Kin recognition in zebrafish: a 24-hour window for olfactory imprinting

Distinguishing kin from non-kin profoundly impacts the evolution of social behaviour. Individuals able to assess the genetic relatedness of conspecifics can preferentially allocate resources towards related individuals and avoid inbreeding. We have addressed the question of how animals acquire the ability to recognize kin by studying the development of olfactory kin preference in zebrafish (Danio rerio). Previously, we showed that zebrafish use an olfactory template to recognize even unfamiliar kin through phenotype matching. Here, we show for the first time that this phenotype matching is based on a learned olfactory imprinting process in which exposure to kin individuals on day 6 post fertilization (pf) is necessary and sufficient for imprinting. Larvae that were exposed to kin before or after but not on day 6 pf did not recognize kin. Larvae isolated from all contact with conspecifics did not imprint on their own chemical cues; therefore, we see no evidence for kin recognition through self-matching in this species. Surprisingly, exposure to non-kin odour during the sensitive phase of development did not result in imprinting on the odour cues of unrelated individuals, suggesting a genetic predisposition to kin odour. Urine-born peptides expressed by genes of the immune system (MHC) are important messengers carrying information about 'self' and 'other'. We suggest that phenotype matching is acquired through a time-sensitive learning process that, in zebrafish, includes a genetic predisposition potentially involving MHC genes expressed in the olfactory receptor neurons.     

Kin recognition in golden hamsters: evidence for kinship odours

Differential treatment of kin and non-kin has been well documented, but much remains unclear about how kin are recognized. If kin are recognized by a phenotype-matching mechanism, there must be a correlation between genetic relatedness and the similarity of cues used for recognition. A habituation technique was used with golden hamsters,Mesocricetus auratus, to investigate the relative similarity of the odour quality of flank gland secretions from siblings and unrelated individuals. Hamsters discriminated between the odours of their own, same-sex siblings but also treated these odours as similar compared to odours of non-siblings (experiment 1). They did not discriminate between the flank gland odours of unfamiliar siblings from another family (experiment 2). They also did not discriminate between the flank gland odours of unfamiliar, paternal half-siblings from another family (experiment 3). These results indicate that subjects perceived odours from genetically similar individuals as similar and provide evidence for kinship odour cues. The discrimination between the flank gland odours of subjects’ own siblings, however, indicates that hamsters learn the subtle differences between the odours of their close kin, probably through experience with siblings in the nest. When only volatile components from flank gland secretions were available to subjects (experiment 4), they again discriminated between the odours of their own siblings, suggesting that the volatile components from the flank gland secretion were sufficient for recognition of individual litter-mates.     

Genetic relatedness assessment through individual odour similarities in mice

Previous research has demonstrated that closer genetic relatedness between individuals (from kin to across species) is associated with greater similarity in the qualities of their individual odours ('odour-genes covariance'). This predictable relationship between individual genotypes and individual odours could enable animals to assess their degree of genetic relatedness to other individuals by comparing the degree of similarity between another individual's odour and their own odour. In two-choice tests with odours of unfamiliar mice from different populations and species, subjects from two species of wild mice, Mus spicilegus and M. musculus , that had been raised in mixed litters of both species spent significantly more time investigating the ano-genital odour of the more genetically similar individual. This differential interest was not affected by common rearing with heterospecifics. These responses are consistent with a self-referencing mechanism enabling differential responses across a wide spectrum of genetic relatedness from kin through populations to heterospecifics. These assessments depend on the degree of similarity between the donor's and the subject's odours rather than on differences between them. Such a parsimonious mechanism may provide a basis for differential responses to conspecifics as opposed to heterospecifics that may function as a premating isolating mechanism.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 78, 595–603.     

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.     

