Sex Bias In Dispersal Research Articles

(Not) far from home: No sex bias in dispersal, but limited genetic patch size, in an endangered species, the Spotted Turtle (Clemmys guttata).

Sex-biased dispersal is common in many animals, with male-biased dispersal often found in studies of mammals and reptiles, including interpretations of spatial genetic structure, ostensibly as a result of male-male competition and a lack of male parental care. Few studies of sex-biased dispersal have been conducted in turtles, but a handful of studies, in saltwater turtles and in terrestrial turtles, have detected male-biased dispersal as expected. We tested for sex-biased dispersal in the endangered freshwater turtle, the spotted turtle (Clemmys guttata) by investigating fine-scale genetic spatial structure of males and females. We found significant spatial genetic structure in both sexes, but the patterns mimicked each other. Both males and females typically had higher than expected relatedness at distances <25 km, and in many distance classes greater than 25 km, less than expected relatedness. Similar patterns were apparent whether we used only loci in Hardy-Weinberg equilibrium (n=7) or also included loci with potential null alleles (n=5). We conclude that, contrary to expectations, sex-biased dispersal is not occurring in this species, possibly related to the reverse sexual dimorphism in this species, with females having brighter colors. We did, however, detect significant spatial genetic structure in males and females, separate and combined, showing philopatry within a genetic patch size of <25 km in C.guttata, which is concerning for an endangered species whose populations are often separated by distances greater than the genetic patch size.

Investigating sex differences in genetic relatedness in great-tailed grackles in Tempe, Arizona to infer potential sex biases in dispersal

In most bird species, females disperse prior to their first breeding attempt, while males remain close to the place they hatched. While explanations for such female bias in natal dispersal have focused on the resource-defense based monogamous mating system that is prevalent in most birds, the factors shaping dispersal decisions are often more complex. Studying species with different social and mating systems can help illuminate the various factors shaping sex biased dispersal. Here, we use genetic approaches to determine whether females and/or males disperse in great-tailed grackles (Quiscalus mexicanus), which have a mating system where the males hold breeding territories that multiple females might choose to place their nest in, but females forage independently of these breeding territories across a wider area. We find that for individuals caught at a single site in Arizona, (i) the average relatedness among all female dyads, but not of males, is higher than average relatedness among other individuals at the site; (ii) that female close relatives are found within shorter distances from each other than pairs of unrelated females, but male close relatives at larger distances than pairs of unrelated males; and (iii) we find a decline in relatedness with increasing spatial distances for females, but not for males. Our findings show that great-tailed grackles offer opportunities to understand how divergent social and mating systems might shape natal philopatry and dispersal, given their reversal of the usual sex-bias in dispersal with females associating with genetic relatives while males are not.

Using strontium isotopes to determine philopatry and dispersal in primates: a case study from Kibale National Park.

Strontium isotope ratios (87Sr/86Sr) allow researchers to track changes in mobility throughout an animal's life and could theoretically be used to reconstruct sex-biases in philopatry and dispersal patterns in primates. Dispersal patterns are a life-history variable that correlate with numerous aspects of behaviour and socio-ecology that are elusive in the fossil record. The present study demonstrates that the standard archaeological method used to differentiate between ‘local’ and ‘non-local’ individuals, which involves comparing faunal isotopic ratios with environmental isotopic minima and maxima, is not always reliable; aspects of primate behaviour, local environments, geologic heterogeneity and the availability of detailed geologic maps may compromise its utility in certain situations. This study instead introduces a different methodological approach: calculating offset values to compare 87Sr/86Sr of teeth with that of bone or local environments. We demonstrate this method's effectiveness using data from five species of primates, including chimpanzees, from Kibale National Park, Uganda. Tooth-to-bone offsets reliably indicate sex-biases in dispersal for primates with small home ranges while tooth-to-environment offset comparisons are more reliable for primates with larger home ranges. Overall, tooth-to-environment offsets yield the most reliable predictions of species' sex-biases in dispersal.

