Trustworthy but not lust-worthy: context-specific effects of facial resemblance

If humans are sensitive to the costs and benefits of favouring kin in different circumstances, a strong prediction is that cues of relatedness will have a positive effect on prosocial feelings, but a negative effect on sexual attraction. Indeed, positive effects of facial resemblance (a potential cue of kinship) have been demonstrated in prosocial contexts. Alternatively, such effects may be owing to a general preference for familiar stimuli. Here, I show that subtly manipulated images of other-sex faces were judged as more trustworthy by the participants they were made to resemble than by control participants. In contrast, the effects of resemblance on attractiveness were significantly lower. In the context of a long-term relationship, where both prosocial regard and sexual appeal are important criteria, facial resemblance had no effect. In the context of a short-term relationship, where sexual appeal is the dominant criterion, facial resemblance decreased attractiveness. The results provide evidence against explanations implicating a general preference for familiar-looking stimuli and suggest instead that facial resemblance is a kinship cue to which humans modulate responses in a context-sensitive manner.     

Reactions to children's faces: Males are more affected by resemblance than females are, and so are their brains

The detection of genetic relatedness (i.e., kinship) affects the social, parental, and sexual behavior of many species. In humans, self-referent phenotype matching based on facial resemblance may indicate kinship, and it has been demonstrated that facial resemblance increases perceptions of trustworthiness and attractiveness [Proc. R. Soc. Lond., B Biol. Sci. 269 (2002) 1307–1312; Proc. R. Soc. Lond., B Biol. Sci. (in press)]. However, investigations of sex differences in reaction to facial resemblance have produced mixed results [Evol. Hum. Behav. 25 (2004) 142–154; Evol. Hum. Behav. 23 (2002) 159–166; Evol. Hum. Behav. 24 (2003) 81–87]. Here, we replicate the effects of Platek et al. [Evol. Hum. Behav. 23 (2002) 159–166] using high-resolution color morphing. We also extend these findings using functional magnetic resonance imaging (fMRI) to demonstrate a possible neural mechanism that may account for the observed sex difference. These data support the hypothesis that human males may use and favor facial resemblance as a paternity cue.     

Visual Phenotype Matching: Cues to Paternity Are Present in Rhesus Macaque Faces

The ability to recognize kin and thus behaviourally discriminate between conspecifics based on genetic relatedness is of importance both in acquiring inclusive fitness benefits and to enable optimal inbreeding. In primates, mechanisms allowing recognition of paternal relatives are of particular interest, given that in these mating systems patrilineal information is unlikely to be available via social familiarity. Humans use visual phenotype matching based on facial features to identify their own and other''s close relatives, and recent studies suggest similar abilities may be present in other species. However it is unclear to what extent familial resemblances remain detectable against the background levels of relatedness typically found within demes in the wild – a necessary condition if facial cues are to function in kin recognition under natural conditions. Here, we experimentally investigate whether parent-offspring relationships are discernible in rhesus macaque (Macaca mulatta) faces drawn from a large free-ranging population more representative of the latter scenario, and in which genetic relatedness has been well quantified from pedigrees determined via molecular markers. We used the human visual system as a means of integrating multiple types of facial cue simultaneously, and demonstrate that paternal, as well as maternal, resemblance to both sons and daughters can be detected even by human observers. Experts performed better than participants who lacked previous experience working with nonhuman primates. However the finding that even naïve individuals succeeded at the task underlines the strength of the phenotypic cues present in faces.     

Cues to paternity: Do partner fidelity and offspring resemblance predict daughter-directed sexual aversions?

Despite the profound influence of relatedness on mating and cooperative behavior in humans, the cues men use to assess paternity and guide offspring-directed behavior have yet to be fully resolved. According to leading theories of kin detection, kinship cues should influence both sexual and altruistic motivations because of fitness consequences associated with inbreeding and welfare tradeoff decisions, respectively. Prior work with paternity assessment, however, has generally evaluated candidate cues solely by demonstrating associations with altruism. Here we (i) replicate past work that found effects of phenotypic resemblance and perceived partner fidelity on offspring investment; and (ii) evaluate whether both phenotypic resemblance and perceived partner fidelity meet the more stringent criteria suggested by theory—that is, whether they also predict inbreeding aversions. We report on two studies, one from a population-based sample of Finnish fathers (N?=?390), the other from a Mechanical Turk sample (N?=?700), and furnish evidence in strong support of perceived partner fidelity as a cue to paternity. Support for resemblance as a cue to paternity was decidedly weaker. We discuss a non-kin-based role that resemblance might play in altruistic decision-making, consider whether men might use additional kinship cues to meet the computational challenges associated with paternity assessment, and provide suggestions for future research.     

