Landscape Genetics of American Beaver in Coastal Oregon

American beaver (Castor canadensis) have been translocated for population restoration, reduction of human-wildlife conflict, and enhancement of ecosystem function. Yet few studies have assessed dispersal of beaver, making it difficult to determine at what scale translocations are appropriate. Genetic studies can provide inferences about gene flow, and thus dispersal. We used a landscape genetic approach to evaluate whether landscape features influenced gene flow among beaver in the Coast Range of western Oregon, USA, using samples collected April–September 2014. We collected genetic samples from live-captured (n = 232), road-killed (n = 2) and trapper-provided (n = 58) tissue samples and genotyped them at 10 microsatellite loci. We mapped records of beaver translocations into or within the study area during the twentieth century to consider the effect of those movements on genetic structure. We used population assignment tests to delimit genetic clusters, evaluated correspondence of those clusters with watershed boundaries and translocation history, and then estimated differentiation between clusters and between watersheds using model-based and model-free approaches. We evaluated how individual genetic differences varied with geographic distance, and investigated related pairs within clusters. We developed landscape resistance models incorporating slope, distance to water, and watershed boundaries at 2 scales, and estimated effective distances between sample locations with least cost path and circuit theoretic analyses. We evaluated the correlation of individual genetic distances with effective distances using a pseudo-bootstrapping approach. Landscape genetic models did not explain spatial variation in genetic structure better than geographic distance, but hierarchical genetic structure corresponded with watershed boundaries and suggested influences from historical translocations. Pairwise individual genetic distances were positively correlated with geographic distances to 61 km; highly-related pairs mostly were detected <1 km apart (median = 1.0 km, = 14.6 ± 2.3 [SE] km, n = 77). We concluded that slope and distance to water did not strongly limit dispersal and gene flow by beaver in this system, but concordance of genetic structure with watershed boundaries suggests that dispersal is more common within than between watersheds. Genetic differentiation of beaver within this topographically complex system was much greater than reported in a study at similar spatial scales in relatively flat topography. We recommend that translocation efforts of American beaver in topographically complex landscapes occur within watersheds when possible but conclude that dispersal can occur across watersheds. © 2021 The Wildlife Society. This article is a U.S. Government work and is in the public domain in the USA.     

phylin: an r package for phylogeographic interpolation

phylin is a package for the r programming environment which offers different methods to spatially interpolate genetic information from phylogeographic data. These interpolations can be used to predict the spatial occurrence of different lineages within a phylogeny using a modified method of kriging, which allows the usage of a genetic distance matrix to derive a model of spatial dependence. phylin improves the available methods to generate interpolated surfaces from a phylogenetic trees by assessing the autocorrelation structure of the genetic information, interpolating the genetic data based on a statistical model, estimating the uncertainty of the predictions and identifying lineage occurrence and contact zones probability without projection of pairwise genetic distances into mid‐points between sample locations. The package also includes methods to plot interpolation surfaces and provide summary tables from the generated data and models. We provide an example of the usefulness of this tool by inferring the spatial occurrence of distinct historical evolutionary lineages of the Lataste's viper (Vipera latastei Boscá, 1878) in the Iberian Peninsula and identifying potential contact areas. The maps of phylogenetic patterns obtained with these methods provide a spatial context to test hypotheses related to processes underlying the geographic distribution of genetic diversity and to inform conservation planning.     

