Contextual cues of genetic relatedness to familiar individuals, such as cosocialization and maternal-perinatal association, modulate prosocial and inbreeding-avoidance behaviors toward specific potential siblings. These findings have been interpreted as evidence that contextual cues of kinship indirectly influence social behavior by affecting the perceived probability of genetic relatedness to familiar individuals. Here, we test a more general alternative model in which contextual cues of kinship can influence the kin-recognition system more directly, changing how the mechanisms that regulate social behavior respond to cues of kinship, even in unfamiliar individuals for whom contextual cues of kinship are absent. We show that having opposite-sex siblings influences inbreeding-relevant perceptions of facial resemblance but not prosocial perceptions. Women with brothers were less attracted to self-resembling, unfamiliar male faces than were women without brothers, and both groups found self-resemblance to be equally trustworthy for the same faces. Further analyses suggest that this effect is driven by younger, rather than older, brothers, consistent with the proposal that only younger siblings exhibit the strong kinship cue of maternal-perinatal association. Our findings provide evidence that experience with opposite-sex siblings can directly influence inbreeding-avoidance mechanisms and demonstrate a striking functional dissociation between the mechanisms that regulate inbreeding and the mechanisms that regulate prosocial behavior toward kin.

Animals, including humans, use olfaction to assess potential social and sexual partners. Although hormones modulate olfactory cues, we know little about whether contraception affects semiochemical signals and, ultimately, mate choice. We examined the effects of a common contraceptive, medroxyprogesterone acetate (MPA), on the olfactory cues of female ring-tailed lemurs (Lemur catta), and the behavioural response these cues generated in male conspecifics. The genital odorants of contracepted females were dramatically altered, falling well outside the range of normal female variation: MPA decreased the richness and modified the relative abundances of volatile chemicals expressed in labial secretions. Comparisons between treatment groups revealed several indicator compounds that could reliably signal female reproductive status to conspecifics. MPA also changed a female's individual chemical 'signature', while minimizing her chemical distinctiveness relative to other contracepted females. Most remarkably, MPA degraded the chemical patterns that encode honest information about genetic constitution, including individual diversity (heterozygosity) and pairwise relatedness to conspecifics. Lastly, males preferentially investigated the odorants of intact over contracepted females, clearly distinguishing those with immediate reproductive potential. By altering the olfactory cues that signal fertility, individuality, genetic quality and relatedness, contraceptives may disrupt intraspecific interactions in primates, including those relevant to kin recognition and mate choice.

There is increasing evidence that human body odors are involved in adaptive behaviors, such as parental attachment in infants or partner choice in adults. The aim of the present study was to investigate changes in body-odor perception around puberty, a period largely ignored for odor-mediated behavioral changes, despite major changes in social needs and in odor emission and perception. Nine families with two children (8 pre-pubertal, aged 7-10, and 10 pubertal, aged 11-18) evaluated body odors of family members and unfamiliar individuals for pleasantness, intensity, and masculinity, and performed a recognition task. The hypothesized emergence of a parent-child mutual aversion for the odor of opposite-sex family members at puberty was not found, contradicting one of the few studies on the topic (Weisfeld et al., J. Exp. Child Psychol. 85:279-295, 2003). However, some developmental changes were observed, including reduced aversion for odor of the same-sex parent, and increased ability of adults, compared to children, to recognize odor of family members. Sex and personality (depressive and aggressive traits) also significantly influenced odor judgments. Further research with larger samples is needed to investigate the poorly explored issue of how olfactory perception of self and family members develops, and how it could correlate with normal reorganizations in social interactions at adolescence.

Abstract Microbiota on the human skin plays a major role in body odour production. The human microbial and chemical signature displays a qualitative and quantitative correlation. Genes may influence the chemical signature by shaping the composition of the microbiota. Recent studies on human skin microbiota, using 16S rRNA gene sequencing, found a high inter- and intrapersonal variation in bacterial species on the human skin, which is relatively stable over time. Human body odours mediate the attraction of mosquitoes to their blood hosts. Odours produced by skin microbiota are attractive to mosquitoes as shown by in vitro studies, and variation in bacterial species on the human skin may explain the variation in mosquito attraction between humans. Detailed knowledge of the ecology and genetics of human skin microbiota is needed in order to unravel the evolutionary mechanisms that underlie the interactions between mosquitoes and their hosts.

