Island isolation and habitat heterogeneity correlate with DNA variation in a marine snail (Littorina saxatilis)

The null assumption of molecular variation is that most of it is neutral to natural selection. This is in contrast to variation in morphological traits that we generally assume is maintained by selection, and therefore often by selection coupled to environmental heterogeneity in time and space. Examples of molecular variation that vary over habitat-shifts, particularly in allozymes, show that the relative impact of non-neutral variation as compared to neutral variation might be substantial in some systems. To assess the importance of habitat-generated variation in relation to variation generated by random processes in nuclear DNA markers at small spatial scales, we compared the effects of island isolation and habitat heterogeneity on genetic substructuring in a rocky shore snail ( Littorina saxatilis ). This species has a restricted migration among islands owing to the lack of free-floating larvae. Earlier studies show that allozymes vary extensively as a consequence of isolation by water barriers among islands, but also as a consequence of divergent selection among different microhabitats within islands. In the DNA markers we observed genetic differentiation owing to island isolation at three of nine loci. In addition, variation at three loci correlated with habitat type, but the correlation for two of the loci was weak. Overall, isolation contributed slightly more to the genetic variation among populations than did habitat-related factors but the difference was small. It is concluded that both island isolation, which interrupts gene flow, and a heterogeneous habitat cause genetic substructuring at the DNA level in L. saxatilis in the studied area, and thus in this species we need to be somewhat concerned about habitat heterogeneity also at DNA loci.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 377–384.     

Genetic consequences of natal dispersal in the colonial lesser kestrel

Dispersal is a life-history trait that plays a fundamental role in population dynamics, influencing evolution, species distribution, and the genetics and structure of populations. In spite of the fact that dispersal has been hypothesized to be an efficient behavioural mechanism to avoid inbreeding, the expected relationship between dispersal and mate relatedness still remains controversial. Here, we examine the genetic consequences of natal dispersal, namely the higher chance of obtaining genetically less similar mates as a result of moving from natal to breeding sites, in a lesser kestrel (Falco naumanni) population. Relatedness between individuals tended to decrease with distance between their breeding colonies, indicating that the study population follows an 'isolation-by-distance' pattern of spatial genetic structure. Such a fine-scale genetic structure generates a scenario in which individuals can potentially increase the chance of obtaining genetically less similar mates by dispersing over larger distances from their natal colony. Using dispersal information and genotypic data, we showed that mate relatedness decreased with natal dispersal distance, an effect that remained significant both while including and excluding philopatric individuals from the data set. These results, together with the well known detrimental consequences of reduced genetic diversity in the study population, suggest that dispersal may have evolved, at least in part, to avoid the negative fitness consequences of mating with genetically similar individuals.     

Genetic differentiation between populations of the European rose hip fly Rhagoletis alternata

We tested for genetic differentiation between populations of Rhagoletis alternata Fall. (Diptera: Tephritidae) on three different host species. We collected larvae from three rose species of the section Caninae ( Rosa canina L., Rosa corymbifera Borkh . , and Rosa rubiginosa L.) from 15 sites across Germany, where the three roses occurred together. Additionally, we sampled three sites in Switzerland. Roses differ in morphology (e.g. leaf glands) as well as phenology. We were able to score nine allozyme loci (five polymorphic). Populations from the three hosts did not differ in genetic variability. We found significant genetic differentiation between populations from different host species. However, the differentiation was very low (0.9%). Hence, we found no indication for host races. Furthermore, surprisingly little geographical structure of genetic differentiation was found between populations of this fruit fly across central Europe. We offer three mutually non-exclusive explanations for these findings. First, gene flow between populations of Rh. alternata is high. Second, the pattern of genetic differentiation is based on a recent expansion of the distributional range . Third, the ongoing gene flow between roses of the section Caninae acts as a hybrid bridge.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 619–625.     