Mate-finding dispersal reduces local mate limitation and sex bias in dispersal.

Sex-biased dispersal (SBD) often skews the local sex ratio in a population. This can result in a shortage of mates for individuals of the less-dispersive sex. Such mate limitation can lead to Allee effects in populations that are small or undergoing range expansion, consequently affecting their survival, growth, stability and invasion speed. Theory predicts that mate shortage can lead to either an increase or a decrease in the dispersal of the less-dispersive sex. However, neither of these predictions have been empirically validated. To investigate how SBD-induced mate limitation affects dispersal of the less-dispersive sex, we used Drosophila melanogaster populations with varying dispersal propensities. To rule out any mate-independent density effects, we examined the behavioural plasticity of dispersal in the presence of mates as well as same-sex individuals with differential dispersal capabilities. In the presence of high-dispersive mates, the dispersal of both male and female individuals was significantly increased. However, the magnitude of this increase was much larger in males than in females, indicating that the former shows greater mate-finding dispersal. Moreover, the dispersal of either sex did not change when dispersing alongside high- or low-dispersive individuals of the same sex. This suggested that the observed plasticity in dispersal was indeed due to mate-finding dispersal, and not mate-independent density effects. Strong mate-finding dispersal can diminish the magnitude of sex bias in dispersal. This can modulate the evolutionary processes that shape range expansions and invasions, depending on the population size. In small populations, mate-finding dispersal can ameliorate Allee effects. However, in large populations, it can dilute the effects of spatial sorting.

Weak population structure in the ant Formica fusca.

Dispersal is a fundamental trait of a species’ biology. High dispersal results in weakly structured or even panmictic populations over large areas, whereas weak dispersal enables population differentiation and strong spatial structuring. We report on the genetic population structure in the polygyne ant Formica fusca and the relative contribution of the dispersing males and females to this. We sampled 12 localities across a ∼35 km2 study area in Finland and generated mitochondrial DNA (mtDNA) haplotype data and microsatellite data. First, we assessed queen dispersal by estimating population differentiation from mtDNA haplotype data. Second, we analysed nuclear DNA microsatellite data to determine overall population genetic substructure in the study area with principal components analysis, Bayesian clustering, hierarchical F statistics and testing for evidence of isolation-by-distance. Third, we directly compared genetic differentiation estimates from maternally inherited mtDNA and bi-parentally inherited DNA microsatellites to test for sex-bias in dispersal. Our results showed no significant spatial structure or isolation by distance in neither mtDNA nor DNA microsatellite data, suggesting high dispersal of both sexes across the study area. However, mitochondrial differentiation was weaker (Fst-mt = 0.0047) than nuclear differentiation (Fst-nuc = 0.027), which translates into a sixfold larger female migration rate compared to that of males. We conclude that the weak population substructure reflects high dispersal in both sexes, and it is consistent with F. fusca as a pioneer species exploiting unstable habitats in successional boreal forests.

Socio-genetic correlates of unbiased sex dispersal in a population of black capuchin monkeys (Sapajus nigritus)

Most social mammal species exhibit male-biased dispersal. Sex bias in dispersal leads to a higher degree of relatedness among individuals of the philopatric sex, thus an atypical dispersal pattern might lead to deviations in the typical within-group kinship structure. Kinship, in turn, influences patterns of social interactions, as widely evident by kin-biased behaviors. We investigated the link between dispersal, relatedness structure, and sociopositive interactions established by adult females of black capuchin monkeys (Sapajus nigritus) living in a population that experiences female dispersal, an unusual pattern for capuchin monkeys. The study was conducted in Parque Estadual Carlos Botelho (PECB), within the Brazilian Atlantic Forest. We addressed dispersal and relatedness patterns by genotyping 20 adults of 3 groups across 9 microsatellite loci. We also sampled the monkeys’ behavior and compared spatial association frequencies and rates of grooming among same- and opposite-sex dyads. There was no difference between males and females in genetic parameters; both males and females show low coefficients of relatedness indicating that neither sex is consistently philopatric. The mean pairwise coefficient of relatedness for co-resident females was not higher than that for co-resident males. Compared to other populations of capuchin monkeys, female bond was weak, as evident by lower spatial association frequencies, reduced rates of grooming and lack of correlation between coefficients of relatedness and measures of dyadic sociopositive interactions. Our findings thus confirm that female dispersal is a habitual process in the capuchin population of PECB, and that, as expected, dispersal by females strongly influences the relatedness structure of the population as well as the affiliative relationships among female groupmates.