Disparate dispersal limitation in Geomalacus slugs unveiled by the shape and slope of the genetic–spatial distance relationship

Long-term dispersal ability is a key species’ trait constraining species ranges and thus large-scale biodiversity patterns. Here we infer the long-term dispersal abilities of three Geomalacus (Gastropoda, Pulmonata) species from their range-wide genetic–spatial distance relationships. This approach follows recent advances in statistical modelling of the analogous pattern at the community level: the distance decay in assemblage similarity. While linear relationships are expected for species with high long-term dispersal abilities, asymptotic relationships are expected for those with more restricted mobility. We evaluated three functional forms (linear, negative exponential and power-law) for the relationship between genetic distance (computed from mitochondrial cox1 sequences, n = 701) and spatial distance. Range fragmentation at present time and at the Last Glacial Maximum was also estimated based on the projection of climatic niches. The power-law function best fit the relationship between genetic and spatial distances, suggesting strong dispersal limitation and long-term population isolation in all three species. However, the differences in slope and explained variance pointed to disparities in dispersal ability among these weak dispersers. Phylogeographic patterns of Geomalacus species are thus largely driven by the same major process (i.e. dispersal limitation), operating at different strengths. This strong dispersal limitation results in geographic clustering of genetic diversity that makes these species highly vulnerable to genetic erosion due to climate change.     

A Comparative Statistical Analysis of Genetic Distances. I - Estimation of Distances and of their Variances: Distribution of the Estimators

We present a Monte-Carlo simulation analysis of the statistical properties of absolute genetic distance and of Nei's minimum and standard genetic distances. The estimation of distances (bias) and of their variances is analysed as well as the distributions of distance and variance estimators, taking into account both gamete and locus samplings. Both of Nei's statistics are non-linear when distances are small and consequently the distributions of their estimators are extremely asymmetrical. It is difficult to find theoretical laws that fit such asymmetrical distributions. Absolute genetic distance is linear and its distributions are better fit by a normal distribution. When distances are medium or large, minimum distance and absolute distance distributions are close to a normal distribution, but those of the standard distance can never be considered as normal. For large distances the jack-knife estimator of the standard distance variance is bad; another standard distance estimator is suggested. Absolute distance, which has the best mathematical properties, is particularly interesting for small distances if the gamete sample size is large, even when the number of loci is small. When both distance and gamete sample size are small, this statistic is biased.     

Decoupling of genetic and phenotypic divergence in a headwater landscape

In stream organisms, the landscape affecting intraspecific genetic and phenotypic divergence is comprised of two fundamental components: the stream network and terrestrial matrix. These components are known to differentially influence genetic structure in stream species, but to our knowledge, no study has compared their effects on genetic and phenotypic divergence. We examined how the stream network and terrestrial matrix affect genetic and phenotypic divergence in two stream salamanders, Gyrinophilus porphyriticus and Eurycea bislineata, in the Hubbard Brook Watershed, New Hampshire, USA. On the basis of previous findings and differences in adult terrestriality, we predicted that genetic divergence and phenotypic divergence in body morphology would be correlated in both species, but structured primarily by distance along the stream network in G. porphyriticus, and by overland distance in E. bislineata. Surprisingly, spatial patterns of genetic and phenotypic divergence were not strongly correlated. Genetic divergence, based on amplified DNA fragment length polymorphisms, increased with absolute geographic distance between sites. Phenotypic divergence was unrelated to absolute geographic distance, but related to relative stream vs. overland distances. In G. porphyriticus, phenotypic divergence was low when sites were close by stream distance alone and high when sites were close by overland distance alone. The opposite was true for E. bislineata. These results show that small differences in life history can produce large differences in patterns of intraspecific divergence, and the limitations of landscape genetic data for inferring phenotypic divergence. Our results also underscore the importance of explicitly comparing how terrestrial and aquatic conditions affect spatial patterns of divergence in species with biphasic life cycles.     

