions. Owing to its simplicity, behaviour based on such a heuristic may in fact be ubiquitous. Finally, we argue that the evolution of exchange and trading of service and commodities among social partners needs greater scientific focus.     

Reciprocal altruism in bats and other mammals

In this paper five conditions are specified which must be met before reciprocal altruism, rather than kin selection, should be invoked. Four purported mammalian examples— social grooming in coati, cluster position in roosting pallid bats, information exchange among greater spear-nosed bats, and blood regurgitation among vampire bats—are examined to determine if reciprocal altruism is necessary to plausibly explain each situation. Results from a computer simulation which apportions the relative selective advantage of vampire bat food sharing to kin selection and reciprocal altruism are then presented. The results demonstrate that the increase in individual survivorship due to reciprocal food sharing events in this species provides a greater increase in inclusive fitness than can be attributed to aiding relatives. This analysis suggests that reciprocal altruism can be selectively more important than kin selection when altruistic behaviors in a relatively large social group occur frequently and provide a major fitness benefit to the recipient even when that recipient is related to the donor.     

Consanguineous marriage among the Fulani

The Fulani are a broad ethnic category of nomadic and seminomadic pastoralists and agropastoralists living in the semiarid Sahel region of sub-Saharan Africa. The Fulani are patrilineal, patrilocal, and moderately polygynous, with arranged first marriages accompanied by the payment of bridewealth, ideally in the form of cattle. Consanguineous marriage is frequent, with first or second cousin marriage preferred. In this paper we present data on levels of consanguineous marriage among the Fulani of northern Burkina Faso and test the hypothesis that inbreeding may be more frequent when there is a scarcity of cattle available, since bridewealth demands are thought to be reduced with close-kin marriage. Among 308 women's marriages, 203 (65.8%) were between kin up to and including second cousins, and 102 (33.1%) were between nonkin. Among 276 men's marriages, 196 (71.0%) were between kin up to and including second cousins, and 77 (27.9%) were between nonkin. The mean population inbreeding coefficient (alpha) was 0.0355 for women, and 0.0374 for men. No increase was found in population levels of inbreeding estimated from marriages contracted after the droughts of 1973 and 1984, which drastically reduced the Fulani's cattle stocks. However, a significantly higher rate of consanguineous marriage was found in families owning the fewest cattle.     

Absence of strongly kin-preferential behavior by adult female sooty mangabeys (Cercocebus atys)

The objective of this study was to test the hypothesis that kin-preferential behavior would be present in sooty mangabeys (Cercocebus atys), a species taxonomically close to baboons and macaques. The affiliative behavior of the adult female members of a large captive group of these mangabeys was analyzed to test this prediction. These females groomed, were approached by, were in proximity to, and were in contact with their kin significantly more than expected, but only when all kin were included in the analysis. Removal of only the mother-infant (less than 1 year) dyadic interactions removed all significant kin effects. Kin-preferential behavior was also absent in affiliative interactions among the adult females as a class. Affiliation between mothers and offspring significantly exceeded that for all other kinship categories (such as siblings, etc.), and affiliation with kin other than offspring did not differ from that with nonkin adults. In their interactions with nonkin, the adult females showed some preference (duration of grooming) for those adult females of similar age, an effect predicted by the intensity of interaction among members of the same age cohort during development. These similar-aged females may also be paternally related. In comparing these results with the existing literature on kin-preferential behavior, two conclusions may be reached: (1) age and degree of kinship are primary factors that must be considered in order to avoid the existing overgeneralization of the importance of kinship in primate social organization, and (2) the role and importance of affiliative behavior in kin-selection theory should perhaps be re-examined in light of questions raised by this study.