Retention of social recognition after hibernation in Belding's ground squirrels

The retention of social memory during long periods of separation, such as hibernation or migration, has not been well documented, despite evidence for long-term social relationships in migrating species or in long-lived sedentary species. We investigated the ability of captive Belding's ground squirrels, Spermophilus beldingi, to remember previously familiar individuals as well as littermates after 9 months of isolation. Before hibernation, young ground squirrels discriminated between odours of familiar and unfamiliar individuals, as shown by greater investigation of a novel individual's odour. The following spring, these yearlings did not respond differentially to odours of previously familiar and unfamiliar individuals, suggesting that memory for familiar conspecifics was lost during hibernation. In contrast, both female and male yearlings continued to discriminate between odours of littermates and previously familiar nonlittermates. Thus, recognition of close kin was maintained during prolonged social isolation, but recognition of familiar, unrelated individuals was not. If re-establishment of familiarity is not costly or if adults rarely interact with the same individuals in successive years, then selection may not favour retention of individual memories of particular conspecifics over the winter. Even though males rarely encounter kin after dispersal, yearling males did recognize their siblings, suggesting that the relative costs of maintaining kin-recognition abilities year-round may be low. Possible mechanisms underlying the formation and maintenance of individual and kin recognition are discussed. Copyright 2000 The Association for the Study of Animal Behaviour.     

“KIN RECOGNITION” AMONG SPADEFOOT TOAD TADPOLES: A SIDE-EFFECT OF HABITAT SELECTION?

Many animals modify their behavior toward unfamiliar conspecifics as a function of their genetic relatedness. A fundamental problem of any kin recognition study is determining what is being recognized and why. For anuran tadpoles, the predominant view is that associating with relatives is kin-selected because these relatives may thereby accrue benefits through increased growth or predation avoidance. An alternative view is that kin associations are simply a side-effect of habitat selection and thus do not represent attempts to identify kin per se. In the laboratory, spadefoot toad tadpoles (Scaphiopus multiplicatus) preferentially associated with unfamiliar siblings over unfamiliar nonsiblings, as do other anurans. However, same age tadpoles also were more likely to orient toward unfamiliar nonsiblings reared on the same food (familiar food) than toward unfamiliar siblings that were reared on unfamiliar food. These results, together with the results of previous tadpole kin recognition studies, suggest that tadpoles orient toward cues learned early in ontogeny, regardless of the cues' source. Tadpoles that preferentially associated with cues learned from their environment at birth would tend to be philopatric. Censuses of 14 natural ponds revealed that tadpole density remained greatest near oviposition sites until four days before metamorphosis. Tadpole philopatry may be advantageous: tadpoles restricted to their natal site had greater growth and survivorship than did their siblings restricted to randomly selected sites elsewhere within the same pond. Thus kin affiliative tendency observed in the laboratory in this and perhaps other species of anurans may be a byproduct of habitat selection. Since kin discrimination in animals is most commonly assayed as orientation toward kin, it follows that many examples of “kin recognition” may not represent true attempts to identify kin as such, but rather may reflect some other recognition system that is under entirely different selective pressures.     

Behavioural development of conspecific odour preferences in bank voles,Clethrionomys glareolus

Biological odours of conspecifics are known to have strong influences on behavioural interaction in bank voles Clethrionomys glareolus. This experiment tested two hypotheses. (1) Olfactory cues from familiar and unfamiliar mature opposite-sex conspecifics differ in their attractiveness to males and females, and their behavioural reactions change with age. (2) A genetically based mechanism is involved in female recognition of kin.In a two-choice preference test, prepubertal males and females were more attracted to familiar than to unfamiliar odours of opposite-sex conspecifics, as manifested by more time spent sniffing familiar voles. As the young reached sexual maturity they shifted their odour preferences. Mature males and females preferred the novel odour of unrelated opposite-sex conspecifics to that of relatives. The results of experiments testing the second hypothesis indicate that females use a genetically based mechanism to recognise their kin. Young and mature females were able to recognise the odour of their biological but socially unknown fathers, and showed the same pattern of behaviour as females in previous experiments.The possible biological functions of kin recognition in bank voles are discussed.     