A test for sex-biased dispersal in the black-faced lion tamarin (Leontopithecus caissara): inferences from microsatellite markers

In social systems with high reproductive skew, the likelihood of breeding may depend on the dispersal ability of non-reproductive members. The aim of the present study was to test through microsatellite genotyping whether males and females of the cooperative breeding callitrichid primate Leontopithecus caissara disperse at similar proportions. Our results indicate that there is no sex-bias in dispersal. However, even-sexed and sex-bias dispersal reported for other callitrichid populations indicate that environmental and demographic attributes may also influence the dispersal pattern. Therefore, dispersal patterns may vary in a more complex fashion than previously thought in callitrichids.

Population genetic structure and direct observations reveal sex-reversed patterns of dispersal in a cooperative bird.

Sex-biased dispersal is pervasive and has diverse evolutionary implications, but the fundamental drivers of dispersal sex biases remain unresolved. This is due in part to limited diversity within taxonomic groups in the direction of dispersal sex biases, which leaves hypothesis testing critically dependent upon identifying rare reversals of taxonomic norms. Here, we use a combination of observational and genetic data to demonstrate a rare reversal of the avian sex bias in dispersal in the cooperatively breeding white-browed sparrow weaver (Plocepasser mahali). Direct observations revealed that (i) natal philopatry was rare, with both sexes typically dispersing locally to breed, and (ii), unusually for birds, males bred at significantly greater distances from their natal group than females. Population genetic analyses confirmed these patterns, as (i) corrected Assignment index (AIc), FST tests and isolation-by-distance metrics were all indicative of longer dispersal distances among males than females, and (ii) spatial autocorrelation analysis indicated stronger within-group genetic structure among females than males. Examining the spatial scale of extra-group mating highlighted that the resulting ‘sperm dispersal’ could have acted in concert with individual dispersal to generate these genetic patterns, but gamete dispersal alone cannot account entirely for the sex differences in genetic structure observed. That leading hypotheses for the evolution of dispersal sex biases cannot readily account for these sex-reversed patterns of dispersal in white-browed sparrow weavers highlights the continued need for attention to alternative explanations for this enigmatic phenomenon. We highlight the potential importance of sex differences in the distances over which dispersal opportunities can be detected.

No sex-biased dispersal in a primate with an uncommon social system-cooperative polyandry.

An influential hypothesis proposed by Greenwood (1980) suggests that different mating systems result in female and male-biased dispersal, respectively, in birds and mammals. However, other aspects of social structure and behavior can also shape sex-biased dispersal. Although sex-specific patterns of kin cooperation are expected to affect the benefits of philopatry and dispersal patterns, empirical evidence is scarce. Unlike many mammals, Saguinus geoffroyi (Geoffroy’s tamarin) has a breeding system in which typically multiple males mate with a single breeding female. Males typically form cooperative reproductive partnerships between relatives, whereas females generally compete for reproductive opportunities. This system of cooperative polyandry is predicted to result in female-biased dispersal, providing an opportunity to test the current hypotheses of sex-biased dispersal. Here we test for evidence of sex-biased dispersal in S. geoffroyi using demographic and genetic data from three populations. We find no sex bias in natal dispersal, contrary to the prediction based on the mating patterns. This pattern was consistent after controlling for the effects of historical population structure. Limited breeding opportunities within social groups likely drive both males and females to disperse, suggesting that dispersal is intimately related to the social context. The integration of genetic and field data revealed that tamarins are another exception to the presumed pattern of male-biased dispersal in mammals. A shift in focus from mating systems to social behavior, which plays a role in most all processes expected to influence sex-bias in dispersal, will be a fruitful target for research both within species and across taxa.