The assessment of relatedness may be crucial in the evolution of socio-sexual behaviour, because it can be associated with fitness benefits mediated by both nepotism and inbreeding avoidance. In this context, one proposed mechanism for kin recognition is 'phenotype matching'; animals might compare phenotypic similarities between themselves and others in order to assess the probability that they are related. Among cues potentially used for kin discrimination, body odours constitute interesting candidates that have been poorly investigated in anthropoid primates so far, because of a mixture of theoretical considerations and methodological/experimental constraints. In this study, we used an indirect approach to examine the similarity in odour signals emitted by related individuals from a natural population of chacma baboons (Papio ursinus). For that purpose, we designed an innovative behavioural tool using mice olfactory abilities in a habituation-discrimination paradigm. We show that:(i) mice can detect odour differences between individuals of same sex and age class in another mammal species, and (ii) mice perceive a higher odour similarity between related baboons than between unrelated baboons. These results suggest that odours may play a role in both the signalling of individual characteristics and of relatedness among individuals in an anthropoid primate. The 'biological olfactometer' developed in this study offers new perspectives to the exploration of olfactory signals from a range of species.

Many species produce odor cues that enable them to be identified individually, as well as providing other socially relevant information. Study of the role of odor cues in the social behavior of great apes is noticeable by its absence. Olfaction has been viewed as having little role in guiding behavior in these species. This study examined whether Western lowland gorillas produce an individually identifiable odor. Odor samples were obtained by placing cloths in the gorilla's den. A delayed matching to sample task was used with human participants (n = 100) to see if they were able to correctly match a target odor sample to a choice of either: 2 odors (the target sample and another, Experiment 1) and 6 odors (the target sample and 5 others, Experiment 2). Participants were correctly able to identify the target odor when given either 2 or 6 matches. Subjects made fewest errors when matching the odor of the silverback, whereas matching the odors of the young gorillas produced most errors. The results indicate that gorillas do produce individually identifiable body odors and introduce the possibility that odor cues may play a role in gorilla social behavior.

Olfaction is often referred to as a multidimensional sense. It is multidimensional in that approximately 1000 different receptor types, each tuned to particular odor aspects, together contribute to the olfactory percept. In humans, however, this percept is nearly unidimensional. Humans can detect and discriminate countless odorants, but can identify few by name. The one thing humans can and do invariably say about an odor is whether it is pleasant or not. We argue that this hedonic determination is the key function of olfaction. Thus, the boundaries of an odor object are determined by its pleasantness, which--unlike something material and more like an emotion--remains poorly delineated with words.

Human sweat contains a mixture of odorants with trigeminal as well as olfactory properties. It has been shown that trigeminal perception is necessary to localize odors and that humans are not able to localize substances that only activate the olfactory system. To analyze the chemosensory properties of human sweat, we studied humans' ability to localize sweat stimuli to the different nostrils. Human sweat was collected during a bicycle workout (20 males) and was then applied to 34 different subjects (17 females) during odor detection and localization experiments by using an olfactometer. During the detection experiment, subjects were instructed to discriminate between sweat stimuli (20) and blanks (10). During the localization experiment, they were assigned to allocate the stimuli to either the right (15) or the left nostril (15). We found that subjects were able to detect the sweat stimuli with moderate to high sensitivity. However, they failed to localize the sweat stimuli to the accurate nostril above chance level. Due to this inability to localize the stimuli, we conclude that human sweat does not activate the intranasal trigeminal system but only the olfactory system.

Although referred to in passing in several places, there have been few attempts to specify the functions of the human olfactory system. This article presents an initial effort at identifying and categorizing these functions, using 3 sources of information as a guide: 1) losses experienced by anosmic participants; 2) olfactory function in other mammals; and 3) capacity, namely, whether the human olfactory system can support the suggested function and whether there is evidence that it does. Three major classes of function were identified, relating to Ingestion (Detection/identification prior to ingestion; Detection of expectancy violations; Appetite regulation; Breast orientation and feeding), Avoiding environmental hazards (Fear related; Disgust related), and Social communication (Reproductive [inbreeding avoidance, fitness detection in prospective mates]; Emotional contagion [fear contagion, stress buffering]). These suggested functions were then examined with respect to 1) issues of ecological validity in human olfactory research; 2) their impact on olfactory loss; and 3) their general and specific implications for the study of human olfaction.