KIN RECOGNITION: FUNCTIONS AND MECHANISMS A REVIEW

1. The aim of this paper has been to review the theory behind kin recognition to examine the benefits individuals obtain by recognizing their kin and to review the mechanisms used by individuals in their recognition of kin. 2. The ability to discriminate between kin and non-kin, and between different classes of kin gives individuals advantages in fitness greater than individuals unable to recognize their kin. Four specific areas of benefit were considered: altruistic behaviour, co-operative behaviour, parental care and mate choice. Finally the possibility that kin recognition has arisen as a byproduct from some other ability was discussed. 3. Mechanisms of kin recognition were considered with respect to three essential components of kin recognition. The cue used to discriminate kin, how individuals classify conspecifics as kin, etc. and how the ability to recognize kin develops. 4. Individuals can use a number of cues to discriminate kin from non-kin. These were divided into cues presented by conspecifics (conspecific cues), of which three types were considered: individual, genetic and group/colony cues, and non-conspecific cues, environmental, state and no cues. Kin recognition could be achieved by use of all these cues. 5. How individuals classify their conspecifics as kin, etc. can be achieved in a number of ways; dishabituation or self-matching, which require no learning of kinship cues, or by phenotype matching or familiarity, both of which require the learning of kinship information. 6. It may be necessary for individuals to acquire information concerning kinship. This may be learned, and can be achieved in a number of ways; physiological imprinting, exposure learning or associative learning. Acquisition by these means is non-selective, in that the cues which are most salient in the individual's environment will be learned. Selectivity can be introduced into this process to increase the probability of acquiring kinship information by a number of means; learning from parents, sensitive periods for learning and prenatal learning. Finally, kinship information could be supplied by recognition genes. 7. A distinction is drawn between cues which are used by an individual in the discrimination of kin, discriminators, and cues which are used by individuals in the acquisition of information about kinship, acquisitors. 8. Experiments used to support previous categories of mechanisms of kin recognition were examined in the light of this discussion and it was found that the results were open to a number of different interpretations and yielded little specific information about the mechanisms of kin recognition. 9. It was concluded that there was much evidence, both theoretical and experimental to support the proposed benefits individuals gain from recognizing kin, but much more research is required before the mechanisms of kin recognition are fully understood.     

Individual odour similarities across species parallel phylogenetic relationships in the S. ehrenbergi superspecies of mole-rats

Research using habituation techniques has shown that rodents from the same kin group, population, or species share similarities in their individual odours that covary with shared genetic similarities between them, that is, the closer their genetic relatedness, the more similar their odours. We assessed similarities in individual odours across four sibling species of subterranean mole-rats from the Spalax ehrenbergi superspecies in Israel. Mole-rats were habituated to the urine odour of a same-sex individual from one species then tested with urine odours of individuals from two different species of the superspecies. Subjects treated urine odours of individuals from more closely genetically related species as similar compared with the odours of individuals from a less closely related species, showing that the covariance between odours and genes extends across species. These similarities in odour also paralleled genetic similarities determined by molecular analysis: odours of descendent species were perceived as similar to those of their closest ancestral species, suggesting that some qualities of the odour of the ancestral species persist in the descendent species. It is generally assumed that during speciation incipient species develop species-specific markers, including, for example, odour markers, to facilitate discrimination of conspecifics from close ancestral heterospecifics. Our findings indicate that similarities in odours across species are more salient than species-specific odour markers. Such findings may also have important implications for mechanisms of species recognition. Copyright 2000 The Association for the Study of Animal Behaviour.     

Social Interaction is Necessary for Discrimination between and Memory for Odours of Close Relatives in Golden Hamsters

The role of social experience in recognition of individuals and their odours is not well understood. In a previous study, hamsters discriminated between the odours of their familiar litter mates, but they did not discriminate between the odours of two males or two females from a different litter that were unfamiliar to them. In this paper the role of social experience in discrimination between odours of male litter mates is examined using habituation techniques. Males, tested 1.5–2 mo after separation from their litter mates, discriminated between the flank gland odours of their foster brothers, but they did not discriminate between flank odours from their unfamiliar brothers that had been reared by another mother (Expt. 1). In Expt. 2, adult males did not discriminate between the flank gland odours of two males from a different litter after a week of exposure across a barrier to their sights, sounds, and odours, but subjects did discriminate between the odours of these same males after five brief encounters with them. A month after the encounters, however, subjects no longer discriminated between these odours. In contrast, adult males discriminated between the flank odours of their brothers 9 mo after separation from them (Expt. 3). These results suggest that hamsters must have direct interactions with closely related individuals to discriminate between their odours because the odours of close kin are so similar. Experience with nest mates results in long-lasting memories for their odours.     

