GENETIC STRUCTURE OF POPULATIONS OF GEOGRAPHICALLY RESTRICTED AND WIDESPREAD SPECIES OF ASTRAGALUS (FABACEAE)

Evolutionary theory predicts that species with small ranges and few individuals will exhibit low levels of genetic polymorphism. We investigated the population genetic structure of two locally endemic and two geographically widespread species of Astragalus. To facilitate direct comparisons among these congeners, three populations of each species were sampled in a consistent manner and scored for allozyme polymorphisms at 12 loci. Genetic polymorphism was lower in restricted A. linifolius and A. osterhouti than in widespread A. pectinatus. However, the restricted species do exhibit a moderate level of isozyme variation, comparable to that of widespread A. pattersoni and higher than has been detected in several other rare plant taxa. As measured by Wright's F statistics, there were no consistent differences between the restricted and widespread taxa with respect to the organization of genetic variation. F_ST values were low in all taxa, indicating little heterogeneity among populations.     

CORRELATION OF PAIRWISE GENETIC AND GEOGRAPHIC DISTANCE MEASURES: INFERRING THE RELATIVE INFLUENCES OF GENE FLOW AND DRIFT ON THE DISTRIBUTION OF GENETIC VARIABILITY

Attempts to relate estimates of regional F_ST to gene flow and drift via Wright's (1931) equation F_ST ≈ 1/ (4Nm + 1) are often inappropriate because most natural sets of populations probably are not at equilibrium (McCauley 1993), as assumed by the island model upon which the equation is based, or ineffective because the influences of gene flow and drift are confounded in the product Nm. Evaluations of the association between genetic (F_ST) and geographic distances separating all pairwise populations combinations in a region allows one to test for regional equilibrium, to evaluate the relative influences of gene flow and drift on population structure both within and between regions, and to visualize the behavior of the association across all degrees of geographic separation. Tests of the model using microsatellite data from 51 populations of eastern collared lizards (Crotaphytus collaris collaris) collected from four distinct geographical regions gave results highly consistent with predicted patterns of association based on regional differences in various historical and ecological factors that affect the amount of drift and gene flow. The model provides a prerequisite for and an alternative to regional F_ST analyses, which often simply assume regional equilibrium, thus potentially leading to erroneous and misleading inferences regarding regional population structure.     

DEMOGRAPHIC GENETICS OF A PIONEER TROPICAL TREE SPECIES: PATCH DYNAMICS,SEED DISPERSAL,AND SEED BANKS

We consider whether changes in population-genetic structure through the life cycle of Cecropia obtusifolia, a tropical pioneer tree, reflect its gap-dependent demography and the role of evolutionary processes that are important for this species. We asked whether the spatial scale at which population-genetic subdivision occurs corresponds to the scale of habitat patchiness created by gap dynamics; whether patterns of seed dispersal and storage in the soil affect spatial genetic patterns; and whether spatial genetic patterns change through the species life cycle. We estimated Wright's F-statistics for six successive life-history stages for individuals grouped into subpopulations according to occurrence in natural gaps, physical proximity, or occurrence within large quadrats. For each life stage, F_ST-statistics were significantly higher when individuals were grouped by gaps, although concordant patterns across life stages for the three grouping methods were obtained. This supports the hypothesis that patchy recruitment in gaps or among-gap heterogeneity influences the species' genetic structure. F-statistics of seeds collected from females before dispersal (tree seeds), seed-rain seeds, soil seeds, seedlings, juveniles, and adults grouped by gaps, were, respectively: F_IT = 0.004, 0.160, 0.121, 0.091, –0.0002, –0.081; F_IS = –0.032, 0.124, 0.118, 0.029, –0.016, –0.083; and F_ST = 0.035, 0.041, 0.003, 0.063, 0.015, 0.002. Spatial genetic differentiation in rain seeds was not significantly lower than that of tree seeds. The loss of genetic structure in the soil seed bank, relative to that found in the seed rain may be due to sampling artifacts, but alternative explanations, such as microsite selection or temporal Wahlund effect are also discussed. If structure among soil seeds is unbiased, the peak in seedling F_ST may be due to microsite selection. F_IS of seeds in the rain and soil were significantly greater than zero. A Wahlund effect is the most likely cause of these positive F_IS values. Such fine-scale substructuring could be caused by correlated seed deposition by frugivores. The decrease in F_IS from seedlings to adults could result from loss of fine-scale genetic structure during stand thinning or from selection.     

