Sex and landscape influence spatial genetic variation in a large fossorial mammal, the Bare-nosed Wombat (Vombatus ursinus).

Dispersal is an important process that is widely studied across species, and it can be influenced by intrinsic and extrinsic factors. Intrinsic factors commonly assessed include the sex and age of individuals, while landscape features are frequently-tested extrinsic factors. Here, we investigated the effects of both sex and landscape composition and configuration on genetic distances among bare-nosed wombats (Vombatus ursinus)-one of the largest fossorial mammals in the world and subject to habitat fragmentation, threats from disease, and human persecution including culling as an agricultural pest. We analyzed a data set comprising 74 Tasmanian individuals (30 males and 44 females), genotyped for 9,064 single-nucleotide polymorphisms. We tested for sex-biased dispersal and the influence of landscape features on genetic distances including land use, water, vegetation, elevation, and topographic ruggedness. We detected significant female-biased dispersal, which may be related to females donating burrows to their offspring due to the energetic cost of excavation, given their large body sizes. Land use, waterbodies, and elevation appeared to be significant landscape predictors of genetic distance. Land use potentially reflects land clearing and persecution over the last 200 years. If our findings based on a limited sample size are valid, retention and restoration of nonanthropogenic landscapes in which wombats can move and burrow may be important for gene flow and maintenance of genetic diversity.

Genetic Analysis Reveals Dispersal Patterns of Japanese Serow in Two Different Habitats of a Mountainous Region.

Dispersal increases the costs of feeding and predation risk in the new environment and is reported to be biased toward habitats similar to the natal region in some mammals. The benefits and costs of dispersal often differ between sexes, and most mammals show male-biased dispersal in relation to a polygamous mating system. Japanese serow is generally a solitary and monogamous species. However, recent studies have shown that the sociality of serows on Mt. Asama differs between habitat types. In the mountain forests with low forage availability, solitary habits and social monogamy were observed, while, in alpine grasslands, female grouping and social polygyny were observed, which is probably due to abundant forage availability. We investigated the effects of habitat characteristics and sociality on the dispersal of serows using fecal and tissue samples from two different habitats on Mt. Asama. The Fst value between the two areas was significantly positive, and the mean relatedness within areas was significantly higher than that between areas, which suggests limited gene flow and natal habitat-biased dispersal. Bayesian clustering analysis showed unidirectional gene flow from forest to grassland, which was probably due to the high forage availability of the grassland. Analyses of the assignment index and mean relatedness did not show male-biased dispersal, even in the grassland, where serows were polygynous. Thus, polygyny in the grassland is not linked to male-biased dispersal. In summary, our study suggests that dispersal patterns in Japanese serows are affected by habitat rather than social differences.

A reanalysis of strontium isotope ratios as indicators of dispersal in South African hominins.

Dispersal patterns in primates have major implications for behavior and sociality but are difficult to reconstruct for fossil species. This study applies novel strontium isotope methodologies that have reliably predicted philopatry and dispersal patterns in chimpanzees and other modern primates to previously published strontium isotope ratios (87Sr/86Sr) of two South African hominins, Australopithecus africanus and Australopithecus robustus. In this study, the difference or 'offset' was calculated between the 87Sr/86Sr of each fossil tooth compared to local bioavailable 87Sr/86Sr as defined by cluster analysis of modern plant isotope ratios. Large teeth (presumably belonging to males) have low offsets from local 87Sr/86Sr proxies, while small teeth (presumably from females) have greater offsets from local 87Sr/86Sr proxies. This supports previous conclusions of male philopatry and female dispersal in both A. africanus and A. robustus. Furthermore, A. robustus shows more extreme differences between presumed males and females compared to A. africanus. This is analogous to differences seen in modern olive baboons compared to chimpanzees and suggests that A. africanus may have had a larger home range than A. robustus. Neither hominin species has 87Sr/86Sr consistent with riparian habitat preferences despite the demonstrated presence of riparian habitats in South Africa at the time.

The disperser dilemma in cooperatively breeding birds.

In most cooperatively breeding birds, individuals do not breed with their natal group members. In order to breed, they have either to disperse into another group or wait for an opposite-sex individual to join their group. In most of these species, females disperse more than males. We develop a dynamic game-theoretic model to account for this asymmetry. When males are physically larger/heavier than females, this allows them to effectively welcome female immigrants into their natal group and overcome the local females' opposition more than vice versa. The model further assumes that the dispersal decision is not confined to a restricted time window, but is rather based on acquired information and responsive to opportunities. The model predicts that (i) females disperse more than males, and (ii) females are willing to tolerate more risks in dispersal than do males. The latter prediction is supported inter alia by the fact that in many cooperatively breeding birds, females disperse at a younger age, and further away from their natal group as compared to dispersing males.

