Sex-biased dispersal in a salmonid fish

We tested the hypothesis that dispersal is sex biased in an unexploited population of brook trout, Salvelinus fontinalis, on Cape Race, Newfoundland, Canada. Based on the assumptions that trout are promiscuous and that reproductive success is limited primarily by either number of mates (males) or fecundity (females), we predicted that males would disperse greater distances than females. We also tested the hypothesis that trout populations comprise stationary and mobile individuals, predicting that males have greater mobility than females. Based on a mark-recapture study of 943 individually tagged fishes, 191 of which were recaptured over 5 years, we find strong support for our hypothesis of male-biased dispersal in brook trout. Averaged among all 11 resampling periods, males dispersed 2.5 times as far as females; during the spawning period only, male dispersal exceeded that by females almost fourfold. Both sexes were heterogeneous with respect to movement, with a lower incidence of mobility among females (29.6%) than males (41.1%); mobile males dispersed six times further than mobile females. We conclude that this sex bias reduces mate competition among male kin and decreases the probability that males will reproduce with related females.     

Sex-biased dispersal and adaptation to marginal habitats

If gene flow occurs through both sexes but only females contribute to population growth, adaptation to marginal (sink) habitats should be differentially affected by male versus female dispersal. Here I address this problem with two models. First, I consider the fate of a rare allele that improves fitness in the marginal habitat but reduces fitness in the core (source) habitat. Then I study the evolution of a polygenic character mediating a trade-off in fitness between the habitats. Both approaches led to qualitatively similar predictions. The effect of a difference in the dispersal rate between the sexes depends on the degree to which immigration from the core habitat boosts the reproductive output from the marginal habitat. This boost is slight if the marginal habitat is able to sustain well a population without immigration. In that case, both female- and male-biased dispersal is more favorable for adaptation to marginal habitats than equal dispersal of both sexes (assuming that the dispersal rate averaged over the sexes is kept constant). In contrast, if the marginal habitat is an absolute sink unable to sustain a population without immigration, the conditions for adaptation to that habitat are least favorable under highly male-biased dispersal and most favorable under highly female-biased dispersal. Under some circumstances, high average (male+female) dispersal is more favorable than low dispersal. Thus, gene flow should not be seen solely as thwarting adaptation to marginal habitats. The results are interpreted in terms of how male and female dispersal affects the relative rate of gene flow from the source to the sink habitat and in the opposite direction. This study predicts that ecological niches of taxa with female-biased dispersal should tend to be broader and more evolutionarily flexible.     

Sex-biased dispersal in the contemporary United States

Females in the contemporary United States disperse farther and are less philopatric than males, a pattern rare among mammals. This difference occurs primarily during the period of first independence following graduation from high School. I examine the patterns and possible causes of sex bias in internal migration using data derived from high school reunion booklets and a survey conducted on a sample of individuals selected from reunion booklets. The bias, which is small but significant, is largely eliminated when locality and socioeconomic factors are controlled. This suggests that these factors affect the sexes differentially. In general, females who move away to college or obtain jobs that are important to their residence are more likely to disperse relatively long distances, while males appear to be more constrained in their dispersal patterns. Female-biased dispersal (virilocal residence) in many nonindustrialized human societies is apparently a consequence of differences between the sexes in patterns of resource accumulation, exemplified by patrilinial inheritance. Similar differences still characterize, to a lesser extent, modern industrialized societies, and may be responsible, at least in part, for the pattern of sex-biased dispersal found in this study.     

Sex-biased dispersal in eastern chipmunks, Tamias striatus

I investigated the effect of male mate competition and inbreeding avoidance on natal dispersal of chipmunks by longitudinally monitoring known individuals from 1986 to 1990. Natal males exhibited greater absolute and effective dispersal distances but dispersed at the same proportion as natal females. Recruitment of juvenile males was negatively affected by density of resident males, but there was no evidence of local mate competition among male kin. Analysis of the spatial distribution of neighbors showed that natal males settled farther from their mothers than did their female siblings and farther than unrelated juvenile males. In addition, mothers apparently tolerated daughters as close neighbors and occasionally shared den sites with grandprogeny. Sexually mature males were never neighbors of their mothers and were never observed at maternal mating bouts. Males may disperse to improve reproductive opportunities by avoiding competition with resident males, and by increasing access to unrelated females. Maternal tolerance of daughters but not sons may result in the close affiliation between mothers and daughters, and indirectly contribute to dispersal of natal males. Hence male-biased dispersal could be a consequence of mate competition and maternal avoidance of incestuous matings. This revised version was published online in July 2006 with corrections to the Cover Date.     