HIERARCHICAL GENETIC STRUCTURE AND GENE FLOW IN MACROGEOGRAPHIC POPULATIONS OF THE EASTERN TENT CATERPILLAR (MALACOSOMA AMERICANUM)

Genetic structure and inferred rates of gene flow in macrogeographic populations of the eastern tent caterpillar Malacosoma americanum were analyzed at two hierarchical scales: local demes and regional subpopulations. Wright's F-statistics were used to estimate population genetic structure using multilocus genotypic data generated electrophoretically. Estimated values of F_ST and the distribution of private alleles were then used to obtain indirect estimates of gene flow. We found modest, though significant, genetic structure at both spatial scales, a pattern consistent with high rates of gene flow over the large distances involved. Modest values obtained for Nei's genetic distance also suggested high levels of gene flow across the range of this species, although some gene-flow restriction resulting from isolation by distance was suggested by a positive regression of genetic distance on geographic distance. The observed homogeneity at enzyme loci across the range of M. americanum parallels the reported uniformity in morphology, suggesting a general absence of local genetic differentiation in this widely distributed species. The genetic homogeneity observed in this wide-ranging insect is discussed in terms of organism-specific environmental experience at different spatial scales. Some organisms occupying apparently heterogeneous environments may ameliorate unsuitable local conditions through microhabitat selection or behavioral modification of their microenvironment. This may be accomplished in M. americanum through group shelter construction and behavioral thermoregulation, closely tying thermoregulation to social biology in this species. If in this way the tent helps produce an effectively homogeneous environment for this species across its extensive range, this system may provide a unique example of how social behavior can influence the distribution of genetic variation in a population.     

MOLECULAR EVOLUTIONARY DIVERGENCE AMONG NORTH AMERICAN CAVE CRICKETS. I. ALLOZYME VARIATION

Forty-nine populations of nine species of North American cave crickets (genera Euhadenoecus and Hadenoecus) have been studied for genetic variation at 41 loci by electrophoresis. Wright's F_ST, Slatkin's Nm^* gene-flow estimator, and Nei's genetic distances (D) have been used to compare closely related species that have different ecological requirements (cave vs. forest species), distribution patterns, and/or different degrees of geographic isolation among populations. Cave and epigean (noncave) species differ greatly in their levels of genetic differentiation. Cave species have lower rates of gene exchange (low Nm, high D, and F_ST) than epigean species. Within cave species the degree of genetic differentiation among populations is correlated with the limestone structure of the area where the species occur. Species or groups of populations inhabiting areas where the limestone is continuous and highly fissured (e.g., H. subterraneus populations in the Mammoth Cave region) are genetically less differentiated than are populations occurring in regions where the limestone distribution is more fragmented, such as the Appalachian Ridge where E. fragilis occurs; this effect is more extreme in Central Tennessee where genetically differentiated E. insolitus populations occur only a few kilometers apart. This suggests that epigean dispersal through forest habitat in cave-dwelling species is negligable. For forest species, the data indicate relatively recent radiation with ongoing gene exchange among populations. For cave species, the distribution of protein polymorphisms is apparently more a function of historical patterns of gene exchange rather than current gene exchange. Phylogenetic relationships were studied using cluster analyses (UPGMA and Wagner algorithms) of Nei's and Edwards' genetic distances and multivariate analysis (correspondence analysis) of the raw allele frequencies. Different algorithms result in branching patterns that are similar but not entirely concordant with one another or with the phylogeny based on morphology.     