Timing and distance of natal dispersal in Asian black bears.

Dispersal has important implications for population ecology and genetics of a species through redistribution of individuals. In most mammals, males leave their natal area before they reach sexual maturity, whereas females are commonly philopatric. Here, we investigate the patterns of natal dispersal in the Asian black bear (Ursus thibetanus) based on data from 550 bears (378 males, 172 females) captured or removed in Gunma and Tochigi prefectures on central Honshu Island, Japan in 2003-2018. We used genetic data and parentage analysis to investigate sex-biased differences in the distance of natal dispersal. We further investigated the age of dispersal using spatial autocorrelation analysis, that is, the change in the correlation between genetic and geographic distances in each sex and age group. Our results revealed that male dispersal distances (mean ± SE = 17.4 ± 3.5 km) were significantly farther than female distances (4.8 ± 1.7 km), and the results were not affected by years of mast failures, a prominent forage source for this population. Based on an average adult female home range radius of 1.8 km, 96% of the males and 50% of the females dispersed. In the spatial autocorrelation analysis, the changes in the relationship between genetic and geographic distances were more pronounced in males compared to females. Males seem to mostly disperse at age 3 regardless of mast productivity, and they gradually disperse far from their home range, but young and inexperienced males may return to their natal home range in years with poor food conditions. The results suggest that factors driving the dispersal process seem to be population structure-based instead of forage availability-based. In females, a significant genetic relationship was observed among all individuals in the group with a minimum age of 6 years within a distance of 2 km, which resulted in the formation of matrilineal assemblages.

(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.

Sex specificity of dispersal behaviour and flight morphology varies among tree hollow beetle species.

BACKGROUND: Flight performance and dispersal behaviour can differ between sexes, resulting in sex-biased dispersal. The primary sex ratio of populations may also explain dispersal bias between sexes, as this bias may evolve with the primary sex ratio to reduce intrasexual competition. Although dispersal bias between sexes is relevant to population dynamics, there are few studies on sex-biased dispersal in insects. We studied the flight performance and dispersal behaviour of seven saproxylic beetle species associated with tree hollows from a sex perspective. We also analysed the possible coevolution of flight performance with the primary sex ratio. METHODS: Wing loading and wing aspect ratio were used as measures of the flight performance of species and sexes. Dispersal behaviour was explored by analysing the frequency of each sex in interception traps versus the primary sex ratio obtained by tree hollow emergence traps using contingency tables and posthoc standardized residuals. A more active flight behaviour was expected for the sex with higher capture frequency in the interception traps. To explore the causes of flight performance bias between sexes, we searched for possible correlations between wing loading or wing aspect ratio and primary sex ratio using Pearson's correlation coefficient. RESULTS: Wing loading and wing aspect ratio differed between species and sexes, with flight performance being higher in males than in females for four of the seven species analysed. Dispersal behaviour and flight performance matched in the case of Elater ferrugineus; males showed higher flight performance and were the most collected sex in the interception traps (more active flyers). In contrast, the higher flight activity of Cetonia carthami aurataeformis females was not correlated with a higher flight performance than that of males. Moreover, we found that a bias in the primary sex ratio towards females is often correlated with a decrease in female flight performance. CONCLUSIONS: We stress that flight performance and dispersal behaviour of sexes do not always go hand in hand. Moreover, the relationship between the sex ratio and flight performance bias between sexes is not driven by competition within the most abundant sex. The inclusion of a sex perspective in insect dispersal studies would be useful to detect dispersal bias between sexes and its causes and would allow for further analysis of its effects on population dynamics.

Genetic diversity and sex-biased dispersal in the brown spotted pitviper (Protobothrops mucrosquamatus): Evidence from microsatellite markers.