Fine-scale spatial genetic structure and gene dispersal in Silene latifolia

Plants are sessile organisms, often characterized by limited dispersal. Seeds and pollen are the critical stages for gene flow. Here we investigate spatial genetic structure, gene dispersal and the relative contribution of pollen vs seed in the movement of genes in a stable metapopulation of the white campion Silene latifolia within its native range. This short-lived perennial plant is dioecious, has gravity-dispersed seeds and moth-mediated pollination. Direct measures of pollen dispersal suggested that large populations receive more pollen than small isolated populations and that most gene flow occurs within tens of meters. However, these studies were performed in the newly colonized range (North America) where the specialist pollinator is absent. In the native range (Europe), gene dispersal could fall on a different spatial scale. We genotyped 258 individuals from large and small (15) subpopulations along a 60 km, elongated metapopulation in Europe using six highly variable microsatellite markers, two X-linked and four autosomal. We found substantial genetic differentiation among subpopulations (global F_ST=0.11) and a general pattern of isolation by distance over the whole sampled area. Spatial autocorrelation revealed high relatedness among neighboring individuals over hundreds of meters. Estimates of gene dispersal revealed gene flow at the scale of tens of meters (5–30 m), similar to the newly colonized range. Contrary to expectations, estimates of dispersal based on X and autosomal markers showed very similar ranges, suggesting similar levels of pollen and seed dispersal. This may be explained by stochastic events of extensive seed dispersal in this area and limited pollen dispersal.     

Genetic structure of brown trout, salmo trutta l., at the southern limit of the distribution range of the anadromous form

Genetic variation at 33 protein loci was investigated in 41 wild brown trout populations from four river basins in Galicia (northwest Spain) to analyse the amount and distribution of genetic diversity in a marginal area, located in the distribution limit of the anadromous form of this species. The genetic diversity detected within populations (H between 0 and 6%) lies within the range quoted for this species in previous reports. The Mino, the most southern river basin analysed, showed a significantly lower genetic diversity and the highest genetic differentiation among the river basins studied. The hierarchical gene diversity analysis showed high population differentiation in a restricted area (GST = 27%), mostly due to differences among populations within basins (GSC = 22%). The reduction of GST observed when the isolated samples were excluded from the analysis (GST = 17%) showed the importance of habitat fragmentation on the heterogeneity detected. Gene flow among populations was comparatively evaluated by three indirect methods, which in general revealed low figures of absolute number of migrants per generation, slightly higher than 1. The gene flow among basins reflected a positive relationship with geographical distance. This trend was confirmed by the significant correlation observed between geographical and genetic distances, including all population pairs, which suggests a component of isolation by distance in brown trout genetic structure. Nevertheless, the nonsignificant intrabasin correlation demonstrates the complexity of genetic relationships among populations in this species. The model of genetic structure in brown trout is discussed in the light of the results obtained.     

Genetic differentiation between individuals

I describe a method of analysis of genetic differentiation which is suitable for the comparison of genetic and demographic estimates of the ‘neighborhood size’– more precisely the product of density and second moment of dispersal distance σ²– in continuous populations sampled at the smallest scale. This method is based on results of models of isolation by distance common to a wide variety of dispersal distances. The performance of this method is tested by simulation for some highly leptokurtic dispersal distributions, and it is applied to a previous study of a kangaroo‐rat (Dipodomys spectabilis) population. In this case, genetic and demographic estimates are within a factor of two from each other. Thus, in line with some previous examples, this study shows that a better agreement may be attained than is usually recognized between genetic and demographic estimates.     

Fine-scale genetic structure and dispersal in cooperatively breeding apostlebirds

In cooperatively breeding species, restricted dispersal of offspring leads to clustering of closely related individuals, increasing the potential both for indirect genetic benefits and inbreeding costs. In apostlebirds (Struthidea cinerea), philopatry by both sexes results in the formation of large (up to 17 birds), predominantly sedentary breeding groups that remain stable throughout the year. We examined patterns of relatedness and fine-scale genetic structure within a population of apostlebirds using six polymorphic microsatellite loci. We found evidence of fine-scale genetic structure within the study population that is consistent with behavioural observations of short-distance dispersal, natal philopatry by both sexes and restricted movement of breeding groups between seasons. Global F(ST) values among breeding groups were significantly positive, and the average level of pairwise relatedness was significantly higher for individuals within groups than between groups. For individuals from different breeding groups, geographical distance was negatively correlated with pairwise relatedness and positively correlated with pairwise F(ST). However, when each sex was examined separately, this pattern was significant only among males, suggesting that females may disperse over longer distances. We discuss the potential for kin selection to influence the evolution and maintenance of cooperative breeding in apostlebirds. Our results demonstrate that spatial genetic structural analysis offers a useful alternative to field observations in examining dispersal patterns of cooperative breeders.     