PHASE III OF WRIGHT'S SHIFTING BALANCE PROCESS AND THE VARIANCE AMONG DEMES IN MIGRATION RATE

Interdemic selection by the differential migration of individuals out from demes of high fitness and into demes of low fitness (Phase III) is one of the most controversial aspects of Wright's Shifting Balance Theory. I derive a relationship between Phase III migration and the interdemic selection differential, S, and show its potential effect on F_ST. The relationship reveals a diversifying effect of interdemic selection by Phase III migration on the genetic structure of a metapopulation. Using experimental metapopulations, I explored the effect of Phase III migration on F_ST by comparing the genetic variance among demes for two different patterns of migration: (1) island model migration and (2) Wright's Phase III migration. Although mean migration rates were the same, I found that the variance among demes in migration rate was significantly higher with Phase III than with island model migration. As a result, F_ST for the frequency of a neutral marker locus was higher with Phase III than it was with island model migration. By increasing F_ST, Phase III enhanced the genetic differentiation among demes for traits not subject to interdemic selection. This feature makes Wright's process different from individual selection which, by reducing effective population size, decreases the genetic variance within demes for all other traits. I discussed this finding in relation to the efficacy of Phase III and random migration for effecting peak shifts, and the contribution of genes with indirect effects to among‐deme variation.     

VOCAL DIALECTS AND THE STRUCTURE OF GEOGRAPHIC VARIATION IN MORPHOLOGICAL AND ALLOZYMIC CHARACTERS IN THE RUFOUS-COLLARED SPARROW,ZONOTRICHIA CAPENSIS

The geographical patterns of variation shown at 20 allozyme and non-enzymatic protein-coding loci, in 8 external, and in 12 skeletal morphological characters in the rufous-collared sparrow, Zonotrichia capensis, were analyzed in order to test the local (genetic) adaptation hypothesis regarding the origin and maintenance of vocal dialects in birds. Approximately 20 males were collected from each of four sites within each of six different dialect zones. There was significant variability in both external and skeletal morphology among all 24 sites and among dialect groups. Average Wright's corrected fixation coefficient (F_ST) was 0.118, indicating significant genetic differentiation among all sites, regardless of dialect. Hierarchical F statistics indicated that only 50% of among site variability was due to a dialect effect. Puna dialect sites were highly differentiated from all other sites with respect to both morphology (external and skeletal measures) and allozyme frequencies. Heterogeneity at the PGM-1 locus among puna scrub sites was the major cause of the high average F_ST across all sites, and within the puna scrub dialect. Average genetic differentiation among non-puna sites (F_ST = 0.018) was similar to differentiation among sites within each of the five non-puna dialect groups (mean F_ST = 0.0132 ± 0.0069). Hierarchical F statistics indicated that none of the among-site differentiation in this subset of samples was due to a dialect effect. These observations are not consistent with the local adaptation hypothesis. All significant genetic heterogeneity occurred among sites in mountainous habitats, and we suggest that topography and patchiness of habitat may have been major factors involved in population differentiation, rather than vocal dialects.     

A MULTISPECIES APPROACH TO THE ANALYSIS OF GENE FLOW IN MARINE SHORE FISHES

Ten species of marine shore fishes with a wide range of life-history strategies were collected from four areas in southern California, U.S.A., and Baja California, Mexico, and examined for patterns of genetic differentiation. Multilocus D and F_ST values (based on 32–42 presumptive gene loci in each species) were both negatively correlated with estimated dispersal capability. These results were robust to variations in the number and type of loci used in the analysis and are compatible with the hypothesis that levels of genetic differentiation in these shore fishes are determined primarily by gene flow and genetic drift. There is no a priori reason to expect the observed correlation to result from natural selection or historical factors. The findings thus suggest that populations of these shore fishes are in at least a quasi-equilibrium with respect to migration, mutation, and genetic drift. Present data were also used to compare estimates of mN_e obtained by three different methods. Estimates based on F_ST values calculated by the methods of Nei and Chesser (F_ST(N)) and Weir and Cockerham (F_ST(W)) were highly correlated, but F_ST(N) ≤ F_ST(W) for every species, leading to generally higher mN_e estimates for Nei and Chesser's method. Estimates of mN_e based on the frequency of private alleles (Slatkin, 1985a) were not as strongly correlated with dispersal capability as were F_ST and D values. A low incidence of private alleles in many species may be responsible for this relatively weak correlation and may limit the general usefulness of Slatkin's method. In spite of their sensitivity to natural selection, F_ST and D may be better indicators of relative gene flow levels for high gene flow species.     