Dispersal plays a vital role in the geographical distribution, population genetic structure, quantity dynamics, and evolution of a species. Sex-biased dispersal is common among vertebrates and many studies have documented a tendency toward male-biased dispersal in mammals and female-biased dispersal in birds. However, dispersal patterns in reptiles remain poorly understood. In this study, we explored the genetic diversity and dispersal patterns of the widely distributed Asian pitviper Protobothrops mucrosquamatus. In total, 16 polymorphic microsatellite loci were screened in 150 snakes (48 males, 44 females, 58 samples without sex information) covering most of their distribution. Microsatellite analysis revealed high genetic diversity in P. mucrosquamatus. Bayesian clustering of population assignment identified two major clusters for all populations, somewhat inconsistent with the mitochondrial DNA phylogeny of P. mucrosquamatus reported in previous research. Analyses based on 92 sex-determined and 37 samples of P. mucrosquamatus from three small sites in Sichuan, China (Mingshan, Yibin, and Zizhong) consistently suggested female-biased dispersal in P. mucrosquamatus, which is the first example of this pattern in snakes. The female-biased dispersal patterns in P. mucrosquamatus may be explained by local resource competition.

Avian Introgression Patterns are Consistent With Haldane's Rule.

According to Haldane's Rule, the heterogametic sex will show the greatest fitness reduction in a hybrid cross. In birds, where sex is determined by a ZW system, female hybrids are expected to experience lower fitness compared to male hybrids. This pattern has indeed been observed in several bird groups, but it is unknown whether the generality of Haldane's Rule also extends to the molecular level. First, given the lower fitness of female hybrids, we can expect maternally inherited loci (i.e., mitochondrial and W-linked loci) to show lower introgression rates than biparentally inherited loci (i.e., autosomal loci) in females. Second, the faster evolution of Z-linked loci compared to autosomal loci and the hemizygosity of the Z-chromosome in females might speed up the accumulation of incompatible alleles on this sex chromosome, resulting in lower introgression rates for Z-linked loci than for autosomal loci. I tested these expectations by conducting a literature review which focused on studies that directly quantified introgression rates for autosomal, sex-linked, and mitochondrial loci. Although most studies reported introgression rates in line with Haldane's Rule, it remains important to validate these genetic patterns with estimates of hybrid fitness and supporting field observations to rule out alternative explanations. Genomic data provide exciting opportunities to obtain a more fine-grained picture of introgression rates across the genome, which can consequently be linked to ecological and behavioral observations, potentially leading to novel insights into the genetic mechanisms underpinning Haldane's Rule.

Spatial population genetic structure and colony dynamics in Damaraland mole-rats (Fukomys damarensis) from the southern Kalahari.

BACKGROUND: Non-random associations within and among groups of social animals can provide valuable insight into the function of group living and the evolution of social behaviour. Damaraland mole-rats (Fukomys damarensis) demonstrate extremely high levels of reproductive skew, and dispersal is considered to be male-biased in onset and frequency, although asymmetry in dispersal distance is yet to be investigated. Dispersal may be positively correlated with increasing favourable environmental conditions, such as rainfall, however, the effects of ecological constraints on dispersal and colony fission-fusion dynamics have not previously been demonstrated on a spatial scale. Here we provide the first spatial population genetic study for this species. We investigated genetic structure in a population of Damaraland mole-rats from the southern Kalahari in South Africa over 3 years, combining observational dispersal data from mark-recapture with population genetic data to evaluate (1) sex-bias in frequency and distance of dispersal in this species, and (2) the effect of rainfall on fission-fusion dynamics of colonies. RESULTS: Our results demonstrate (1) that both males and females favour local dispersal but on rare occasions may disperse over distances greater than 400 m, (2) that males may disperse over greater distances than females, and (3) that males more frequently immigrate into established neighbouring colonies than females, who predominantly disperse by colony fission, i.e. multiple individuals "budding" from their native colony into a neighbouring territory, thereby establishing new colonies. Furthermore, our results demonstrate (4) elevated dispersal and colony fission in association with increased rainfall, supporting the hypothesis that rainfall may play a significant role in the maintenance and/or disruption of reproductive skew in Damaraland mole-rat populations. CONCLUSION: This study represents the first fine-scale spatial population genetic study in Damaraland mole-rats, and provides relevant insights into colony fission-fusion dynamics in a social and cooperatively breeding species.

Weak spatial-genetic structure in a native invasive, the southern pine beetle (Dendroctonus frontalis), across the eastern United States.