Mate finding, Allee effects and selection for sex-biased dispersal.

Although dispersal requires context-dependent decision-making in three distinct stages (emigration, transit, immigration), these decisions are commonly ignored in simple models of dispersal. For sexually reproducing organisms, mate availability is an important factor in dispersal decisions. Difficulty finding mates can lead to an Allee effect where population growth decreases at low densities. Surprisingly, theoretical studies on mate finding and on sex-biased dispersal produce opposing predictions: in the former, one sex is predicted to move less if the other sex evolves to search more, whereas in the latter, mate-finding difficulties can select for less sex bias in dispersal when mate finding occurs after dispersal. Here, we develop a pair of models to examine the joint evolution of dispersal and settlement behaviour. Our first model resolves the apparent contradiction from the mate search and dispersal literatures. Our second model demonstrates that the relationship between mating system and sex-biased dispersal is more complex than a simple contrast between resource defence monogamy and female defence polygyny. Our results highlight that a key factor is the timing of mating relative to dispersal (before, during, or after). We also show that although movement has the potential to alleviate a mate-finding Allee effect, in some cases, it can actually exacerbate the effect.

The population structure of Glossina fuscipes fuscipes in the Lake Victoria basin in Uganda: implications for vector control

BackgroundGlossina fuscipes fuscipes is the primary vector of trypanosomiasis in humans and livestock in Uganda. The Lake Victoria basin has been targeted for tsetse eradication using a rolling carpet initiative, from west to east, with four operational blocks (3 in Uganda and 1 in Kenya), under a Pan-African Tsetse and Trypanosomiasis Eradication Campaign (PATTEC). We screened tsetse flies from the three Ugandan PATTEC blocks for genetic diversity at 15 microsatellite loci from continental and offshore populations to provide empirical data to support this initiative.MethodsWe collected tsetse samples from 11 sites across the Lake Victoria basin in Uganda. We performed genetic analyses on 409 of the collected tsetse flies and added data collected for 278 individuals in a previous study. The flies were screened across 15 microsatellite loci and the resulting data were used to assess the temporal stability of populations, to analyze patterns of genetic exchange and structuring, to estimate dispersal rates and evaluate the sex bias in dispersal, as well as to estimate demographic parameters (NE and NC).ResultsWe found that tsetse populations in this region were stable over 4-16 generations and belong to 4 genetic clusters. Two genetic clusters (1 and 2) corresponded approximately to PATTEC blocks 1 and 2, while the other two (3 and 4) fell within PATTEC block 3. Island populations grouped into the same genetic clusters as neighboring mainland sites, suggesting presence of gene flow between these sites. There was no evidence of the stretch of water separating islands from the mainland forming a significant barrier to dispersal. Dispersal rates ranged from 2.5 km per generation in cluster 1 to 14 km per generation in clusters 3 and 4. We found evidence of male-biased dispersal. Few breeders are successfully dispersing over large distances. Effective population size estimates were low (33–310 individuals), while census size estimates ranged from 1200 (cluster 1) to 4100 (clusters 3 and 4). We present here a novel technique that adapts an existing census size estimation method to sampling without replacement, the scheme used in sampling tsetse flies.ConclusionOur study suggests that different control strategies should be implemented for the three PATTEC blocks and that, given the high potential for re-invasion from island sites, mainland and offshore sites in each block should be targeted at the same time.