PATTERNS OF REPRODUCTION,GENETIC DIVERSITY,AND GENETIC DIFFERENTIATION IN CALIFORNIA POPULATIONS OF THE GENICULATE CORALLINE ALGA LITHOTHRIX ASPERGILLUM (RHODOPHYTA)1

The reproductive composition and genetic diversity of populations of the red seaweed Lithothrix aspergillum Gray (O. Corallinales) were studied at three southern California sites (Shaw's Cove and Treasure Island, Laguna Beach; Indian Rock, Santa Catalina Island) and at a fourth site (Bodega Bay) located in northern California. Sexually reproducing populations were confined to southern California. Diploid individuals were numerically dominant over haploid (gametophytic) individuals at all sites. Intertidal and subtidal subpopulations from Shaw's Cove differed in their reproductive profiles. Most intertidal specimens found on emersed surfaces were densely branched, turf-forming, and bore tetrasporangial (68.6%), carposporangial (11.4%), or spermatangial (5.7%) conceptacles, reflecting a sexual life history; none produced asexual bispores. In contrast, 74.3% of the larger, loosely branched subtidal specimens bore bisporangial conceptacles indicative of asexual reproduction. Nearly 70% of the Indian Rock thalli showed no evidence of conceptacle formation. Only asexual, diploid bispore-producing thalli were obtained from the Bodega Bay site. Genetic diversity (mean number of alleles per locus, percent of polymorphic loci, and average expected heterozygosity) of diploid L. aspergillum populations varied with life-history characteristics and geographic location. A total of 30 alleles was inferred from zymograms of 16 loci examined by starch-gel electrophoresis; of these loci, 11 were polymorphic. The genetic diversity of sexual, diploid populations of L. aspergillum (alleles per locus [A/L] = 1.4-1.5; percent polymorphic loci [%P] = 37.5-50.0) was relatively high compared with other red seaweeds. Lowest diversity (A/L = 1.0; %P = 0.0) occurred in the exclusively asexual Bodega Bay population which consisted of genetic clones. All sexual L. aspergillum populations deviated significantly from Hardy-Wein-berg expectations due to lower than expected heterozygosity. Genetic differentiation (Wright's F_statistic [F_ST]; Nei's Genetic Distance [D]) among sexually reproducing southern California populations was low (F_ST= 0.030) on a local scale (ca. 5 km), suggesting high levels of gene flow, but high genetic differention (F_ST= 0.390 and 0.406) occurred among southern California populations separated by ca. 70 km. Very high genetic differentiation (F_ST= 0.583–0.683) was obtained between northern and southern California populations separated by 700–760 km. Our genetic and reproductive data suggest that the L. aspergillum population from Bodega Bay is sustained by perennation, vegetative propagation, or asexual reproduction by bispores and may represent an isolated remnant or a population established by a founder event.     

Historic and recent fragmentation coupled with altitude affect the genetic population structure of one of the world's highest tropical tree line species