Darwin was struck by the many similarities between humans and other primates and believed that these similarities were the product of common ancestry. He would be even more impressed by the similarities if he had known what we have learned about primates over the last 50 years. Genetic kinship has emerged as the primary organizing force in the evolution of primate social organization and the patterning of social behaviour in non-human primate groups. There are pronounced nepotistic biases across the primate order, from tiny grey mouse lemurs (Microcebus murinus) that forage alone at night but cluster with relatives to sleep during the day, to cooperatively breeding marmosets that rely on closely related helpers to rear their young, rhesus macaque (Macaca mulatta) females who acquire their mother's rank and form strict matrilineal dominance hierarchies, male howler monkeys that help their sons maintain access to groups of females and male chimpanzees (Pan troglodytes) that form lasting relationships with their brothers. As more evidence of nepotism has accumulated, important questions about the evolutionary processes underlying these kin biases have been raised. Although kin selection predicts that altruism will be biased in favour of relatives, it is difficult to assess whether primates actually conform to predictions derived from Hamilton's rule: br > c. In addition, other mechanisms, including contingent reciprocity and mutualism, could contribute to the nepotistic biases observed in non-human primate groups. There are good reasons to suspect that these processes may complement the effects of kin selection and amplify the extent of nepotistic biases in behaviour.

In this paper, we investigated the sexual activity levels of several subtypes of middle adolescents (age 14-15 years). The subtype profiles were based on dominance-popularity status and a range of behaviors associated with dominance and popularity. In addition, gender differences in behavioral profiles were examined among dominant-popular, sexually active young adolescents. Results showed that socially dominant and popular young adolescent boys who exhibited a highly aggressive profile were more sexually active than their low-status and non-aggressive male peers; dominant-popular girls who were very attractive and gossips were more sexually active than their female peers. The results are discussed from an evolutionary psychological framework.

In this study, we examined the patterns of sex differences in men and women married to each other in five cultures (China, Russia, Turkey, UK, and the U.S.) to look for universal patterns in behavioral dimorphisms and for cultural variability in those patterns. Over 400 couples in each cultural group completed the 235-item Marriage and Relationship Questionnaire on various aspects of marriage, appropriately translated for each culture. Sex differences were anticipated in responses related to female choosiness, labor performed, emotional expressiveness, interest in sex, physical attractiveness, and jealousy. To measure male-female differences in each culture, t-tests were utilized, and effect sizes were calculated. Significant sex differences (p < .05, two-tailed) emerged in all six areas examined, although cultural differences were also seen in the patterns. For example, on items relevant to female choosiness, women in most, but not all, cultures were more likely than their husbands to endorse these statements:"I have thought of divorcing my spouse" and "My parents played a role in choosing my spouse." In China, where scores on emotional expressiveness were low, sex differences disappeared in the category related to emotions. Results suggest that long-term marriage exhibits a balance between homogamy and dimorphism serving reproductive interests. Moreover, culture may moderate this balance for particular sex differences.

An evolutionary model of human behavior should privilege emotions: essential, phylogenetically ancient behaviors that learning and decision making only subserve. Infants and non-mammals lack advanced cognitive powers but still survive. Decision making is only a means to emotional ends, which organize and prioritize behavior. The emotion of pride/shame, or dominance striving, bridges the social and biological sciences via internalization of cultural norms.

Animal and human studies have shown that individuals choose mates partly on the basis of similarity, a tendency referred to as homogamy. Several authors have suggested that a specific innate recognition mechanism, phenotypic matching, allows the organism to detect similar others by their resemblance to itself. However, several objections have been raised to this theory on both empirical and theoretical grounds. Here, we report that homogamy in humans is attained partly by sexual imprinting on the opposite-sex parent during childhood. We hypothesized that children fashion a mental model of their opposite-sex parent's phenotype that is used as a template for acquiring mates. To disentangle the effects of phenotypic matching and sexual imprinting, adopted daughters and their rearing families were examined. Judges found significant resemblance on facial traits between daughter's husband and her adoptive father. Furthermore, this effect may be modified by the quality of the father-daughter relationship during childhood. Daughters who received more emotional support from their adoptive father were more likely to choose mates similar to the father than those whose father provided a less positive emotional atmosphere.