Genetic spatial autocorrelation can readily detect sex‐biased dispersal

Sex-biased dispersal is expected to generate differences in the fine-scale genetic structure of males and females. Therefore, spatial analyses of multilocus genotypes may offer a powerful approach for detecting sex-biased dispersal in natural populations. However, the effects of sex-biased dispersal on fine-scale genetic structure have not been explored. We used simulations and multilocus spatial autocorrelation analysis to investigate how sex-biased dispersal influences fine-scale genetic structure. We evaluated three statistical tests for detecting sex-biased dispersal: bootstrap confidence intervals about autocorrelation r values and recently developed heterogeneity tests at the distance class and whole correlogram levels. Even modest sex bias in dispersal resulted in significantly different fine-scale spatial autocorrelation patterns between the sexes. This was particularly evident when dispersal was strongly restricted in the less-dispersing sex (mean distance <200 m), when differences between the sexes were readily detected over short distances. All tests had high power to detect sex-biased dispersal with large sample sizes (n ≥ 250). However, there was variation in type I error rates among the tests, for which we offer specific recommendations. We found congruence between simulation predictions and empirical data from the agile antechinus, a species that exhibits male-biased dispersal, confirming the power of individual-based genetic analysis to provide insights into asymmetries in male and female dispersal. Our key recommendations for using multilocus spatial autocorrelation analyses to test for sex-biased dispersal are: (i) maximize sample size, not locus number; (ii) concentrate sampling within the scale of positive structure; (iii) evaluate several distance class sizes; (iv) use appropriate methods when combining data from multiple populations; (v) compare the appropriate groups of individuals.

Sex-Biased Dispersal and Volcanic Activities Shaped Phylogeographic Patterns of Extant Orangutans (genus: Pongo)

The Southeast Asian Sunda archipelago harbors a rich biodiversity with a substantial proportion of endemic species. The evolutionary history of these species has been drastically influenced by environmental forces, such as fluctuating sea levels, climatic changes, and severe volcanic activities. Orangutans (genus: Pongo), the only Asian great apes, are well suited to study the relative impact of these forces due to their well-documented behavioral ecology, strict habitat requirements, and exceptionally slow life history. We investigated the phylogeographic patterns and evolutionary history of orangutans in the light of the complex geological and climatic history of the Sunda archipelago. Our study is based on the most extensive genetic sampling to date, covering the entire range of extant orangutan populations. Using data from three mitochondrial DNA (mtDNA) genes from 112 wild orangutans, we show that Sumatran orangutans, Pongo abelii, are paraphyletic with respect to Bornean orangutans (P. pygmaeus), the only other currently recognized species within this genus. The deepest split in the mtDNA phylogeny of orangutans occurs across the Toba caldera in northern Sumatra and, not as expected, between both islands. Until the recent past, the Toba region has experienced extensive volcanic activity, which has shaped the current phylogeographic patterns. Like their Bornean counterparts, Sumatran orangutans exhibit a strong, yet previously undocumented structuring into four geographical clusters. However, with 3.50 Ma, the Sumatran haplotypes have a much older coalescence than their Bornean counterparts (178 kya). In sharp contrast to the mtDNA data, 18 Y-chromosomal polymorphisms show a much more recent coalescence within Sumatra compared with Borneo. Moreover, the deep geographic structure evident in mtDNA is not reflected in the male population history, strongly suggesting male-biased dispersal. We conclude that volcanic activities have played an important role in the evolutionary history of orangutans and potentially of many other forest-dwelling Sundaland species. Furthermore, we demonstrate that a strong sex bias in dispersal can lead to conflicting patterns in uniparentally inherited markers even at a genus-wide scale, highlighting the need for a combined usage of maternally and paternally inherited marker systems in phylogenetic studies.

Distinct physiological and behavioural functions for parental alleles of imprinted Grb10.

Imprinted genes, defined by their preferential expression of a single parental allele, represent a subset of the mammalian genome and often have key roles in embryonic development1, but also post-natal functions including energy homeostasis2 and behaviour3, 4. When the two parental alleles are unequally represented within a social group (when there is sex-bias in dispersal and/or variance in reproductive success)5, 6, imprinted genes may evolve to modulate social behaviour, although to date no such instance is known. Predominantly expressed from the maternal allele during embryogenesis, Grb10 encodes an intracellular adapter protein that can interact with a number of receptor tyrosine kinases and downstream signalling molecules7. Here we demonstrate that within the brain Grb10 is expressed from the paternal allele from fetal life into adulthood and that ablation of this expression engenders increased social dominance specifically among other aspects of social behaviour, a finding supported by the observed increase in allogrooming by paternal Grb10 deficient animals. Grb10 is, therefore, the first example of an imprinted gene that regulates social behaviour. It is also currently alone in exhibiting imprinted expression from each of the parental alleles in a tissue specific manner, as loss of the peripherally expressed maternal allele leads to significant fetal and placental overgrowth. Thus, Grb10 is to date a unique imprinted gene, able to influence distinct physiological processes, fetal growth and adult behaviour, due to actions of the two parental alleles in different tissues.