Effects of landscape and history on diversification of a montane, stream-breeding amphibian

Aim The aim of this study was to understand the roles of landscape features in shaping patterns of contemporary and historical genetic diversification among populations of the Andean tree frog (Hypsiboas andinus) across spatial scales. Location Andes mountains, north‐western Argentina, South America. Methods Mitochondrial DNA control region sequences were utilized to assess genetic differentiation among populations and calculate population pair‐wise genetic distances. Three models of movement, namely traditional straight‐line distance and two effective distances based on habitat classification, were examined to determine which of these explained the most variation in pair‐wise population genetic differentiation. The two habitat classifications were based on digital vegetation and hydrology layers that were generated from a 90‐m resolution digital elevation model (DEM) and known relationships between elevation and habitat. Mantel tests were conducted to test for correlations between geographic and genetic distance matrices and to estimate the percentage variation explained by each type of geographic distance. To investigate the location of possible barriers to gene flow, we used Monmonier’s maximum difference algorithm as implemented in barrier 2.2. Results At both geographic scales, effective distances explained more variation in genetic differentiation than did straight‐line distance. The least‐cost distances based on the simple classification performed better than the more detailed habitat classification. We controlled for the effects of historical range fragmentation determined from previous nested clade analyses, and therefore evaluated the effect of different distances on the genetic variation attributable to more recent factors. Effective distances identified populations that were highly divergent as a result of isolation in unsuitable habitats. The proposed locations of barriers to gene flow identified using Monmonier’s maximum difference algorithm corresponded well with earlier analyses and supported findings from our partial Mantel tests. Main conclusions Our results indicate that landscape features have been important in both historical and contemporary genetic structuring of populations of H. andinus at both large and small spatial scales. A landscape genetic perspective offers novel insights not provided by traditional phylogeographic studies: (1) effective distances can better explain patterns of differentiation in populations, especially in heterogeneous landscapes where barriers to dispersal may be common; and (2) least‐cost path analysis can help to identify corridors of movement between populations that are biologically more realistic.     

Isolation by distance explains genetic structure of Buggy Creek virus,a bird-associated arbovirus

Many of the arthropod-borne viruses (arboviruses) show extensive genetic variability and are widely distributed over large geographic areas. Understanding how virus genetic structure varies in space may yield insight into how these pathogens are adapted to and dispersed by different hosts or vectors, the relative importance of mutation, drift, or selection in generating genetic variability, and where and when epidemics or epizootics are most likely to occur. However, because most arboviruses tend to be sampled opportunistically and often cannot be isolated in large numbers at a given locale, surprisingly little is known about their spatial genetic structure on the local scale at which host/vector/virus interactions typically occur. Here, we examine fine-scale spatial structure of two sympatric lineages of Buggy Creek virus (BCRV, Togaviridae), an alphavirus transmitted by the ectoparasitic swallow bug (Oeciacus vicarius) to colonially nesting cliff swallows (Petrochelidon pyrrhonota) and invasive house sparrows (Passer domesticus) in North America. Data from 377 BCRV isolates at cliff swallow colony sites in western Nebraska showed that both virus lineages were geographically structured. Most haplotypes were detected at a single colony or were shared among nearby colonies, and pair-wise genetic distance increased significantly with geographic distance between colony sites. Genetic structure of both lineages is consistent with isolation by distance. Sites with the most genetically distinct BCRV isolates were occupied by large numbers of house sparrows, suggesting that concentrations of invasive sparrows may represent foci for evolutionary change in BCRV. Our results show that bird-associated arboviruses can show genetic substructure over short geographic distances.     

Ecological connectivity assessment in a strongly structured fire salamander (Salamandra salamandra) population