Aim To assess the effects of altitude and historic and recent forest fragmentation on the genetic diversity and structure of the wind‐pollinated tropical tree line species Polylepis incana. Location One of the highest mountain forest regions of the world, located in the Eastern Cordillera of the Ecuadorian Andes. Methods We compared genetic diversity and structure of adult trees with those of seedlings (n= 118 in both cases) in nine forest stands spanning an altitudinal gradient from 3500 to 4100 m a.s.l. using amplified fragment length polymorphisms (AFLPs). Genetic diversity was calculated as percentage of polymorphic bands (P) and Nei's expected heterozygosity (He); genetic differentiation was assessed using analysis of molecular variance, Φ_ST statistics and Bayesian cluster analysis. Results Estimates of genetic diversity at the population level were significantly lower in seedlings than in adults. Genetic diversity (He‐value) was, in both cases, negatively correlated to altitude and positively correlated to population size in the seedlings. Genetic differentiation of the seedlings was approximately as high (φ_ST= 0.298) as that of the adults (φ_ST= 0.307), and geographical differentiation was clearly reflected in both AFLP profiles, with mountain ridges acting as barriers to gene flow. Main conclusions Our study provides evidence of a historic upslope migration of P. incana in central Ecuador. In addition, it highlights the detrimental effects of unexpectedly strong genetic isolation, both recent and historical, particularly for our wind‐pollinated species where the distance between forest stands was less than 25 km. We therefore additionally propose that in habitats with pronounced high‐mountain landscape structures, gene flow may be hampered to such an extent that species have a more pronounced sensitivity to habitat fragmentation, even among populations of wind‐pollinated trees.     

Background selection and FST: Consequences for detecting local adaptation

Background selection is a process whereby recurrent deleterious mutations cause a decrease in the effective population size and genetic diversity at linked loci. Several authors have suggested that variation in the intensity of background selection could cause variation in F_ST across the genome, which could confound signals of local adaptation in genome scans. We performed realistic simulations of DNA sequences, using recombination maps from humans and sticklebacks, to investigate how variation in the intensity of background selection affects F_ST and other statistics of population differentiation in sexual, outcrossing species. We show that, in populations connected by gene flow, Weir and Cockerham's (1984; Evolution, 38 , 1358) estimator of F_ST is largely insensitive to locus‐to‐locus variation in the intensity of background selection. Unlike F_ST, however, d_XY is negatively correlated with background selection. Moreover, background selection does not greatly affect the false‐positive rate in F_ST outlier studies in populations connected by gene flow. Overall, our study indicates that background selection will not greatly interfere with finding the variants responsible for local adaptation.     

High population differentiation and genetic variation in the endangered Mexican pine Pinus Rzedowskii (Pinaceae)

Pinus rzedowskii is an endangered pine species from Michoaca´n (central Me´xico), which has been previously reported from only three localities. Classified within the subgenus Strobus, it exhibits intermediate morphological characters between subgenera Strobus and Pinus. We analyzed genetic aspects that could shed light on the evolution and conservation of this species. The genetic structure of nine populations was examined using 14 isozyme loci. Pinus rzedowskii has a relatively high level of genetic variation with 46.8% of the loci assayed being polymorphic, a total of 35 alleles, and a mean heterozygosity per population of 0.219. We calculated Wright's F_ST statistic to estimate gene flow indirectly and to evaluate whether or not there was genetic structuring among populations. We found a marked differentiation among populations (F_ST = 0.175) and significant inbreeding (F_IS = 0.247). No pattern of isolation by distance was found. We also constructed a dendrogram based on a genetic distance matrix to obtain an overview of the possible historical relationships among populations. Finally, we found a convex relationship between the genetic distance among populations and the number of ancestral lineages, suggesting that demographically this species has not been at risk recently. Although endangered, with small and fragmented populations, P. rzedowskii shows higher levels of genetic variation than other conifer species with larger populations or similar conservation status.     