Partitioning nuclear and chloroplast variation at multiple spatial scales in the neotropical epiphytic orchid, Laelia rubescens

Insights into processes that lead to the distribution of genetic variation within plant species require recognition of the importance of both pollen and seed movement. Here we investigate the contributions of pollen and seed movement to overall gene flow in the Central American epiphytic orchid, Laelia rubescens. Genetic diversity and structure were examined at multiple spatial scales in the tropical dry forest of Costa Rica using nuclear (allozymes) and chloroplast restriction fragment length polymorphism (RFLP) markers, which were found to be diverse (allozymes, P = 73.3%; HE = 0.174; cpDNA, HE = 0.741). Nuclear genetic structure (FSTn) was low at every spatial scale (0.005-0.091). Chloroplast markers displayed more structure (0.073-0.254) but relatively similar patterns. Neither genome displayed significant isolation-by-distance. Pollen and seed dispersal rates did not differ significantly from one another (mp/ms = 1.40) at the broadest geographical scale, among sites throughout Costa Rica. However, relative contributions of pollen and seeds to gene flow were scale-dependent, with different mechanisms determining the dominant mode of gene flow at different spatial scales. Much seed dispersal is highly localized within the maternal population, while some seeds enter the air column and are dispersed over considerable distances. At the intermediate scale (10s to 100s of metres) pollinators are responsible for substantial pollen flow. This species appears capable of distributing its genes across the anthropogenically altered landscape that now characterizes its Costa Rican dry forest habitat.     

Drivers of genetic differentiation in a generalist insect‐pollinated herb across spatial scales

The isolation‐by‐distance model (IBD) predicts that genetic differentiation among populations increases with geographic distance. Yet, empirical studies show that a variety of ecological, topographic and historical factors may override the effect of geographic distance on genetic variation. This may particularly apply to species with narrow but highly heterogeneous distribution ranges, such as those occurring along elevational gradients. Using nine SSR markers, we study the genetic differentiation of the montane pollination‐generalist herb, Erysimum mediohispanicum. Because the effects of any given factor may depend on the geographic scale considered, we investigate the contribution of different environmental and historical factors at three different spatial scales. We evaluate five competing models that put forward the role of geographic distance, local environmental factors [biotic interactions (IBEb) and climatic variables (IBEa)], landscape resistance (IBR) and phylogeographic patterns (IBP), respectively. We find significant IBD regardless of the spatial scale and the genetic distance estimator considered. However, IBEa and IBP also play a prominent role in shaping genetic differentiation patterns at the larger spatial scales, and IBR is significant at the fine spatial scale. Overall, our results highlight the importance of combining different estimators, statistical approaches and spatial scales to disentangle the relative importance of the various ecological factors contributing to the shaping of genetic divergence patterns in natural populations.     

Genetic differentiation associated with commercial traffic in the Polynesian mosquito, Aedes polynesiensis Marks 1951

The population structure of the Polynesian mosquito Aedes polynesiensis was investigated using electrophoretic data from two polymorphic protein loci. Considerable differentiation was observed both within and between islands in different archipelagos (Society, Tuamotu, Austral). Gene flow evaluated by F_st estimates was independent of geographic distance between islands but related to commercial traffic intensity. The results are discussed in view of recent findings on the variability of susceptibility to insecticides and of suitability as a vector for the nematode Wuchereria bancrofti.     