Small populations are more prone to extinction if the dispersal among them is not adequately maintained by ecological connections. The degree of isolation between populations could be evaluated measuring their genetic distance, which depends on the respective geographic (isolation by distance, IBD) and/or ecological (isolation by resistance, IBR) distances. The aim of this study was to assess the ecological connectivity of fire salamander Salamandra salamandra populations by means of a landscape genetic approach. The species lives in broad‐leaved forest ecosystems and is particularly affected by fragmentation due to its habitat selectivity and low dispersal capability. We analyzed 477 biological samples collected in 47 sampling locations (SLs) in the mainly continuous populations of the Prealpine and Eastern foothill lowland (PEF) and 10 SLs in the fragmented populations of the Western foothill (WF) lowland of Lombardy (northern Italy). Pairwise genetic distances (Chord distance, D_C) were estimated from allele frequencies of 16 microsatellites loci. Ecological distances were calculated using one of the most promising methodology in landscape genetics studies, the circuit theory, applied to habitat suitability maps. We realized two habitat suitability models: one without barriers (EcoD) and a second one accounting for the possible barrier effect of main roads (EcoD_b). Mantel tests between distance matrices highlighted how the Log‐D_C in PEF populations was related to log‐transformed geographic distance (confirming a prevalence of IBD), while it was explained by the Log‐EcoD, and particularly by the Log‐EcoD_b, in WF populations, even when accounting for the confounding effect of geographic distance (highlighting a prevalence of IBR). Moreover, we also demonstrated how considering the overall population, the effect of Euclidean or ecological distances on genetic distances acting at the level of a single group (PEF or WF populations) could not be detected, when population are strongly structured.     

Relative importance of habitat connectivity in shaping the genetic profiles of two southern African elephant‐shrews

Aim When interpreting genetic patterns across a landscape it is surprisingly difficult to disentangle the effects of landscape connectivity from those of species biology. Here, the spatial distributions of genetic variation of two sympatric elephant‐shrew species, the western rock elephant‐shrew (Elephantulus rupestris) and the round‐eared elephant‐shrew (Macroscelides proboscideus), are determined and compared. We selected these species because they have similar biologies but differ markedly in habitat use, the rationale being that differences in their genetic structure should be a result largely of landscape variables directly or indirectly affecting dispersal rather than of the biology of the species. Location South Africa and Namibia. Methods Mitochondrial sequence data (control region and cytochrome b) were used to describe the phylogeographic structure of these elephant‐shrew species across their distribution. To determine whether genetic variation is significantly structured, spatial analyses of molecular variation were performed. Isolation‐by‐distance versus alternative patterns of genetic structure was investigated using a Mantel test. Results Our analyses indicated an overall structured genetic profile for E. rupestris, a species closely associated with rocky outcrops. This was in contrast to a pattern mostly of isolation‐by‐distance across the distribution of M. proboscideus, a species found on gravel plains. Main conclusions Specific landscape features will differentially affect gene flow (both historical and current), and therefore also the spatial genetic structure, of species with markedly different habitat requirements. The genetic profiles for the two species included here support predictions based on the connectivity of their respective occupied habitats. The results also support the more general prediction that species with a naturally clustered distribution (such as E. rupestris) should have a more structured genetic pattern than those having a more continuous distribution (M. proboscideus).     

Landscape genetics reveals unique and shared effects of urbanization for two sympatric pool‐breeding amphibians

Metapopulation‐structured species can be negatively affected when landscape fragmentation impairs connectivity. We investigated the effects of urbanization on genetic diversity and gene flow for two sympatric amphibian species, spotted salamanders (Ambystoma maculatum) and wood frogs (Lithobates sylvaticus), across a large (>35,000 km²) landscape in Maine, USA, containing numerous natural and anthropogenic gradients. Isolation‐by‐distance (IBD) patterns differed between the species. Spotted salamanders showed a linear and relatively high variance relationship between genetic and geographic distances (r = .057, p < .001), whereas wood frogs exhibited a strongly nonlinear and lower variance relationship (r = 0.429, p < .001). Scale dependence analysis of IBD found gene flow has its most predictable influence (strongest IBD correlations) at distances up to 9 km for spotted salamanders and up to 6 km for wood frogs. Estimated effective migration surfaces revealed contrasting patterns of high and low genetic diversity and gene flow between the two species. Population isolation, quantified as the mean IBD residuals for each population, was associated with local urbanization and less genetic diversity in both species. The influence of geographic proximity and urbanization on population connectivity was further supported by distance‐based redundancy analysis and multiple matrix regression with randomization. Resistance surface modeling found interpopulation connectivity to be influenced by developed land cover, light roads, interstates, and topography for both species, plus secondary roads and rivers for wood frogs. Our results highlight the influence of anthropogenic landscape features within the context of natural features and broad spatial genetic patterns, in turn supporting the premise that while urbanization significantly restricts interpopulation connectivity for wood frogs and spotted salamanders, specific landscape elements have unique effects on these two sympatric species.     