The southern pine beetle, Dendroctonus frontalis, is a native pest of pine trees that has recently expanded its range into the northeastern United States. Understanding its colonization, dispersal, and connectivity will be critical for mitigating negative economic and ecological impacts in the newly invaded areas. Characterization of spatial-genetic structure can contribute to this; however, previous studies have reached different conclusions about regional population genetic structure, with one study reporting a weak east-west pattern, and the most recent reporting an absence of structure. Here we systematically assessed several explanations for the absence of spatial-genetic structure. To do this, we developed nine new microsatellite markers and combined them with an existing 24-locus data matrix for the same individuals. We then reanalyzed this full dataset alongside datasets in which certain loci were omitted with the goal of creating more favorable signal to noise ratios. We also partitioned the data based on the sex of D. frontalis individuals, and then employed a broad suite of genotypic clustering and isolation-by-distance (IBD) analyses. We found that neither inadequate information content in the molecular marker set, nor unfavorable signal-to-noise ratio, nor insensitivity of the analytical approaches could explain the absence of structure. Regardless of dataset composition, there was little evidence for clusters (i.e., distinct geo-genetic groups) or clines (i.e., gradients of increasing allele frequency differences over larger geographic distances), with one exception: significant IBD was repeatedly detected using an individual-based measure of relatedness whenever datasets included males (but not for female-only datasets). This is strongly indicative of broad-scale female-biased dispersal, which has not previously been reported for D. frontalis, in part owing to logistical limitations of direct approaches (e.g., capture-mark-recapture). Weak spatial-genetic structure suggests long-distance connectivity and that gene flow is high, but additional research is needed to understand range expansion and outbreak dynamics in this species using alternate approaches.

Cooperative interactions among females can lead to even more extraordinary sex ratios.

Hamilton's local mate competition theory provided an explanation for extraordinary female-biased sex ratios in a range of organisms. When mating takes place locally, in structured populations, a female-biased sex ratio is favored to reduce competition between related males, and to provide more mates for males. However, there are a number of wasp species in which the sex ratios appear to more female biased than predicted by Hamilton's theory. It has been hypothesized that the additional female bias in these wasp species results from cooperative interactions between females. We investigated theoretically the extent to which cooperation between related females can interact with local mate competition to favor even more female-biased sex ratios. We found that (i) cooperation between females can lead to sex ratios that are more female biased than predicted by local competition theory alone, and (ii) sex ratios can be more female biased when the cooperation occurs from offspring to mothers before dispersal, rather than cooperation between siblings after dispersal. Our models formally confirm the verbal predictions made in previous experimental studies, which could be applied to a range of organisms. Specifically, cooperation can help explain sex ratio biases in Sclerodermus and Melittobia wasps, although quantitative comparisons between predictions and data suggest that some additional factors may be operating.

Blow flies, synanthropy and sex ratio: Are the deviations in the sex proportion linked to human transformation of landscapes?

In most species, several factors like time of emergence, age at maturation, reproductive life span, survival of males and females, mating behavior, differential resource use, and migration patterns may affect the adult sex ratio. Anthropogenic landscape transformation is known to change diversity, favoring colonization by exotic species but other populational parameters, such as the sex ratio, have not been assessed. The aim of the present study was to describe the sex-ratio patterns of adult necrophagous blow flies captured using carrion-baited traps along habitats representing different levels of human impact. We describe the sex bias for four species: the exotic Chrysomya albiceps, Chrysomya megacephala, Chrysomya putoria, and the native Cochliomyia macellaria. Three types of habitats were selected: highly anthropized (urbanized), moderately anthropized (rural), and native forest (natural) within a humid subtropical ecoregion in Northeastern Argentina. We found an overall trend to female bias among the four species when considering the total number of each sex. However, our study showed a changing sex-ratio pattern along the gradient of human influence. Our results indicate that a higher likelihood of trapping more males in unsuitable habitats seems to be widespread among blow flies. Urban exploiters, such as Ch. megacephala and Ch. putoria, locally shifted their trends, becoming male-biased in natural habitats. The opposite trend was detected in the urban avoider Co. macellaria (which shifted to male-biased in urban habitats). The exception was Ch. albiceps, whose sex proportion did not shift to a male-biased sex ratio. The results of this study highlight the changing sex-ratio patterns displayed by Calliphoridae in response to different conditions along a human-influence gradient. The biological traits and underlying mechanisms promoting the intraspecific changes of the sex ratio are discussed.

Dispersal and mating patterns determine the fate of naturally dispersed populations: evidence from Bombina orientalis.