Kin discrimination and altruism in the larvae of a solitary insect

Kin selection theory predicts altruism between related individuals, which requires the ability to recognize kin from non-kin. In insects, kin discrimination associated with altruistic behaviour is well-known in clonal and social species but in very few solitary insects. Here, we report that the solitary larvae of a non-social insect Aleochara bilineata Gyll. (Coleoptera; Staphylinidae) show kin discrimination and sibling-directed altruistic behaviour. Larvae superparasitize more frequently the hosts parasitized by non-kin individuals than those hosts parasitized by siblings. Kin discrimination probably occurs by self-referent phenotype matching, where an individual compares its own phenotype with that of a non-familiar related individual, a mechanism rarely demonstrated in animals. The label used to recognize kin from non-kin corresponds to substances contained in the plug placed on the hosts by the resident larvae during the parasitization process. Kin competition induced by a limited larval dispersion may have favoured the evolution of kin recognition in this solitary species.     

Nestmate recognition in sweat bees (Lasioglossum zephyrum): Does an individual recognize its own odour or only odours of its nestmates?

We investigated the olfactory mechanism by which guard bees of Lasioglossum zephyrum decide whether to admit conspecifics to their nests. First we set up colonies of young bees, consisting of sisters from a single family or a mixture of bees from two distinct families. These bees were then introduced into colonies other than their own. Our experimental evidence shows that guards learn the odours of their nestmates, then accept or reject other bees on the basis of the similarity of the latters' odours to those of the guards' nestmates. Guards act as though they do not use their own odour as a reference for nestmate recognition. This recognition mechanism enables individuals with different odours to live together; it may also enhance the operation of kin selection by providing a more complete basis for discriminating relatives from non-relatives. No evidence was found that nestmates acquire one another's odours. Such lack of odour transfer may be characteristic of early stages in the evolution of recognition mechanisms.     

Recognition of individual,sex and species odours by salamanders of the Plethodon glutinosus-P. Jordani complex

Salamanders of the Plethodon glutinosus-P. jordani complex were tested for the ability to distinguish conspecific, sex-specific, and heterospecific chemical cues. Male and female P. glutinosus preferred substrates previously occupied by conspecifics over their own, but randomly chose between substrates marked by male or female conspecifics. This suggests that while these salamanders are able to distinguish between their own and conspecific substrate odours, they are unable to identify sex by means of substrate odours. Experiments using an olfactometer showed that male P. jordani, male and female P. glutinosus, and an electrophoretically distinct and non-hybridizing sympatric phenotype in the P. glutinosus complex (here called species A), all preferred female airborne odours over male airborne odours. This demonstrates that these salamanders can identify sex by means of airborne odours. Male P. glutinosus and species A both preferred conspecific female odours over heterospecific female odours in olfactometer experiments. These results suggest an important role for olfaction in the sexual and social behaviour of these salamanders, particularly as a pre-mating isolating mechanism.     

Young female pigs can discriminate individual differences in odours from conspecific urine

The ability to discriminate odour cues from different conspecifics has been demonstrated in a variety of small mammalian species. We used a habituation-dishabituation procedure to investigate whether 10-week-old female pigs,Sus scrofa , are able to discriminate between urinary odours from similar-aged conspecifics that were unfamiliar (not encountered for at least 7 weeks). We also examined whether environmental factors can affect the ease with which urine from different individuals is discriminated. Subjects receiving urine samples from the same unfamiliar individual in two successive 2-min exposures separated by a 15-min interval showed habituation in their investigation response to the urine in the second exposure. This habituation was maintained in a third 2-min exposure, 15 min later, if the urine sample was again from the same individual. However, if the urine sample was from a different unfamiliar individual, there was a dishabituation of the investigation response. This was taken to indicate an ability to discriminate the two samples. These data indicate that young pigs could use urinary cues to discriminate other individuals. Effective individual discrimination should facilitate the formation and maintenance of stable social groupings but could be disrupted if, for example, animals from the same group share common odour cues that mask individually distinctive scents. However, urine samples from individuals living in the same group appeared to be no more difficult to discriminate than those from individuals living in different groups. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