Sex-biased dispersal and the speed of two-sex invasions

Population models that combine demography and dispersal are important tools for forecasting the spatial spread of biological invasions. Current models describe the dynamics of only one sex (typically females). Such models cannot account for the sex-related biases in dispersal and mating behavior that are typical of many animal species. In this article, we construct a two-sex integrodifference equation model that overcomes these limitations. We derive an explicit formula for the invasion speed from the model and use it to show that sex-biased dispersal may significantly increase or decrease the invasion speed by skewing the operational sex ratio at the invasion's low-density leading edge. Which of these possible outcomes occurs depends sensitively on complex interactions among the direction of dispersal bias, the magnitude of bias, and the relative contributions of females and males to local population growth.     

Sex-biased dispersal and inbreeding avoidance in birds and mammals

Sex differences in dispersal distance are widespread in birds and mammals, but the predominantly dispersing sex differs consistently between the classes. There has been persistent debate over the relative importance of two factors - intrasexual competition and inbreeding avoidance - in producing sex-biased dispersal, and over the sources of the difference in dispersal patterns between the two classes. Recent studies cast new light on these questions.     

Two-patch population models with adaptive dispersal: the effects of varying dispersal speeds

The population-dispersal dynamics for predator–prey interactions and two competing species in a two patch environment are studied. It is assumed that both species (i.e., either predators and their prey, or the two competing species) are mobile and their dispersal between patches is directed to the higher fitness patch. It is proved that such dispersal, irrespectively of its speed, cannot destabilize a locally stable predator–prey population equilibrium that corresponds to no movement at all. In the case of two competing species, dispersal can destabilize population equilibrium. Conditions are given when this cannot happen, including the case of identical patches.     

Sex-biased dispersal of adults mediates the evolution of altruism among juveniles

Population viscosity has been proposed as an important mechanism for the evolution of cooperation. The idea is that if individuals do not disperse far during the course of their lives, they will tend to interact with their genealogical relatives, which may give kin-selected benefits for cooperation. However, in the simplest model of population structure, the evolution of cooperation is unaffected by the rate of dispersal, owing to dispersal also mediating competition between social partners. This surprising result has generated much research interest in recent years. Here I show that dispersal does matter if there is a sex difference in dispersal rate, even when the expression of cooperation is not conditional upon the actor's dispersal status or sex. In particular, I show that cooperation among juveniles is relatively favoured when there is a small sex bias in adult dispersal in favour of the sex with the greatest variance in reproductive success, and is relatively disfavoured when this sex bias is large or in the opposite direction. This is because dispersal by individuals of each sex can have different consequences for the genetic structure of the population.     

Sex-biased dispersal in western lowland gorillas (Gorilla gorilla gorilla)

We explored two hypotheses related to potential differences between sexes in dispersal behaviour in western lowland gorillas (Gorilla gorilla gorilla). Direct observations suggest that immature females have more opportunities to move between breeding groups than immature males. The distribution of kin dyadic relationships within and between groups does not, however, support this hypothesis. At larger geographical scales, dispersal is likely to be easier for males than females because of the solitary phase most blackbacks experience before founding their own breeding group. However, previous work indicates that males settle preferentially close to male kin. By specifically tracing female and male lineages with mitochondrial and Y-chromosomal genetic markers, we found that male gorillas in the 6000 km2 area we surveyed form a single population whereas females are restricted to the individual sites we sampled and do not freely move around this area. These differences are more correctly described as differences in dispersal distances, rather than differences in dispersal rates between sexes (both sexes emigrate from their natal group in this species). Differences in resource competition and dispersal costs between female and male gorillas are compatible with the observed pattern, but more work is needed to understand if these ultimate causes are responsible for sex-biased dispersal distances in western lowland gorillas.     