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.     

Facial resemblance increases the attractiveness of same-sex faces more than other-sex faces

Our reactions to facial self-resemblance could reflect either specialized responses to cues of kinship or by-products of the general perceptual mechanisms of face encoding and mere exposure. The adaptive hypothesis predicts differences in reactions to self-resemblance in mating and prosocial contexts, while the by-product hypothesis does not. Using face images that were digitally transformed to resemble participants, I showed that the effects of resemblance on attractiveness judgements depended on both the sex of the judge and the sex of the face being judged: facial resemblance increased attractiveness judgements of same-sex faces more than other-sex faces, despite the use of identical procedures to manipulate resemblance. A control experiment indicated these effects were caused neither by lower resemblance of other-sex faces than same-sex faces, nor by an increased perception of averageness or familiarity of same-sex faces due to prototyping or mere exposure affecting only same-sex faces. The differential impact of self-resemblance on our perception of same-sex and other-sex faces supports the hypothesis that humans use facial resemblance as a cue of kinship.     

Human Ability to Recognize Kin Visually Within Primates

The assessment of relatedness is a key determinant in the evolution of social behavior in primates. Humans are able to detect kin visually in their own species using facial phenotypes, and facial resemblance in turn influences both prosocial behaviors and mating decisions. This suggests that cognitive abilities that allow facial kin detection in conspecifics have been favored in the species by kin selection. We investigated the extent to which humans are able to recognize kin visually by asking human judges to assess facial resemblance in 4 other primate species (common chimpanzees, western lowland gorillas, mandrills, and chacma baboons) on the basis of pictures of faces. Humans achieved facial interspecific kin recognition in all species except baboons. Facial resemblance is a reliable indicator of relatedness in at least chimpanzees, gorillas, and mandrills, and future work should explore if the primates themselves also share the ability to detect kin facially.     

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.     

Facial resemblance enhances trust

Organisms are expected to be sensitive to cues of genetic relatedness when making decisions about social behaviour. Relatedness can be assessed in several ways, one of which is phenotype matching: the assessment of similarity between others' traits and either one's own traits or those of known relatives. One candidate cue of relatedness in humans is facial resemblance. Here, I report the effects of an experimental manipulation of facial resemblance in a two-person sequential trust game. Subjects were shown faces of ostensible playing partners manipulated to resemble either themselves or an unknown person. Resemblance to the subject's own face raised the incidence of trusting a partner, but had no effect on the incidence of selfish betrayals of the partner's trust. Control subjects playing with identical pictures failed to show such an effect. In a second experiment, resemblance of the playing partner to a familiar (famous) person had no effect on either trusting or betrayals of trust.     

Does Facial Resemblance Enhance Cooperation?

Facial self-resemblance has been proposed to serve as a kinship cue that facilitates cooperation between kin. In the present study, facial resemblance was manipulated by morphing stimulus faces with the participants'' own faces or control faces (resulting in self-resemblant or other-resemblant composite faces). A norming study showed that the perceived degree of kinship was higher for the participants and the self-resemblant composite faces than for actual first-degree relatives. Effects of facial self-resemblance on trust and cooperation were tested in a paradigm that has proven to be sensitive to facial trustworthiness, facial likability, and facial expression. First, participants played a cooperation game in which the composite faces were shown. Then, likability ratings were assessed. In a source memory test, participants were required to identify old and new faces, and were asked to remember whether the faces belonged to cooperators or cheaters in the cooperation game. Old-new recognition was enhanced for self-resemblant faces in comparison to other-resemblant faces. However, facial self-resemblance had no effects on the degree of cooperation in the cooperation game, on the emotional evaluation of the faces as reflected in the likability judgments, and on the expectation that a face belonged to a cooperator rather than to a cheater. Therefore, the present results are clearly inconsistent with the assumption of an evolved kin recognition module built into the human face recognition system.     