Clump spacing in a desert dwarf shrub community

A Monte Carlo method based on Ripley's K function-a cumulative function related to the number of plants encountered at different distances from other plants-is used to test the null hypothesis of random distribution of shrub clumps in a desert dwarf shrub community in Namaqualand, South Africa, where Psilocaulon arenosum is the dominant shrub. The method takes into account the apparent regularity of pattern caused by the finite size (up to 2 m diameter) of the clumps. It is shown that the clump centres are significantly aggregated (compared to random expectation) at distances on the order of 1 m. Such aggregation is expected, as a simple result of regeneration near to seed sources, if the time between catastrophic droughts is short in relation to the time required for development of a non-aggregated or regular pattern determined by moisture competition. No significant regulatiry was detected at distances of 3 m or less. One subplot showed regularity above 3 m, but this pattern was not shown by the other subplot and may not be a competition effect. These results support a hypothesis of aggregation caused by regeneration pattern decaying slowly toward randomness as larger individuals compete.We thank C. J. F. ter Braak for suggesting the method of spatial analysis; 1. Noy-Meir, C. J. F. ter Braak and two unnamed reviewers for helpful comments; the Netherlands Science Research Council (ZWO) for financial support to I. C. Prentice (grant B84-218); and Prof. D. von Willert for material support to M. J. A. Werger.     

Species delimitation and geography

Despite the importance of the geographical arrangement of populations for the inference of species boundaries, only a few approaches that integrate spatial information into species delimitation have thus far been developed. Persistent differentiation of sympatric groups of individuals is the best criterion for species status. Species delimitation becomes more prone to error if allopatric metapopulations are considered because it is often difficult to assess whether observed differences between allopatric metapopulations would be sufficient to prevent the fusion of these metapopulations upon contact. We propose a novel approach for testing the hypothesis that the multilocus genetic distances between individuals or populations belonging to two different candidate species are not larger than expected based on their geographical distances and the relationship of genetic and geographical distances within the candidate species. A rejection of this null hypothesis is an argument for classifying the two studied candidate species as distinct species. Case studies show that the proposed tests are suitable to distinguish between intra‐ and interspecific differentiation. The regression approach proposed here is more appropriate for testing species hypotheses with regard to isolation by distance than (partial) Mantel tests. Our tests assume a linear relationship between genetic and (transformed) geographical distances. This assumption can be compromised by a high genetic variability within populations as found in a case study with microsatellite markers.     

Trophy hunting and perceived risk in closed ecosystems: Flight behaviour of three gregarious African ungulates in a semi‐arid tropical savanna

Although being an important conservation tool in Africa, trophy hunting is known to influence risk perception in wildlife species, thus affecting the behaviour and fitness of most targeted species. We studied the effects of trophy hunting on the flight behaviour of impala (Aepyceros melampus), greater kudu (Tragelaphus strepsiceros) and sable (Hippotragus niger) in two closed ecosystems, Cawston Ranch (hunting area) and Stanley and Livingstone Private Game Reserve (tourist area), western Zimbabwe. Using standardized field procedures, we assessed the flight behavioural responses of the three species in two seasons: non‐hunting (December–March) and hunting (April–November) between March 2013 and November 2014. We tested the effect of habitat, group size, sex, season, start distance and alert distance on flight initiation distance using linear mixed models. Habitat, group size sex and alert distance did not have any effect on flight initiation distance for the three species. The three species were more alert and displayed longer flight initiation distances in the hunting area compared with the tourist area. Flight initiation distances for the three species were higher during the hunting season for the hunting area and low during the non‐hunting season. Flight distances of the three species did not differ between the hunting area and the tourist area. We concluded that trophy hunting increased perceived risk of wild ungulates in closed hunting areas, whereas ungulates in non‐hunting areas are less responsive and somehow habituated to human presence. Management plans should include minimum approach distances by tourists as well as establishing seasonal restrictions on special zones to promote species viability. Research aimed at integrating behavioural responses with physiological aspects of target species should be promoted to ensure that managers are able to deal with the behavioural trade‐offs of trophy hunting at local and regional scale.     