Dispersal, gene flow, and population genetic structure in the pygmy rabbit (Brachylagus idahoensis)

Dispersal and gene flow are significant components of metapopulation structure and dynamics. We examined gene flow in the pygmy rabbit (Brachylagus idahoensis), a sagebrush specialist that occurs in small, isolated populations in the Great Basin region and is believed to have limited dispersal abilities. We examined genetic diversity, gene flow, and population genetic structure, and investigated the effects of putative barriers to gene flow in 8 sample locations within 2 study areas in Idaho during 2002–2006. Potential barriers included secondary roads, highways, creeks, and agricultural pastures. Distance between sample locations ranged from 0.5 to 32 km. We expected that gene flow would be limited and that moderate levels of genetic structure would be apparent among sample locations separated by barriers and distances over several kilometers. However, analyses based on 15 microsatellite loci from 239 rabbits revealed low to moderate levels of genetic differentiation among sample locations. Bayesian cluster analyses suggested that sample locations separated by ,14 km comprised 1 genetic cluster with high levels of gene flow among sample locations within clusters. We documented no significant evidence of a sex bias in dispersal based on sex-specific FST among sample locations, and relatedness (r), mean assignment index (mAI), and variance of assignment index (vAI) within sample locations; however, spatial autocorrelation suggested that juvenile females might disperse farther than males. These results indicate that dispersal capabilities of pygmy rabbits greatly exceed previous estimates, and that creeks and roads were not effective barriers to gene flow in our study areas. DOI: 10.1644/09-MAMM-A032R.1.

Microsatellite Analyses Provide Evidence of Male-Biased Dispersal in the Radiated Tortoise Astrochelys radiata (Chelonia: Testudinidae)

Dispersal is a major force in shaping the genetic structure and dynamics of species; thus, its understanding is critical in formulating appropriate conservation strategies. In many species, sexes do not face the same evolutionary pressures, and consequently dispersal is often asymmetrical between males and females. This is well documented in birds and mammals but has seldom been investigated in other taxa, including reptiles and, more specifically, nonmarine chelonians. In these species, nest-site fidelity observations are frequent but still remain to be associated with natal homing. Here, we tested for sex-biased dispersal in the radiated tortoise (Astrochelys radiata) from southern Madagascar. Using data from 13 microsatellite markers, we investigated patterns of relatedness between sexes in 2 populations. All Mantel tests indicated significant isolation by distance at the individual level in females but not in males. Furthermore, spatial autocorrelation analyses and 2 analytical approaches designed to assess general trends in sex-specific dispersal also supported male-biased dispersal. On the other hand, comparisons of overall genetic structure among sampling sites did not provide conclusive support for greater philopatry in females, but these tests may have low statistical power because of methodological and biological constraints. Radiated tortoises appear to be both polyandrous and polygynous, and evolutionary processes that may lead to a sex bias in dispersal are discussed with respect to tortoise breeding biology. Female natal homing is hypothesized as a key trait explaining greater female philopatry in A. radiata. These findings highlight the necessity of additional research on natal homing in tortoises, a behavioral trait with direct implications for conservation.