鳄蜥对熟悉和陌生个体气味的辨别

个体辨别对于减少同种争斗以及配偶选择具有重要意义。我们用棉棒粘取鳄蜥(Shinisaurus crocodilurus)尿液作为气味源,以香水作为对照,测定鳄蜥对熟悉个体气味、陌生个体气味以及香水的舔舌次数和舔舌潜伏期,来探讨鳄蜥通过化学信息辨别熟悉和陌生个体的能力。结果显示,不论是雌性还是雄性,对不同个体尿液的舔舌次数均显著高于对香水的,舔舌潜伏期显著短于香水的;尽管雄性对陌生同性个体气味与熟悉同性个体气味的舔舌次数无显著差异,但对前者的舔舌潜伏期显著短于后者;雄性对陌生雌性气味的舔舌次数显著多于熟悉雌性气味的,对前者的舔舌潜伏期显著短于后者;雌性对陌生雄性气味的舔舌潜伏期显著短于对熟悉雄性气味的;雄鳄蜥对陌生雌性气味的舔舌次数显著多于雌鳄蜥对陌生雄性的。结果表明,鳄蜥能辨别同种个体的化学信息,并能通过化学信息来辨别熟悉和陌生个体,推测鳄蜥的这种辨别能力对其领域分配以及繁殖交配有重要作用。     

Paternal kin discrimination: the evidence and likely mechanisms

One of the most important assumptions of kin selection theory is that individuals behave differently towards kin than non-kin. In mammals, there is strong evidence that maternal kin are distinguished from non-kin via familiarity. However, little is known about whether or not mammals can also recognize paternal kin as many female mammals, including primates, mate with multiple males near the time of conception, potentially concealing paternal kinship. Genetic data in several mammalian species with a promiscuous mating system and male-biased dispersal reveal a high skew in male reproduction which leads to co-residing paternal half-siblings. In most primates, individuals also form stable bisexual groups creating opportunities for males to interact with their offspring. Here I consider close paternal kin co-resident in the same social group, such as father-offspring and paternal half-siblings (i.e. animals sharing the same father but who were born to different mothers) and review mammalian studies of paternal kin discrimination. Furthermore, I summarize the most likely mechanisms of paternal kin discrimination (familiarity and phenotype matching). When familiarity is the underlying mechanism, mothers and/or the sire could mediate familiarity among paternal half-siblings as well as between fathers and offspring assuming mothers and/or fathers can assess paternity. When animals use phenotype matching, they might use their fathers' template (when the father is present) or self (when the father is absent) to assess paternal kinship in others. Available evidence suggests that familiarity and phenotype matching might be used for paternal kin discrimination and that both mechanisms might apply to a wide range of social mammals characterized by a high skew in male reproduction and co-residence of paternal kin. Among primates, suggested evidence for phenotype matching can often have an alternative explanation, which emphasizes the crucial importance of controlling for familiarity as a potential confounding variable. However, the mechanism/s used to identify paternal kin might differ within a species (as a function of each individual's specific circumstances) as well as among species (depending upon the key sensory modalities of the species considered). Finally, I discuss the possible cues used in paternal kin discrimination and offer suggestions for future studies.     

Group relatedness and kin discrimination in honey bees Apis mellifera L.