GENETIC CONSEQUENCES OF SEED DISPERSAL IN THREE SYMPATRIC FOREST HERBS. II. MICROSPATIAL GENETIC STRUCTURE WITHIN POPULATIONS

Significant spatial genetic differentiation over short distances was detected by F-statistics and spatial autocorrelation within populations of the temperate forest herbs Cryptotaenia canadensis, Osmorhiza claytonii and Sanicula odorata (Apiaceae). Differences among the three species were consistent with estimates of their seed-dispersal abilities. Populations of Cryptotaenia, with the most limited seed dispersal, are characterized by genetic structure at smaller spatial scales than those of Osmorhiza or Sanicula, as indicated by higher estimates of θ(F_st), larger autocorrelation coefficients, and correlograms with more distant x-intercepts. Although spatial autocorrelation was somewhat more sensitive to the distribution of rare alleles than F-statistics, the two methods were generally concordant. Genetic structure was more pronounced, and inbreeding coefficients larger, in low-density, patchy populations than in a high-density site. Observed patterns of spatial autocorrelation, particularly for Cryptotaenia, were in agreement with expectations based on simulations of isolation by distance. The magnitude of observed autocorrelations was less than those typically produced in computer-simulation studies, but this discrepancy between empirical and theoretical results probably is derived from a lack of genetic and demographic equilibrium in natural populations. Isolation by distance can be an important evolutionary force organizing spatial genetic structure in plant populations, particularly in predominantly self-fertilizing species such as those studied here.     

Dispersal ability and habitat requirements determine landscape‐level genetic patterns in desert aquatic insects

Species occupying the same geographic range can exhibit remarkably different population structures across the landscape, ranging from highly diversified to panmictic. Given limitations on collecting population‐level data for large numbers of species, ecologists seek to identify proximate organismal traits—such as dispersal ability, habitat preference and life history—that are strong predictors of realized population structure. We examined how dispersal ability and habitat structure affect the regional balance of gene flow and genetic drift within three aquatic insects that represent the range of dispersal abilities and habitat requirements observed in desert stream insect communities. For each species, we tested for linear relationships between genetic distances and geographic distances using Euclidean and landscape‐based metrics of resistance. We found that the moderate‐disperser Mesocapnia arizonensis (Plecoptera: Capniidae) has a strong isolation‐by‐distance pattern, suggesting migration–drift equilibrium. By contrast, population structure in the flightless Abedus herberti (Hemiptera: Belostomatidae) is influenced by genetic drift, while gene flow is the dominant force in the strong‐flying Boreonectes aequinoctialis (Coleoptera: Dytiscidae). The best‐fitting landscape model for M. arizonensis was based on Euclidean distance. Analyses also identified a strong spatial scale‐dependence, where landscape genetic methods only performed well for species that were intermediate in dispersal ability. Our results highlight the fact that when either gene flow or genetic drift dominates in shaping population structure, no detectable relationship between genetic and geographic distances is expected at certain spatial scales. This study provides insight into how gene flow and drift interact at the regional scale for these insects as well as the organisms that share similar habitats and dispersal abilities.     