Contemporary human‐altered landscapes and oceanic barriers reduce bumble bee gene flow

Much of the world's terrestrial landscapes are being altered by humans in the form of agriculture, urbanization and pastoral systems, with major implications for biodiversity. Bumble bees are one of the most effective pollinators in both natural and cultivated landscapes, but are often the first to be extirpated in human‐altered habitats. Yet, little is known about the role of natural and human‐altered habitats in promoting or limiting bumble bee gene flow. In this study, I closely examine the genetic structure of the yellow‐faced bumble bee, Bombus vosnesenskii, across the southwestern US coast and find strong evidence that natural oceanic barriers, as well as contemporary human‐altered habitats, limit bee gene flow. Heterozygosity and allelic richness were lower in island populations, while private allelic richness was higher in island populations compared to mainland populations. Genetic differentiation, measured for three indices across the 1000 km study region, was significantly greater than the null expectation (F_ST = 0.041, F’_ST = 0.044 and D_est = 0.155) and correlated with geographic distance. Furthermore, genetic differentiation patterns were most strongly correlated with contemporary (2011) not past (2006, 2001) resistance maps calibrated for high dispersal limitation over oceans, impervious habitat and croplands. Despite the incorporation of dramatic elevation gradients, the analyses reveal that oceans and contemporary human land use, not mountains, are the primary dispersal barriers for B. vosnesenskii gene flow. These findings reinforce the importance of maintaining corridors of suitable habitat across the distribution range of native pollinators to promote their persistence and safeguard their ability to provide essential pollination services.     

Evolution of putative barrier loci at an intermediate stage of speciation with gene flow in campions (Silene)

Understanding the origin of new species is a central goal in evolutionary biology. Diverging lineages often evolve highly heterogeneous patterns of genetic differentiation; however, the underlying mechanisms are not well understood. We investigated evolutionary processes governing genetic differentiation between the hybridizing campions Silene dioica (L.) Clairv. and S. latifolia Poiret. Demographic modelling indicated that the two species diverged with gene flow. The best‐supported scenario with heterogeneity in both migration rate and effective population size suggested that a small proportion of the loci evolved without gene flow. Differentiation (F _ST) and sequence divergence (d _XY) were correlated and both tended to peak in the middle of most linkage groups, consistent with reduced gene flow at highly differentiated loci. Highly differentiated loci further exhibited signatures of selection. In between‐species population pairs, isolation by distance was stronger for genomic regions with low between‐species differentiation than for highly differentiated regions that may contain barrier loci. Moreover, differentiation landscapes within and between species were only weakly correlated, suggesting that linked selection due to shared recombination and gene density landscapes is not the dominant determinant of genetic differentiation in these lineages. Instead, our results suggest that divergent selection shaped the genomic landscape of differentiation between the two Silene species, consistent with predictions for speciation in the face of gene flow.     

GENETIC STRUCTURE OF GIANT CLAM (TRIDACNA MAXIMA) POPULATIONS IN THE WEST PACIFIC IS NOT CONSISTENT WITH DISPERSAL BY PRESENT-DAY OCEAN CURRENTS

The Pacific marine biota, particularly species with long planktonic larval stages, are thought to disperse widely throughout the Pacific via ocean currents. The little genetic data available to date has supported this view in that little or no significant regional differentiation of populations has been found over large geographical distances. However, recent data from giant clams has demonstrated not only significant regional differentiation of populations, but routes of gene flow that run perpendicular to the main present-day ocean currents. Extensive surveys of genetic variation at eight polymorphic loci in 19 populations of the giant clam Tridacna maxima, sampled throughout the West and Central Pacific, confirmed that the patterns of variation seen so far in T. gigas were not unique to that species, and may reflect a fundamental genetic structuring of shallow-water marine taxa. Populations of T. maxima within highly connected reef systems like the Great Barrier Reef were panmictic (average F_ST < 0.003), but highly significant genetic differences between reef groups on different archipelagos (average F_ST = 0.084) and between West and Central Pacific regions (average F_ST = 0.156) were found. Inferred gene flow was high (N_em usually > 5) between the Philippines and the Great Barrier Reef, between the Philippines and Melanesia (the Solomon Islands and Fiji), and between the Philippines and the Central Pacific island groups (Marshall Islands, Kiribati, Tuvalu and Cook Islands). Gene flow was low between these three sets of island chains (N_em < 2). These routes of gene flow are perpendicular to present-day ocean currents. It is suggested that the spatial patterns of gene frequencies reflect past episodes of dispersal at times of lower sea levels which have not been erased by subsequent dispersal by present-day circulation. The patterns are consistent with extensive dispersal of marine species in the Pacific, and with traditional views of dispersal from the Indo-Malay region. However, they demonstrate that dispersal along present-day ocean surface currents cannot be assumed, that other mechanisms may operate today or that major dispersal events are intermittent (perhaps separated by several thousands of years), and that the nature and timing of dispersal of Pacific marine species is more complex than has been thought.     