BACKGROUND: In contrast to the explosive increase of a population following biological invasion, natural dispersal, i.e., when a population disperses from its original range into a new range, is a passive process that is affected by resources, the environment, and other factors. Natural dispersal is also negatively impacted by genetic drift and the founder effect. Although the fates of naturally dispersed populations are unknown, they can adapt evolutionarily over time to the new environment. Can naturally dispersed populations evolve beneficial adaptive strategies to offset these negative effects to maintain their population in a stable state? RESULTS: The current study addressed this question by focusing on the toad Bombina orientalis, the population of which underwent natural dispersal following the Last Glacial Maximum in Northeast Asia. Population genetic approaches were used to determine the genetic structure, dispersal pattern, and mating system of the population of B. orientalis in northeast China (Northern population). The results showed that this northern population of B. orientalis is a typical naturally dispersed population, in which the stable genetic structure and high level of genetic diversity of the population have been maintained through the long-distance biased dispersal behavior of males and the pattern of promiscuity within the population. CONCLUSIONS: Our findings suggest that naturally dispersed populations can evolve effective adaptive strategies to maintain a stable population. Different species may have different strategies. The relevance of these maintenance mechanisms for naturally dispersed populations provide a new perspective for further understanding the processes of speciation and evolution.

Selection against immigrants in wild seabird populations.

Immigration is a major demographic parameter shaping population dynamics and is an important driver of eco-evolutionary patterns, but the fitness consequences for individuals following their settlement to a new population (immigrants) remain poorly tested in wild animal populations, particularly among long-lived species. Here we show that immigrants have a lower fitness than residents in three wild seabird populations (wandering albatross Diomedea exulans, southern fulmar Fulmarus glacialoides, snow petrel Pagodroma nivea). Across all species and during a 32-year period, immigrants made on average -9 to 29% fewer breeding attempts, had 5-31% fewer fledglings, had 2-16% lower breeding success and produced 6-46% fewer recruits. Female immigration and male residency were also favored through differences in breeding performance. We provide evidence for selection against immigrants in wild populations of long-lived species and our results are consistent with female-biased dispersal in birds being driven by asymmetric limiting resources and the competitive ability of dispersers vs. non-dispersers.

Genetic structure of recently fragmented suburban populations of European stag beetle.

Habitat loss and fragmentation due to urbanization can negatively affect metapopulation persistence when gene flow among populations is reduced and population sizes decrease. Inference of patterns and processes of population connectivity derived from spatial genetic analysis has proven invaluable for conservation and management. However, a more complete account of population dynamics may be obtained by combining spatial and temporal sampling. We, therefore, performed a genetic study on European stag beetle (Lucanus cervus L.) populations in a suburban context using samples collected in three locations and during the period 2002-2016. The sampling area has seen recent landscape changes which resulted in population declines. Through the use of a suite of F ST, clustering analysis, individual assignment, and relatedness analysis, we assessed fine scale spatiotemporal genetic variation within and among habitat patches using 283 individuals successfully genotyped at 17 microsatellites. Our findings suggested the three locations to hold demographically independent populations, at least over time scales of relevance to conservation, though with higher levels of gene flow in the past. Contrary to expectation from tagging studies, dispersal appeared to be mainly female-biased. Although the life cycle of stag beetle suggests its generations to be discrete, no clear temporal structure was identified, which could be attributed to the varying duration of larval development. Since population bottlenecks were detected and estimates of effective number of breeders were low, conservation actions are eminent which should include the establishment of suitable dead wood for oviposition on both local and regional scales to increase (re)colonization success and connectivity among current populations.

Density-dependent sex-biased development of macroptery in a water strider.

In wing-polymorphic insects, wing morphs differ not only in dispersal capability but also in life history traits because of trade-offs between flight capability and reproduction. When the fitness benefits and costs of producing wings differ between males and females, sex-specific trade-offs can result in sex differences in the frequency of long-winged individuals. Furthermore, the social environment during development affects sex differences in wing development, but few empirical tests of this phenomenon have been performed to date. Here, I used the wing-dimorphic water strider Tenagogerris euphrosyne to test how rearing density and sex ratio affect the sex-specific development of long-winged dispersing morphs (i.e., sex-specific macroptery). I also used a full-sib, split-family breeding design to assess genetic effects on density-dependent, sex-specific macroptery. I reared water strider nymphs at either high or low densities and measured their wing development. I found that long-winged morphs developed more frequently in males than in females when individuals were reared in a high-density environment. However, the frequency of long-winged morphs was not biased according to sex when individuals were reared in a low-density environment. In addition, full-sib males and females showed similar macroptery incidence rates at low nymphal density, whereas the macroptery incidence rates differed between full-sib males and females at high nymphal density. Thus complex gene-by-environment-by-sex interactions may explain the density-specific levels of sex bias in macroptery, although this interpretation should be treated with some caution. Overall, my study provides empirical evidence for density-specific, sex-biased wing development. My findings suggest that social factors as well as abiotic factors can be important in determining sex-biased wing development in insects.