Changes of effective gene dispersal distances by pollen and seeds across successive life stages in a tropical tree

Pollen and seed dispersal are the two key processes in which plant genes move in space, mostly mediated by animal dispersal vectors in tropical forests. Due to the movement patterns of pollinators and seed dispersers and subsequent complex spatial patterns in the mortality of offspring, we have little knowledge of how pollinators and seed dispersers affect effective gene dispersal distances across successive recruitment stages. Using six highly polymorphic microsatellite loci and parentage analyses, we quantified pollen dispersal, seed dispersal, and effective paternal and maternal gene dispersal distances from pollen‐ and seed‐donors to offspring across four recruitment stages within a population of the monoecious tropical tree Prunus africana in western Kenya. In general, pollen‐dispersal and paternal gene dispersal distances were much longer than seed‐dispersal and maternal gene dispersal distances, with the long‐distance within‐population gene dispersal in P. africana being mostly mediated by pollinators. Seed dispersal, paternal and maternal gene dispersal distances increased significantly across recruitment stages, suggesting strong density‐ and distance‐dependent mortality near the parent trees. Pollen dispersal distances also varied significantly, but inconsistently across recruitment stages. The mean dispersal distance was initially much (23‐fold) farther for pollen than for seeds, yet the pollen‐to‐seed dispersal distance ratio diminished by an order of magnitude at later stages as maternal gene dispersal distances disproportionately increased. Our study elucidates the relative changes in the contribution of the two processes, pollen and seed dispersal, to effective gene dispersal across recruitment. Overall, complex sequential processes during recruitment contribute to the genetic make‐up of tree populations. This highlights the importance of a multistage perspective for a comprehensive understanding of the impact of animal‐mediated pollen and seed dispersal on small‐scale spatial genetic patterns of long‐lived tree species.     

Estimating dispersal distributions at multiple scales: within‐colony and among‐colony dispersal rates,distances and directions in European Shags Phalacrocorax aristotelis

Knowledge of the rate, distance and direction of dispersal within and among breeding areas is required to understand and predict demographic and genetic connectivity and resulting population and evolutionary dynamics. However dispersal rates, and the full distributions of dispersal distances and directions, are rarely comprehensively estimated across all spatial scales relevant to wild populations. We used re‐sightings of European Shags Phalacrocorax aristotelis colour‐ringed as chicks on the Isle of May (IoM), UK, to quantify rates, distances and directions of dispersal from natal to subsequent breeding sites both within IoM (within‐colony dispersal) and across 27 other breeding colonies covering 1045 km of coastline (among‐colony dispersal). Additionally, we used non‐breeding season surveys covering 895 km of coastline to estimate breeding season detection probability and hence potential bias in estimated dispersal parameters. Within IoM, 99.6% of individuals dispersed between their natal and observed breeding nest‐site. The distribution of within‐colony dispersal distances was right‐skewed; mean distance was shorter than expected given random settlement within IoM, yet some individuals dispersed long distances within the colony. The distribution of within‐colony dispersal directions was non‐uniform but did not differ from expectation given the spatial arrangement of nest‐sites. However, 10% of all 460 colour‐ringed adults that were located breeding had dispersed to a different colony. The maximum observed dispersal distance (170 km) was much smaller than the maximum distance surveyed (690 km). The distribution of among‐colony dispersal distances was again right‐skewed. Among‐colony dispersal was directional, and differed from random expectation and from the distribution of within‐colony dispersal directions. Non‐breeding season surveys suggested that the probability of detecting a colour‐ringed adult at its breeding location was high in the northeastern UK (98%). Estimated dispersal rates and distributions were therefore robust to incomplete detection. Overall, these data demonstrate skewed and directionally divergent dispersal distributions across small (within‐colony) and large (among‐colony) scales, indicating that dispersal could create genetic and demographic connectivity within the study area.     