This article describes current evolutionary research on adolescent sexual and romantic behavior. It first reviews functional explanations for basic sex differences in behavior. As in other pair-bonding mammals, women seek dominant males, and men seek and guard young, fertile females. Recent work is then described on adolescent competitiveness, mate selection, and pair bonding. The outcomes of even childhood social competition can be profound, with healthy, early-maturing, attractive children deriving lifelong benefits. Adolescent competition is intense among girls as well as boys. Depression is more common for boys with few sex partners, and for girls with many. Based on cross-cultural data and on analysis of pubertal changes in girls and boys, it can be concluded that adolescents have an evolved propensity for early sexual experimentation, followed by more judicious mate choice. Yet the bond with a girl's first sex partner is often profound. Amorous infatuations are intense in both sexes and usually mitigate within 3 years, for plausible adaptive reasons. Early menarche and unmarried motherhood impose developmental disadvantages on children but may be an evolved adaptation for stressful family conditions and a shortage of marriageable men. Gaining an understanding of normal adolescence can help in diagnosing, preventing, and treating problematic behaviors.

Marriage is universal, and pair bonding is found in other species too with highly dependent young. So marriage functions as a reproductive social arrangement that traditionally involved the extended family. The sexes are not identical in their biological contributions to children's survival, so they seek somewhat different attributes in a mate. Men seek a young, attractive, sexually faithful bride. Women seek a man who is older, taller, and (as in many other species) socially dominant. Both sexes prefer a kind, healthy, attractive, similar mate who is emotionally attached to them. A spouse who fails to maintain sufficiently high mate value is vulnerable to divorce. Infertility and sexual dissatisfaction predict divorce, as does death of a child, but the more children, the stabler the marriage. Cross-cultural data suggest that cruel or subdominant men (e.g., poor providers) and unfaithful women are prone to divorce. Marriages in which the wife dominates the husband in economic contributions, nonverbal behavior, and decision making tend to be less satisfying. In societies in which wives are economically independent of husbands, divorce rates are high. As women's economic power has risen with industrialization, divorce rates have climbed. Economic and fitness considerations also help explain cultural differences in polygyny, age at marriage, arranged marriage, concern with the bride's sexual chastity, and marriage ceremonies. Other factors also affect marital dynamics, such as state subsidies to families, the sex ratio, and influence of the couple's parents.

In social insects, workers trade personal reproduction for indirect fitness returns from helping their mother rear collateral kin. Colony membership is generally used as a proxy for kin discrimination, but the question remains whether recognition allows workers to discriminate between kin and nonkin regardless of colony affiliation. We investigated whether workers of the ant Formica fusca can identify their mother when fostered with their mother, their sisters, a hetero-colonial queen or hetero-colonial workers. We found that workers always displayed less aggression towards both their mother and their foster queen, as compared to an unfamiliar hetero-colonial queen. In support of this finding, workers maintain their colony hydrocarbon profile regardless of foster regime, yet show modifications when exposed to different environments. This indicates that recognition entails environmental and genetic components, which allow both discrimination of kin in the absence of prior contact and learning of recognition cues based on group membership.

ABSTRACT: BACKGROUND: Like other vertebrates, primates recognize their relatives, primarily to minimize inbreeding. Although associative, social learning is typically credited for discrimination of familiar kin, discrimination of unfamiliar kin remains unexplained. As sex-biased dispersal in long-lived species cannot consistently prevent encounters between unfamiliar kin, inbreeding remains a threat and mechanisms to avoid it beg explanation. Using a molecular approach that combined analyses of biochemical and microsatellite markers in 17 female and 19 male ring-tailed lemurs (Lemur catta), we describe odor-gene covariance to establish the feasibility of olfactory-mediated kin recognition. RESULTS: Despite derivation from different genital glands, labial and scrotal secretions shared about 170 of their respective 338 or 203 semiochemicals. In addition, these semiochemicals encoded information about genetic relatedness within and between the sexes. Although the sexes showed opposite seasonal patterns in signal complexity, the odor profiles of related individuals converged in the competitive breeding season, particularly in mixed-sex pairs. Thus, a mutual olfactory signal of genetic relatedness appeared specifically when such information would be crucial for preventing inbreeding. CONCLUSIONS: We suggest that signal convergence between the sexes may reflect strong selective pressures on kin recognition, whereas signal convergence within the sexes may arise as its by-product or function independently to prevent competition between unfamiliar relatives. The link between an individual's genome and its olfactory signals could be mediated by biosynthetic pathways producing polymorphic semiochemicals or by carrier proteins modifying the individual bouquet of olfactory cues. In conclusion, we unveil a possible olfactory mechanism of kin recognition that has specific relevance to understanding inbreeding avoidance and nepotistic behavior observed in free-ranging primates, and broader relevance to understanding the mechanisms of vertebrate olfactory communication.