Lack of conspicuous sex-biased dispersal patterns at different spatial scales in an Asian endemic goose species breeding in unpredictable steppe wetlands.

Dispersal affects the spatial distribution and population structure of species. Dispersal is often male-biased in mammals while female-biased in birds, with the notable exception of the Anatidae. In this study, we tested genetic evidence for sex-biased dispersal (SBD) in the Swan Goose Anser cygnoides, an Asian endemic and IUCN vulnerable species, which has been increasingly restricted to breeding on Mongolian steppe wetlands. We analyzed the genotypes of 278 Swan Geese samples from 14 locations at 14 microsatellite loci. Results from assignment indices, analysis of molecular variance, and five other population descriptors all failed to support significant SBD signals for the Swan Goose at the landscape level. Although overall results showed significantly high relatedness within colonies (suggesting high levels of philopatry in both sexes), local male genetic structure at the 1,050 km distance indicated greater dispersal distance for females from the eastern sector of the breeding range. Hence, local dispersal is likely scale-dependent and female-biased within the eastern breeding range. These findings are intriguing considering the prevailing expectation for there to be female fidelity in most goose species. We suggest that while behavior-related traits may have facilitated the local genetic structure for the Swan Goose, several extrinsic factors, including the decreasing availability of the nesting sites and the severe fragmentation of breeding habitats, could have contributed to the absence of SBD at the landscape level. The long-distance molt migration that is typical of goose species such as the Swan Goose may also have hampered our ability to detect SBD. Hence, we urge further genetic sampling from other areas in summer to extend our results, complemented by field observations to confirm our DNA analysis conclusions about sex-specific dispersal patterns at different spatial scales in this species.

Dispersal evolution diminishes the negative density dependence in dispersal.

In many organisms, dispersal varies with the local population density. Such patterns of density-dependent dispersal (DDD) are expected to shape the dynamics, spatial spread, and invasiveness of populations. Despite their ecological importance, empirical evidence for the evolution of DDD patterns remains extremely scarce. This is especially relevant because rapid evolution of dispersal traits has now been empirically confirmed in several taxa. Changes in DDD of dispersing populations could help clarify not only the role of DDD in dispersal evolution, but also the possible pattern of subsequent range expansion. Here, we investigate the relationship between dispersal evolution and DDD using a long-term experimental evolution study on Drosophila melanogaster. We compared the DDD patterns of four dispersal-selected populations and their non-selected controls. The control populations showed negative DDD, which was stronger in females than in males. In contrast, the dispersal-selected populations showed DDD, where neither males nor females exhibited DDD. We compare our results with previous evolutionary predictions that focused largely on positive DDD, and highlight how the direction of evolutionary change depends on the initial DDD pattern of a population. Finally, we discuss the implications of DDD evolution for spatial ecology and evolution.

Dispersal and genetic structure in a tropical small mammal, the Bornean tree shrew (Tupaia longipes), in a fragmented landscape along the Kinabatangan River, Sabah, Malaysia.

BACKGROUND: Constraints in migratory capabilities, such as the disruption of gene flow and genetic connectivity caused by habitat fragmentation, are known to affect genetic diversity and the long-term persistence of populations. Although negative population trends due to ongoing forest loss are widespread, the consequence of habitat fragmentation on genetic diversity, gene flow and genetic structure has rarely been investigated in Bornean small mammals. To fill this gap in knowledge, we used nuclear and mitochondrial DNA markers to assess genetic diversity, gene flow and the genetic structure in the Bornean tree shrew, Tupaia longipes, that inhabits forest fragments of the Lower Kinabatangan Wildlife Sanctuary, Sabah. Furthermore, we used these markers to assess dispersal regimes in male and female T. longipes. RESULTS: In addition to the Kinabatangan River, a known barrier for dispersal in tree shrews, the heterogeneous landscape along the riverbanks affected the genetic structure in this species. Specifically, while in larger connected forest fragments along the northern riverbank genetic connectivity was relatively undisturbed, patterns of genetic differentiation and the distribution of mitochondrial haplotypes in a local scale indicated reduced migration on the strongly fragmented southern riverside. Especially, oil palm plantations seem to negatively affect dispersal in T. longipes. Clear sex-biased dispersal was not detected based on relatedness, assignment tests, and haplotype diversity. CONCLUSION: This study revealed the importance of landscape connectivity to maintain migration and gene flow between fragmented populations, and to ensure the long-term persistence of species in anthropogenically disturbed landscapes.