In addressing problems concerning kin discrimination and the evolutionary significance of inclusive fitness in highly eusocial insects, the genetic relatedness between groups, such as individuals belonging to the same patriline within colonies, is of interest. Based on the definitions for individual genetic relationship, several new measurements of group relatedness are derived and their genetical meaning is discussed. In a metabolic bioassay for the documentation of kin discrimination, carbon dioxide production was used to quantify reactions of groups of honey bees to volatile odours of other groups. Worker test groups, which had not experienced their hive environment, were able to discriminate between related and unrelated groups of workers within and between colonies. Test groups responded in a similar way to odours of genotypically mixed groups with related and unrelated workers and to the odours of pure groups of the less related subgroup. Thus, the reaction to group odours that function as discrimination labeles is not a linear response as predicted by an additive gestalt odour model. Since experienced bees collected from the colony could not discriminate between the various patrilines, the observed phenomenon resembles nest mate recognition rather than within-colony kin recognition.     

Mechanisms of kin discrimination inferred from pedigrees and the spatial distribution of mates

Where animals avoid inbreeding, different mechanisms of kin discrimination can leave different 'signatures' in the patterns of observed mate relationship. For example, consider a species with no paternal care. If a female avoids mating with familiar individuals, one would expect a deficit of offspring whose parents are maternal half-siblings, but paternal half-siblings would be unfamiliar with each other and thus have offspring at the frequency expected by chance. If spatial cues are used to avoid inbreeding, a female would be expected to produce few offspring with males (even unrelated males) living near her birth site. We searched for these and other signatures with data from a long-term study of banner-tailed kangaroo rats, Dipodomys spectabilis, in Arizona, USA, using a combination of intensive censusing, mapping of available dens, microsatellite-based parentage determination, and a randomization routine that determines the numbers of offspring expected if females in the population mate indiscriminately among the males available to them. The data are consistent with the hypothesis that kangaroo rats discriminate kin by familiarity developed via association early in life, rather than by using spatial cues or self-referential phenotype matching. Our approach should be widely applicable as a means of assessing the degree to which kin discrimination exists (in contexts like nepotism as well as inbreeding avoidance) and in inferring what cues animals use to assess categories of relationship.     

Odour–genes covariance within a natural population of subterranean Spalax galili blind mole rats

Previous research, using habituation techniques with multiple rodent species as subjects, has demonstrated (from kin to across species) the greater perceptual similarity in the qualities of individual odours of more closely genetically related individuals ('odour–genes covariance', abbreviated 'OGC'). This predictable relationship between individual genotypes and individual odours has only been assessed in rodents in which the genetic similarities and dissimilarities are known either through laboratory breeding or by selecting odour donors from different populations or species. To study OGC within a natural population of rodents, we genotyped Spalax galili blind mole rats using ten microsatellite markers, and conducted pairwise genetic distance comparisons to estimate genetic similarity and to calculate the relatedness coefficient between pairs of individuals. We then used habituation-generalization techniques to assess whether perceived odour similarities covaried with the genetic similarities we had identified. Indeed, animals treated odours from genetically closer donors as more similar than odours from less genetically similar donors. The results suggest an accurate and subtle ability to resolve genetically determined odour distinctions among familially unrelated animals within a population. Graded differential responses to conspecific odours based on degrees of similarity between other individuals' odours and one's own have been demonstrated in kinship and species discriminations and preferences. The evidence presented here provides the basis for hypothesizing a similar process that could promote optimal outbreeding and inclusive fitness within populations of conspecifics. The evolutionary implications of these findings are discussed.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 483–490.     

Kin discrimination in the social lizard Egernia saxatilis (Scincidae)

The ability to discriminate kin from nonkin is critical forthe evolution of kin-based sociality. Black rock skinks, Egerniasaxatilis, are viviparous lizards that typically live in "nuclearfamilies" consisting of an adult male, adult female, and oneor more cohorts of juveniles. Laboratory trials showed thatjuvenile lizards can discriminate between the scent of adultsfrom their own social group versus that of unfamiliar adults.Experiments in which we translocated individuals among familygroups revealed that this discriminatory ability was based onfamiliarity with other individuals rather than genetic relatedness.For example, neither "fostered" juveniles nor their mothersdisplayed any scent-based kin discrimination when brought togetherafter 2 months' separation. Thus, unlike the closely related(and also social) Egernia striolata, black rock skinks basekin discrimination on familiarity rather than genotypic similarity.