Only a handful of cases of chromosomally integrated human herpesvirus 6 (CI-HHV-6) have been reported, suggesting that this phenomenon is rare. We here present a familial case of HHV-6 variant A (HHV-6A) transmission through a generation, which was identified in the setting of allogeneic hematopoietic stem cell transplantation (HSCT). A 31-year-old man with myelodysplastic syndrome underwent allogeneic HSCT from a human leukocyte antigen-identical sibling, and was found to be continuously yielding high copy numbers of HHV-6A DNA in plasma evaluated by real-time polymerase chain reaction (PCR). Antiviral therapy with ganciclovir or foscarnet failed to decrease the copy numbers. HHV-6A DNA was detected in the patient's buccal mucosa and hair follicles, and was also detected in the plasma, whole blood, and buccal mucosa of the patient's father and 2 siblings, but not in his mother. The sequences of HHV-6A DNA isolated from all family members were identical. Since monitoring of HHV-6 by PCR has been widely introduced to the field of HSCT, transplant physicians should be aware of such an alternative form of HHV-6 transmission, particularly when HHV-6A is detected.

Kin recognition, an evolutionary phenomenon ubiquitous among phyla, is thought to promote an individual's genes by facilitating nepotism and avoidance of inbreeding. Whereas isolating and studying kin recognition mechanisms in humans using auditory and visual stimuli is problematic because of the high degree of conscious recognition of the individual involved, kin recognition based on body odors is done predominantly without conscious recognition. Using this, we mapped the neural substrates of human kin recognition by acquiring measures of regional cerebral blood flow from women smelling the body odors of either their sister or their same-sex friend. The initial behavioral experiment demonstrated that accurate identification of kin is performed with a low conscious recognition. The subsequent neuroimaging experiment demonstrated that olfactory based kin recognition in women recruited the frontal-temporal junction, the insula, and the dorsomedial prefrontal cortex; the latter area is implicated in the coding of self-referent processing and kin recognition. We further show that the neuronal response is seemingly independent of conscious identification of the individual source, demonstrating that humans have an odor based kin detection system akin to what has been shown for other mammals. Hum Brain Mapp 2009.(c) 2008 Wiley-Liss, Inc.

Maloney and Dal Martello [Maloney, L.T., Dal Martello, M.F.(2006). Kin recognition and the perceived facial similarity of children. Journal of Vision, 6(10), 1047-1056. http://www.journalofvision.org/6/10/4/] reported that similarity ratings of pairs of related and unrelated children were almost perfect predictors of the probability that those children were judged as being siblings by a second group of observers. Surprisingly, similarity ratings were poor predictors of whether a pair was same-sex or opposite-sex, suggesting that people ignore cues that are uninformative about kinship when making similarity judgments of faces. Using adult sibling faces, we find that similarity ratings for same-sex pairs were significantly higher than for opposite-sex pairs, suggesting that similarity judgments of adult faces are not entirely synonymous with kinship judgments.

We investigated the ability of rabbit pups to display preferences towards various elements of their postnatal environment during the stage of confinement in the nest. Subjects were submitted to a two-choice test during the first week after birth to assess if they could detect and discriminate between does, litters of pups, or nesting materials of the same developmental stage. On D1 and D7, pups were attracted to any lactating doe, litter, or nest when compared to an empty compartment. When two stimuli were opposed, pups preferred their own nest to an alien one on D1 and D7 but not their mother nor their siblings when opposed to alien does or pups. However, additional tests indicated that this lack of preference for kin conspecifics resulted from a predominant attraction to thermal cues over individual odors. Indeed, pups were strongly attracted to a warm compartment (37 degrees C) than to a cooler one (20 degrees C) and once thermal cues were controlled for in the testing situation, the pups were specifically attracted to odors of their own mother's hair and of their siblings. No preference was observed towards the mother's uterine secretions. In conclusion, pups can recognize olfactory cues emanating from their mother and their siblings the day after birth. The preference for nesting materials would reflect in major part the combined attraction to maternal and sibling odors present in the nest.(c) 2008 Wiley Periodicals, Inc. Dev Psychobiol 50: 542-553, 2008.