Sex-biased dispersal in a rare butterfly and the implications for its conservation

The survival of many species may be dependent on their ability to exist in human-altered landscapes within metapopulations; in turn, metapopulation persistence is dictated by the ability of individuals to move effectively among patches to promote recolonization. The Taylor’s checkerspot butterfly (Euphydryas editha taylori) is a species that does not naturally occur in fragmented landscapes, yet it is now restricted to a handful of small isolated prairie habitats. Current recovery plans aim to establish a stable metapopulation; however, to date little is known about the species’ ability to move across the landscape. In 2010 and 2011, we conducted marking, tracking and boundary surveys to explore the movement dynamics of adults within two sites in Oregon, USA. Over the survey period, we marked 136 male butterflies, tracked 174 individuals and observed the behavior of 1,576 individual butterflies at site boundaries. Our study revealed a significant sex-bias in the movement dynamics of the Taylor’s checkerspot in both suitable habitat and surrounding matrix. Males were highly motile, whereas females appeared sedentary, rarely moving from their natal site. The limited dispersal behavior of females indicates that populations cannot persist naturally in a metapopulation and thus are at high risk of extinction. Based on our findings, we recommend that managers take proactive measures to increase or enable dispersal (including translocation) to existing and/or restored sites.     

Sex-biased dispersal and natal philopatry in the diamondback terrapin, Malaclemys terrapin

Nesting ecology and population studies indicate that diamondback terrapins (Malaclemys terrapin) exhibit nest site fidelity and high habitat fidelity. However, genetic studies indicate high levels of gene flow. Because dispersal affects the genetics and population dynamics of a species, we used six highly polymorphic microsatellite markers to investigate sex-biased dispersal and natal philopatry of M. terrapin in Barnegat Bay, NJ. We compared results of spatial autocorrelation analysis, assignment methods and Wright's F(ST) estimators to a mark-recapture analysis. Mark-recapture analysis over a 4-year period indicated that most individuals have relatively small home ranges (<2 km), with mature females displaying greater home ranges than males. Goodness of fit analysis of our mark-recapture study indicated that some juvenile males were likely transient individuals moving through our study location. Mean assignment indices and first-generation migrant tests indicated that mature males were more prone to disperse than mature females, but first-generation migrant tests indicated that per capita there are more female than male dispersers. Thus, the relative importance of males and females on gene flow in terrapin populations may change in relation to population sex ratios. Spatial autocorrelation analysis indicated that mature females exhibited natal philopatry to nesting beaches, but first-generation migrant tests indicated that a small number of females failed to nest on natal beaches. Finally, we discuss the important conservation implications of male-biased dispersal and natal philopatry in the diamondback terrapin.     

Sex-biased natal dispersal in Hokkaido brown bears revealed through mitochondrial DNA analysis

Understanding natal dispersal patterns is fundamental in the ecology and conservation biology of large wild carnivores. In this study, we used two approaches to determine genetic variation and dispersal patterns of brown bears in the Shiretoko Peninsula, eastern Hokkaido, Japan. The first approach was a large-scale genetic analysis. We analyzed haplotypes from the mitochondrial DNA (mtDNA) control region of 760 individual samples collected throughout the peninsula during 1998–2016. We detected seven haplotypes, including two that were confirmed for the first time. In females, the distribution of haplotypes was geographically structured, whereas haplotypes in males were distributed widely throughout the peninsula. Only some males in the lower peninsula had haplotypes that were not detected within the peninsula. The second approach was a local-scale genetic analysis, including intensive focal sampling in the Rusha area, a special wildlife protection area on the peninsula. Proportions of mtDNA haplotypes in adult bears were investigated and compared between the sexes. Although more than half of the females had the same haplotype, males had more diverse haplotypes, suggesting that they came to the Rusha area from other regions. Thus, our study revealed that mtDNA haplotype distribution has been maintained by female philopatry, and that bears exhibit male-biased dispersal. Furthermore, the lower peninsula appears to act as a contact zone between the peninsula and mainland Hokkaido, which is important for maintaining genetic diversity.     

Sex-biased dispersal, haplodiploidy and the evolution of helping in social insects

In his famous haplodiploidy hypothesis, W. D. Hamilton proposed that high sister-sister relatedness facilitates the evolution of kin-selected reproductive altruism among Hymenopteran females. Subsequent analyses, however, suggested that haplodiploidy cannot promote altruism unless altruists capitalize on relatedness asymmetries by helping to raise offspring whose sex ratio is more female-biased than the population at large. Here, we show that haplodiploidy is in fact more favourable than is diploidy to the evolution of reproductive altruism on the part of females, provided only that dispersal is male-biased (no sex-ratio bias or active kin discrimination is required). The effect is strong, and applies to the evolution both of sterile female helpers and of helping among breeding females. Moreover, a review of existing data suggests that female philopatry and non-local mating are widespread among nest-building Hymenoptera. We thus conclude that Hamilton was correct in his claim that 'family relationships in the Hymenoptera are potentially very favourable to the evolution of reproductive altruism'.     