Kin and kinship psychology both influence cooperative coordination in Yasawa,Fiji

Genes shared through common ancestry are among the oldest social bonds. Despite these ancient roots, humans often co-opt the psychology of genetic relatedness and extend it to genetically unrelated others through culturally-acquired kinship systems. We investigate how genealogical relatedness and kinship norms might mutually support or oppose each other within a known kin network in Yasawa, Fiji. Yasawans' reliance on intensive, kin-based cooperation for daily life makes Yasawan kinship an interesting test case to compare the effects of genealogy and kinship norms. Confirming qualitative ethnographic claims, we find that Yasawan kin terms can be described in two dimensions of respect/closeness and joking/authority. Individual players use different strategies for genealogical relatives and non-relatives by making economic game choices that are increasingly beneficial to partners who share a higher percentage of genes through common ancestry. However, pairs of players are most successful in coordinating their game choices despite conflicting self-interests based upon kinship norms relevant to hierarchy. Thus, while genealogical relatedness may boost generosity, the extra behavioral structuring from kinship norms facilitates more productive but difficult coordinated action even when communication is not possible.     

Women's attractiveness judgments of self-resembling faces change across the menstrual cycle

Two lines of reasoning predict that women's preferences for people exhibiting cues to kinship will be lower in the follicular phase than in the luteal phase of the menstrual cycle. Women may avoid kinship cues during the follicular phase when they are most fertile due to the costs of inbreeding. Alternatively, women may seek kinship cues during the luteal phase as a byproduct of the benefits of associating with kin during pregnancy, which is also characterized by high progesterone. We find that preferences for facial resemblance, a putative kinship cue, follow this predicted pattern and are positively correlated with estimated progesterone levels based on cycle day. Neither estimated estrogen levels nor conception risk predicted preferences for self-resemblance, and the cyclic shift was stronger for preferences for female faces than male faces. These findings lead to the possibility that this cyclic change in preference for self-resemblance may be a byproduct of a hormonal mechanism for increasing affiliative behavior toward kin during pregnancy rather than a mechanism for preventing inbreeding during fertile periods.     

Kinship affects investment by helpers in a cooperatively breeding bird

Helping behaviour in cooperative breeding systems has been attributed to kin selection, but the relative roles of direct and indirect fitness benefits in the evolution of such systems remain a matter of debate. In theory, helpers could maximize the indirect fitness benefits of cooperation by investing more in broods with whom they are more closely related, but there is little evidence for such fine-scale adjustment in helper effort among cooperative vertebrates. In this study, we used the unusual cooperative breeding system of the long-tailed tit Aegithalos caudatus to test the hypothesis that the provisioning effort of helpers was positively correlated with their kinship to broods. We first use pedigrees and microsatellite genotypes to characterize the relatedness between helpers and breeders from a 14 year field study. We used both pedigree and genetic approaches because long-tailed tits have access to pedigree information acquired through social relationships, but any fitness consequences will be determined by genetic relatedness. We then show using both pedigrees and genetic relatedness estimates that alloparental investment by helpers increases as their relatedness to the recipients of their care increases. We conclude that kin selection has played a critical role in moulding the investment decisions of helpers in this cooperatively breeding species.     

The Genetic Relatedness in Groups of Joint-Nesting Taiwan Yuhinas: Low Genetic Relatedness with Preferences for Male Kin

The relative importance of direct and indirect fitness and, thus, the role of kinship in the evolution of social behavior is much debated. Studying the genetic relatedness of interacting individuals is crucial to improving our understanding of these issues. Here, we used a seven-year data set to study the genetic structure of the Taiwan yuhina (Yuhina brunneciceps), a joint-nesting passerine. Ten microsatellite loci were used to investigate the pair-wised relatedness among yuhina breeding group members. We found that the average genetic relatedness between same-sex group members was very low (0.069 for male dyads and 0.016 for female dyads). There was also a low ratio of closely-related kin (r>0.25) in the cooperative breeding groups of yuhinas (21.59% and 9.68% for male and female dyads, respectively). However, the relatedness of male dyads within breeding groups was significantly higher than female dyads. Our results suggest that yuhina cooperation is maintained primarily by direct fitness benefits to individuals; however, kin selection might play a role in partner choice for male yuhinas. Our study also highlights an important, but often neglected, question: Why do animals form non-kin groups, if kin are available? We use biological market theory to propose an explanation for group formation of unrelated Taiwan yuhinas.     