Genetic differentiation and speciation among four Phyllotreta species (Coleoptera: Chrysomelidae)

Among the four closely related Phyllotreta species P. dilatata Thomson, 1866, P. flexuosa (Illiger, 1794), P. ochripes (Curtis, 1837) and P. tetrastigma (Comolli, 1837) several systematic problems occur due to the existence of colour pattern variation and morphological similarity. All four species feed on Brassicaceae, but P. tetrastigma , P. flexuosa and P. dilatata are monophagous in the field, whereas P. ochripes is oligophagous. Genetic differentiation and systematic relationships of the four species are studied by means of allozyme electrophoresis of 10 variable loci, with a total of 30 allelomorphs. Of the six alleles that are present only in a single species, none is diagnostic. Rogers modified genetic distance and Cavalli-Sforza & Edwards arc distance between all populations are calculated and Distance Wagner trees are constructed, revealing three major groups. P. ochripes is separated first, being the most distant species. The three other species are more closely related, and within this group P. dilatata is separated from the other two. The last group contains the populations of P. tetrastigma and P. flexuosa. In the differentiation between the different species host plant shifts and consequent genetic differentiation have been important.     

Genetic structure of the endangered perennial plant Eryngium alpinum (Apiaceae) in an alpine valley

We investigated the genetic structure of Eryngium alpinum (Apiaceae) in an Alpine valley where the plant occurs in patches of various sizes. In a conservation perspective, our goal was to determine whether the valley consists of one or several genetic units. Habitat fragmentation and previous observations of restricted pollen/seed dispersal suggested pronounced genetic structure, but gene dispersal often follows a leptokurtic distribution, which may lead to weak genetic structure. We used nine microsatellite loci and two nested sampling designs (50 × 50 m grid throughout the valley and 2 × 2 m grid in two 50 × 10 m quadrats). Within the overall valley, F -statistics and Bayesian approaches indicated high genetic homogeneity. This result might be explained by: (1) underestimation of long-distance pollen/seed dispersal by in situ experiments and (2) too recent fragmentation events to build up genetic structure. Spatial autocorrelation revealed isolation by distance on the overall valley but this pattern was much more pronounced in the 50 × 10 m quadrats sampled with a 2-m mesh. This was probably associated with limited primary seed dispersal, leading to the spatial clustering of half-sibs around maternal plants. We emphasize the interest of nested sampling designs and of combining several statistical tools.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 667–677.     

Genetic diversity and genetic structure of populations of Ranunculus japonicus Thunb. (Ranunculaceae)

Abstract In order to clarify the genetic diversity and population structure of Ranunculus japonicus , allozymic analysis was conducted on 60 populations in southwestern Japan. Considerable genetic variati ons were detected among the populations of R. japonicus . The genetic diversities within species ( H _es = 0.215) and within populations ( H _ep = 0.172) were slightly higher than those of other perennial herbs with widespread distribution and outcrossing plants. Significantly higher values of fixation index were detected in some populations, which might have arisen from restricted mating partners. The majority of genetic variation (approx. 80%) resided within a population and a moderate level of genetic differentiation ( G _ST = 0.203) was observed among populations. The F _ST value (0.203) suggests the existence of a substantial population structure in this species. The highly significant correlation between geographic distance and F _ST values indicates that isolation by distance has played an important role in the construction of the genetic structure of this species.     

Quantifying population structure on short timescales

Quantifying the contribution of the various processes that influence population genetic structure is important, but difficult. One of the reasons is that no single measure appropriately quantifies all aspects of genetic structure. An increasing number of studies is analysing population structure using the statistic D, which measures genetic differentiation, next to G_ST, which quantifies the standardized variance in allele frequencies among populations. Few studies have evaluated which statistic is most appropriate in particular situations. In this study, we evaluated which index is more suitable in quantifying postglacial divergence between three‐spined stickleback (Gasterosteus aculeatus) populations from Western Europe. Population structure on this short timescale (10 000 generations) is probably shaped by colonization history, followed by migration and drift. Using microsatellite markers and anticipating that D and G_ST might have different capacities to reveal these processes, we evaluated population structure at two levels: (i) between lowland and upland populations, aiming to infer historical processes; and (ii) among upland populations, aiming to quantify contemporary processes. In the first case, only D revealed clear clusters of populations, putatively indicative of population ancestry. In the second case, only G_ST was indicative for the balance between migration and drift. Simulations of colonization and subsequent divergence in a hierarchical stepping stone model confirmed this discrepancy, which becomes particularly strong for markers with moderate to high mutation rates. We conclude that on short timescales, and across strong clines in population size and connectivity, D is useful to infer colonization history, whereas G_ST is sensitive to more recent demographic events.     