Linking seed dispersal and genetic structure of trees: a biogeographical approach

Aim Natural and human‐induced differences in frugivore assemblages can influence the seed dispersal distances of trees. An important issue in seed dispersal systems is to understand whether differences in seed dispersal distances also affect the genetic structure of mature trees. One possible approach to test for a relationship between seed dispersal and the genetic structure of mature trees is to compare the genetic structure of two closely related tree species between two biogeographical regions that differ in frugivore assemblages and seed dispersal distances. Previous studies on two Commiphora species revealed that Commiphora guillauminii in Madagascar has a much lower seed dispersal distance than Commiphora harveyi in South Africa. We tested whether the lower seed dispersal distance might have caused decreased gene flow, resulting in a stronger genetic structure in Madagascar than in South Africa. Location Madagascar and South Africa. Methods Using amplified fragment length polymorphism markers we investigated the genetic structure of 134 trees in Madagascar and 158 trees in South Africa at a local and a regional spatial scale. Results In concordance with our hypothesis, kinship analysis suggests that gene flow was restricted mostly to 3 km in Madagascar and to 30 km in South Africa. At the local spatial scale, the genetic differentiation among groups of trees within sample sites was marginally significantly higher in Madagascar (F_ST = 0.069) than in South Africa (F_ST = 0.021). However, at a regional spatial scale genetic differentiation was lower in Madagascar (F_ST = 0.053) than in South Africa (F_ST = 0.163). Main conclusions Our results show that lower seed dispersal distances of trees were linked to higher genetic differentiation of trees only at a local spatial scale. This suggests that seed dispersal affects the genetic population structure of trees at a local, but not at a regional, spatial scale.     

Long‐distance dispersal in a fire‐ and livestock‐protected savanna

Savannas are highly diverse and dynamic environments that can shift to forest formations due to protection policies. Long‐distance dispersal may shape the genetic structure of these new closed forest formations. We analyzed eight microsatellite loci using a single‐time approach to understand contemporary pollen and effective seed dispersal of the tropical tree, Copaifera langsdorffii Desf. (Fabaceae), occurring in a Brazilian fire‐ and livestock‐protected savanna. We sampled all adult trees found within a 10.24 ha permanent plot, young trees within a subplot of 1.44 ha and open‐pollinated seeds. We detected a very high level of genetic diversity among the three generations in the studied plot. Parentage analysis revealed high pollen immigration rate (0.64) and a mean contemporary pollen dispersal distance of 74 m. In addition, half‐sib production was 1.8 times higher than full‐sibs in significant higher distances, indicating foraging activity preference for different trees at long distances. There was a significant and negative correlation between diameter at breast height (DBH) of the pollen donor with the number of seeds (r = ?0.640, P‐value = 0.032), suggesting that pollen donor trees with a higher DBH produce less seeds. The mean distance of realized seed dispersal (recruitment kernel) was 135 m due to the large home range dispersers (birds and mammals) in the area. The small magnitude of spatial genetic structure found in young trees may be a consequence of overlapping seed shadows and increased tree density. Our results show the positive side of closed canopy expansion, where animal activities regarding pollination and seed dispersal are extremely high.     

High gene flow between populations of Macrotermes michaelseni (Isoptera, Termitidae)

Termite alates are thought to be poor active flyers, and this should lead to considerable genetic differentiation on small spatial scales. However, using four microsatellite loci for the termite Macrotermes michaelseni we found low values of genetic differentiation (F_ST) across a spatial scale of even more than 50 km. Genetic differentiation between populations increased with spatial distance up to 50 km. Furthermore, up to this distance, the scatter around the linear regression of genetic differentiation versus spatial distance increased with spatial distance. This suggests that across such spatial distances gene flow and genetic drift are of about equal importance, and near equilibrium. Using a regional F_ST as well as the distance between populations with non-significant F_ST-values (up to 25 km), gene flow is sufficiently high so that populations may be regarded as panmictic on spatial scales of 25 to 50 km. The apparent contradiction between dispersal distances observed in the field and estimates of gene flow from genetic markers may be due to the masses of swarming alates. Assuming a leptokurtic distribution of dispersal distances, atleast some alates are expected to travel considerable distances, most likely by passive drift. Received 25 January 2005; revised 11 April 2005; accepted 26 April 2005.