Procellariiform seabirds wander the world's oceans aided by olfactory abilities rivaling those of any animal on earth. Over the past 15 years, I have been privileged to study the sensory ecology of procellariiforms, focusing on how olfaction contributes to behaviors, ranging from foraging and navigation to individual odor recognition, in a broader sensory context. We have developed a number of field techniques for measuring both olfactory- and visually based behaviors in chicks and adults of various species. Our choice of test odors has been informed by long-term dietary studies and geochemical data on the production and distribution of identifiable, scented compounds found in productive waters. This multidisciplinary approach has shown us that odors provide different information over the ocean depending on the spatial scale. At large spatial scales (thousands of square kilometers), an olfactory landscape superimposed upon the ocean surface reflects oceanographic or bathymetric features where phytoplankton accumulate and an area-restricted search for prey is likely to be successful. At small spatial scales (tens to hundreds of square kilometers), birds use odors and visual cues to pinpoint and capture prey directly. We have further identified species-specific, sensory-based foraging strategies, which we have begun to explore in evolutionary and developmental contexts. With respect to chemical communication among individuals, we have shown that some species can distinguish familiar individuals by scent cues alone. We are now set to explore the mechanistic basis for these discriminatory abilities in the context of kin recognition, and whether or not the major histocompatibility complex is involved.

Kin recognition is important in animal social systems. However, though plants often compete with kin, there has been as yet no direct evidence that plants recognize kin in competitive interactions. Here we show in the annual plant Cakile edentula, allocation to roots increased when groups of strangers shared a common pot, but not when groups of siblings shared a pot. Our results demonstrate that plants can discriminate kin in competitive interactions and indicate that the root interactions may provide the cue for kin recognition. Because greater root allocation is argued to increase below-ground competitive ability, the results are consistent with kin selection.

Drawing on the data from the longitudinal Binuclear Family Study, 173 grown children were interviewed 20 years after their parents' divorce. This article addresses two basic questions:(1) What impact does the relationship between parents have on their children 20 years after the divorce? and (2) When a parent remarries or cohabits, how does it impact a child's sense of family? The findings show that the parental subsystem continues to impact the binuclear family 20 years after marital disruption by exerting a strong influence on the quality of relationships within the family system. Children who reported that their parents were cooperative also reported better relationships with their parents, grandparents, stepparents, and siblings. Over the course of 20 years, most of the children experienced the remarriage of one or both parents, and one third of this sample remembered the remarriage as more stressful than the divorce. Of those who experienced the remarriage of both of their parents, two thirds reported that their father's remarriage was more stressful than their mother's. When children's relationships with their fathers deteriorated after divorce, their relationships with their paternal grandparents, stepmother, and stepsiblings were distant, negative, or nonexistent. Whether family relationships remain stable, improve, or get worse is dependent on a complex interweaving of many factors. Considering the long-term implications of divorce, the need to emphasize life course and family system perspectives is underscored.

Recent research on kin investment shows a matrilateral bias as a function of paternity uncertainty. Kin investment, however, is a special case of kin altruism. We thus hypothesize that psychological adaptations have evolved to regulate cousin-directed altruism according to predictably variable levels of paternity uncertainty in different categories of cousins. We develop a formal mathematical model that predicts that individuals should be most willing to act altruistically towards their mother's sister's (MoSis) children and least willing to act altruistically towards their father's brother's (FaBro) children. Altruism towards father's sister's (FaSis) and mother's brother's (MoBro) children are predicted to fall in between. An empirical study (N=195), assessing expressed altruistic proclivities, confirmed the predictions from the model. Participants expressed willingness-to-help following the descending order:(i) MoSis children,(ii) MoBro children,(iii) FaSis children, and (iv) FaBro children. The psychological variables of emotional closeness, empathic concern and contact frequency showed precisely the same pattern across distinct cousins, providing convergent confirmation of the model. The results support the hypothesis of cousin-specific adaptations sensitive to varying probabilities of paternity uncertainty.