Sex-biased dispersal patterns depend on the spatial scale in a social rodent

Dispersal is a fundamental process in ecology because it influences the dynamics, genetic structure and persistence of populations. Furthermore, understanding the evolutionary causes of dispersal pattern, particularly when they differ between genders, is still a major question in evolutionary ecology. Using a panel of 10 microsatellite loci, we investigated at different spatial scales the genetic structure and the sex-specific dispersal patterns in the common vole Microtus arvalis, a small colonial mammal. This study was conducted in an intensive agricultural area of western France. Hierarchical F_ST analyses, relatedness and assignment tests suggested (i) that females are strongly kin-clustered within colonies; (ii) that dispersal is strongly male-biased at a local scale; and (iii) long-distance dispersal is not rare and more balanced between genders. We conclude that males migrate continuously from colony to colony to reproduce, whereas females may disperse just once and would be mainly involved in new colony foundation.     

洞庭湖区自然野化麋鹿种群的偏性扩散行为

大多数鹿科动物表现出偏向雄性的扩散行为。麋鹿(Elaphurus davidianus)于20世纪初在中国灭绝,1985年被重引入中国。1998年长江流域特大洪水导致部分麋鹿个体从湖北石首麋鹿国家级自然保护区围栏内外逸,最终在洞庭湖区形成自然野化种群。野化麋鹿是否也存在偏性扩散行为?1995—2012年,采用样带调查法、分区直数法、特殊个体识别法和访问调查法,按性别组成将扩散群分为雄性群、混合群和雌性群,对洞庭湖区自然野化麋鹿种群的扩散行为进行了研究,共记录到118次扩散。其中,有65次扩散可确认扩散群的类型;60次可确认扩散群准确的个体数。结果显示:(1)雄性麋鹿较雌性更倾向于扩散。雄性群的扩散频次高于雌性群和混合群;50%的雄性扩散群仅由单一成年雄体组成。(2)雄性麋鹿的扩散能力最强,雄性群、混合群和雌性群的平均扩散距离分别为(13.73±8.74)km、(11.05±4.16)km和(8.95±2.16)km,但三者之间的差异不显著(χ2=1.896,df=2,P=0.387)。雌性群和混合群的扩散距离均短于25km,而5.88%的雄性群扩散距离长于25 km。(3)雄性群的平均个体数与混合群的差异显著(F=5.324,df=24,P=0.000.05),与雌性群的差异不显著(F=9.830,df=35,P=0.813),而混合群与雌性群之间的差异显著(F=48.085,df=55,P=0.000.05)。(4)50.00%的雄性群和53.57%的混合群选择芦苇草地作为扩散目的地的生境。洞庭湖区自然野化麋鹿种群与其他野生鹿科动物一样,也存在"雄鹿较雌鹿更倾向于扩散"的现象,雄性的扩散能力比雌性强,雌性的扩散可能依赖于雄性。以上结果对麋鹿的野外放归和自然野化种群的管理与保护具有重要的意义。     

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.     

Sex-biased dispersal in a migratory bat: a characterization using sex-specific demographic parameters

We studied the noctule bat (Nyctalus noctula), in which the mitochondrial F(ST) is about 10 times that revealed by nuclear markers, to address two questions. We first verified whether random dispersal of one sex is compatible with highly contrasted mitochondrial and nuclear population structures. Using computer simulations, we then assessed the power of multilocus population differentiation tests when the expected population structure departs only slightly from panmixia. Using an island model with sex-specific demographic parameters, we found that random male dispersal is consistent with the population structure observed in the noctule. However, other parameter combinations are also compatible with the data. We computed the minimum sex bias in dispersal (at least 69% of the dispersing individuals are males), a result that would not be available if we had used more classical population genetic models. The power of multilocus population differentiation tests was unexpectedly high, the tests being significant in almost 100% of the replicates, although the observed population structure infered from nuclear markers was extremely low (F(ST) = 0.6%).