Sex-specific associative learning cues and inclusive fitness benefits in the Seychelles warbler

In cooperative breeding vertebrates, indirect fitness benefits would be maximized by subordinates that accurately assess their relatedness to group offspring and preferentially help more closely related kin. In the Seychelles warbler (Acrocephalus sechellensis), we found a positive relationship between subordinate-nestling kinship (determined using microsatellite marker genotypes) and provisioning rates, but only for female subordinates. Female subordinates that helped were significantly more related to the nestlings than were nonhelpers, and the decision to help appears to be based on associative learning cues. High levels of female infidelity means that subordinates cannot trust their legitimacy through the male line, consequently they appear to use the continued presence of the primary female, but not the primary male, as a reliable cue to determine when to feed nestlings. By using effective discrimination, female subordinates are able to maximize the indirect benefits gained within a cooperative breeding system otherwise driven primarily by direct breeding benefits.     

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.     

Differential facial resemblance of young children to their parents: who do children look like more?

In humans, paternal investment is highly variable and is modulated by paternity uncertainty. Facial phenotypic similarity between a father and a child is one possible paternity indicator. However, whether such paternal-biased traits are expressed in children is unclear, as previous empirical results are contradictory. Therefore, we quantified the facial resemblance between a child and each of his or her parents, from birth to 6 years old. Resemblance was assessed from pictures of the face by nonrelated judges. We found that, at all ages, children resemble both their parents more than would be expected by chance, although there is a differential resemblance toward one or the other parent depending on the age and sex of the child. For newborns, boys and girls resemble their mothers more, this differential resemblance persisting through time for girls. For boys, an inversion occurs and they resemble their fathers more between 2 and 3 years of age. The resemblance ascribed by the parents shows that, at birth, mothers ascribe a resemblance to the father, as previously found, although assessment by external judges revealed the opposite. These results suggest that facial appearance is a cue for kin recognition between a father and a child. Patterns of differential resemblance are discussed within the context of evolutionary theories on parental investment.     

Whom do the sparrows follow? The effect of kinship on social preference in house sparrow flocks

Non-aggressive social interactions between group-mates, e.g. maintenance of spatial proximity or activity synchrony are basic elements of a species’ social structure, and were found to be associated with important fitness consequences in group-living animals. In the establishment of such affiliative relationships, kinship has often been identified as one of the key predictors, but this has rarely been studied in simple social groups such as flocks of gregarious birds. In this study we investigated whether kinship affects social preference, as measured by the tendency to associate with others during various social activities, in captive house sparrow (Passer domesticus) flocks where birds could interact with differently related flock-mates. We found that preference between flock-mates was correlated with familiarity from early nestling period: same-brood siblings followed their sib initiating new activities more often than non-sib birds. The strength of association between birds also tended to correlate with genetic relatedness, but this was mainly due to the effect of siblings’ affiliation. Thus we concluded that house sparrows prefer the company of their siblings during social activities even well after fledging, which may facilitate kin-biased behaviours.     

Parent's expectations and experiences of resemblance through donor conception

In the field of donor conception, matching is a practice that involves choosing gametes or embryos in such a way as to minimize physical differences between the donor and the recipients. This practice serves to create resemblance between child and social parent/s, to meet the societal norm of biological relationship and legitimize the family form.

This study focused on parents’ experiences and expectations of the clinical practice of matching and (non)resemblance with their donor-offspring. Results are based on accounts of 18 heterosexual donor-conceiving parents (11 couples) with a donor-offspring between 8 and 32 years old.

Findings suggest that resemblance is more than just a strategy to achieve social legitimization of the donor-conceived family. Resemblance also plays a role in the creation of family bonds and kinship. This suggests that the social reasoning behind matching should at least be complemented by a psychological understanding of relatedness and connectedness.