Phylogeography of five Polytrichum species within Europe

Using allozymes and microsatellites we have analysed the genetic structure among European populations for several Polytrichum species to infer relevant factors, such as historical events or gene flow, that have shaped their genetic structure. As we observed low levels of genetic differentiation among populations, and no decreasing levels of genetic variation with increasing latitude within most of the examined species, no genetic evidence was obtained for a step-wise recolonization of Europe from southern refugia after the latest glacial period for P. commune , P. uliginosum , P. formosum and P. piliferum . The near absence of population substructuring within these species does indicate that extensive spore dispersal is the most important factor determining the genetic structure among European Polytrichum populations. Gene flow levels have apparently been sufficient to prevent genetic differentiation among populations caused by genetic drift, and to wipe out any genetic structure caused by the postglacial recolonization process. On the other hand, increased genetic differentiation of alpine P. formosum populations suggests that mountain ranges might restrict gene flow significantly among Polytrichum populations. In contrast to most examined Polytrichum species, P. juniperinum showed high levels of genetic differentiation and a profound genetic structure. Assuming that gene flow is not more restricted in P. juniperinum , these findings suggest that this species has recolonized Europe after the latest glacial period from two different refugia, one possibly being the British Isles.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society 2003, 78, 203–213.     

Small-scale spatial genetic structure in the Central African rainforest tree species Aucoumea klaineana: a stepwise approach to infer the impact of limited gene dispersal, population history and habitat fragmentation

Under the isolation-by-distance model, the strength of spatial genetic structure (SGS) depends on seed and pollen dispersal and genetic drift, which in turn depends on local demographic structure. SGS can also be influenced by historical events such as admixture of differentiated gene pools. We analysed the fine-scale SGS in six populations of a pioneer tree species endemic to Central Africa, Aucoumea klaineana. To infer the impacts of limited gene dispersal, population history and habitat fragmentation on isolation by distance, we followed a stepwise approach consisting of a Bayesian clustering method to detect differentiated gene pools followed by the analysis of kinship-distance curves. Interestingly, despite considerable variation in density, the five populations situated under continuous forest cover displayed very similar extent of SGS. This is likely due to an increase in dispersal distance with decreased tree density. Admixture between two gene pools was detected in one of these five populations creating a distinctive pattern of SGS. In the last population sampled in open habitat, the genetic diversity was in the same range as in the other populations despite a recent habitat fragmentation. This result may due to the increase of gene dispersal compensating the effect of the disturbance as suggested by the reduced extent of SGS estimated in this population. Thus, in A. klaineana, the balance between drift and dispersal may facilitate the maintenance of genetic diversity. Finally, from the strength of the SGS and population density, an indirect estimate of gene dispersal distances was obtained for one site: the quadratic mean parent-offspring distance, sigma(g), ranged between 210 m and 570 m.     

Genetic and morphological divergence reveals local subdivision of perch (Perca fluviatilis L.)

The level of gene flow is an important factor influencing genetic differentiation between populations. Typically, geographic distance is considered to be the major factor limiting dispersal and should thus only influence the degree of genetic divergence at larger spatial scales. However, recent studies have revealed the possibility for small-scale genetic differentiation, suggesting that the spatial scale considered is pivotal for finding patterns of isolation by distance. To address this question, genetic and morphological differentiation were studied at two spatial scales (range 2–13 km and range 300 m to 2 km) in the perch ( Perca fluviatilis L.) from the east coast archipelago of Sweden, using seven microsatellite loci and geometric morphometrics. We found highly significant genetic differentiation between sampled locations at both scales. At the larger spatial scale, the distance per se was not affecting the level of divergence. At the small scale, however, we found subtle patterns of isolation by distance. In addition, we also found morphological divergence between locations, congruent with a spatial separation at a microgeographic scale, most likely due to phenotypic plasticity. The present study highlights the importance of geographical scale and indicates that there might be a disparity between the dispersal capacity of a species and the actual movement of genes. Thus, how we view the environment and possible barriers to dispersal might have great implications for our ability to fully understand the evolution of genetic differentiation, local adaptation, and, in the end, speciation.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 746–758.