Reverse sex‐biased philopatry in a cooperative bird: genetic consequences and a social cause

The genetic structure of a group or population of organisms can profoundly influence the potential for inbreeding and, through this, can affect both dispersal strategies and mating systems. We used estimates of genetic relatedness as well as likelihood-based methods to reconstruct social group composition and examine sex biases in dispersal in a Costa Rican population of white-throated magpie-jays (Calocitta formosa, Swainson 1827), one of the few birds suggested to have female-biased natal philopatry. We found that females within groups were more closely related than males, which is consistent with observational data indicating that males disperse upon maturity, whereas females tend to remain in their natal territories and act as helpers. In addition, males were generally unrelated to one another within groups, suggesting that males do not disperse with or towards relatives. Finally, within social groups, female helpers were less related to male than female breeders, suggesting greater male turnover within groups. This last result indicates that within the natal group, female offspring have more opportunities than males to mate with nonrelatives, which might help to explain the unusual pattern of female-biased philopatry and male-biased dispersal in this system. We suggest that the novel approach adopted here is likely to be particularly useful for short-term studies or those conducted on rare or difficult-to-observe species, as it allows one to establish general patterns of philopatry and genetic structure without the need for long-term monitoring of identifiable individuals.

Sex‐biased natal dispersal and inbreeding avoidance in American black bears as revealed by spatial genetic analyses

We tested the hypothesis that sex-biased natal dispersal reduces close inbreeding in American black bears, a solitary species that exhibits nearly complete male dispersal and female philopatry. Using microsatellite DNA and spatial data from reproductively mature bears (>or= 4 years old), we examined the spatial genetic structure of two distinct populations in New Mexico from 1993 to 2000. As predicted, relatedness (r) and the frequency of close relationships (parent-offspring or full siblings) decreased with distance among female dyads, but little change was observed among male or opposite-sex dyads. Neighbouring females were more closely related than neighbouring males. The potential for inbreeding was low. Most opposite-sex pairs that lived sufficiently close to facilitate mating were unrelated, and few were close relatives. We found no evidence that bears actively avoided inbreeding in their selection of mates from this nearby pool, as mean r and relationship frequencies did not differ between potential and actual mating pairs (determined by parentage analysis). These basic patterns were apparent in both study areas despite a nearly two-fold difference in density. However, the sex bias in dispersal was less pronounced in the lower-density area, based on proportions of bears with male and female relatives residing nearby. This result suggests that male bears may respond to reduced competition by decreasing their rate or distance of dispersal. Evidence supports the hypothesis that inbreeding avoidance is achieved by means of male-biased dispersal but also indicates that competition (for mates or resources) modifies dispersal patterns.

Sex Differences in Dispersal and the Evolution of Helping and Harming

In this article, we explore the impact of sex-biased dispersal on local relatedness and on selection for helping and harming behavior among males and females. We show that in a patch-structured population, when there is a marked sex bias in dispersal, selection will almost always favor harming behavior among individuals of the sex more prone to dispersal. This result holds regardless of the effects of mating skew or overlapping generations. Selection may well also favor helping behavior among individuals of the philopatric sex, particularly if there is generational overlap, but this is less likely to occur if individuals of the philopatric sex compete more intensely for fewer breeding opportunities. In this last case, if generational overlap is low and mating skew pronounced, the result may be selection for harming behavior among both males and females. In general, the rate of dispersal and the level of relatedness among individuals of one sex do not reliably predict their level of helping or harming behavior; selection on either males or females depends on the dispersal of both sexes.

Sex‐Biased Dispersal: A Result of a Sex Difference in Breeding Site Availability

Sex-biased dispersal is often explained by assuming that the resource-defending sex pays greater costs of moving from a familiar area. We hypothesize that sex-biased dispersal may also be caused by a sex bias in breeding site availability. In avian resource-defense mating systems, site availability is often more constrained for females: males can choose from all vacant sites, whereas females are restricted to sites defended by males. Using data on breeding dispersal of a migratory passerine, we show that average number of available breeding options and availability of the previous year's territory was greater for males than females. The female bias in site unavailability may explain the female bias in probability of breeding dispersal because there was no sex bias in dispersal among birds with their previous year's territory available. We suggest that sex biases in the availability of breeding options may be an important factor contributing to observed variation in sex-biased dispersal patterns.