THE EFFECTIVE SIZE OF A HIERARCHICALLY STRUCTURED POPULATION

A knowledge of the effective size of a population (N_e) is important in understanding its current and future evolutionary potential. Unfortunately, the effective size of a hierarchically structured population is not, in general, equal to the sum of its parts. In particular, the inbreeding structure has a major influence on N_e. Here I link N_e to Wright's hierarchical measures of inbreeding, F_IS and F_ST, for an island-structured population (or metapopulation) of size N_T. The influence of F_ST depends strongly on the degree to which island productivity is regulated. In the absence of local regulation (the interdemic model), interdemic genetic drift reduces N_e. When such drift is combined with local inbreeding under otherwise ideal conditions, the effects of F_IS and F_ST are identical: increasing inbreeding either within or between islands reduces N_e, with N_e = N_T/[(1 + F_IS)(1 + F_ST) ? 2F_ISF_ST]. However, if islands are all equally productive because of local density regulation (the traditional island model), then N_e = N_T/[(1 + F_IS)(1 –F_ST)] and the effect of F_ST is reversed. Under the interdemic model, random variation in the habitat quality (and hence productivity) of islands act to markedly decrease N_e. This variation has no effect under the island model because, by definition, all islands are equally productive. Even when no permanent island structure exists, spatial differences in habitat quality can significantly increase the overall variance in reproductive success of both males and females and hence lower N_e. Each of these basic results holds when other nonideal factors are added to the model. These factors, deviations from a 1:1 sex ratio, greater than Poisson variance in female reproductive success, and variation in male mating success due to polygynous mating systems, all act to lower N_e. The effects of male and female variance on N_e have important differences because only females affect island productivity. Finally, it is noted that to use these relationships, F_IS and F_ST must be estimated according to Wright's definition (and corrected to have a zero expectation under the null model). A commonly used partitioning (θ, θ_g) can be biased if either island size or the number of islands is small.     

Fine scale patterns of migration and gene flow in the endangered mound spring snail, <Emphasis Type="Italic">Fonscochlea accepta</Emphasis> (Mollusca:Hydrobiidae) in arid Australia

Naturally patchy ecosystems are models for other systems currently undergoing anthropogenic habitat fragmentation. Understanding patterns of gene flow in these model systems can help us manage species and ecosystems threatened by human impacts. The mound springs of central Australia represent such a natural model ecosystem, supporting a unique aquatic fauna distributed within an inhospitable arid landscape. Moreover, these springs are being impacted by over extraction of groundwater, providing a unique opportunity to look at dispersal in a patchy habitat that is changing. The present study represents the first fine scale analysis of gene flow under different scenarios of habitat connectivity for the endangered mound spring snail, Fonscochlea accepta. Within a single spring group pairwise estimates of F _ST between springs were very low (ave 0.015) with no association found between genetic distance and a series of geographical distance matrices based on the degree of habitat connectivity among the springs: results implying unstructured dispersal and limited population isolation. However, results from Bayesian assignment tests showed that on average approximately 97% of snails were assigned to their spring of origin. In a preliminary analysis at broader geographic scales (among spring groups) the results from F _ST estimates, Mantel correlation analyses and assignment tests all suggest much stronger and geographically correlated population structuring. While varying results from F-statistics and Bayesian analyses stem from the different information they utilise, together they provide data on contemporary and historical estimates of gene flow and the influence of landscape dynamics on the spatial genetic patterning of the springs.     

POPULATION STRUCTURE AND LEVELS OF GENE FLOW IN THE MEDITERRANEAN LAND SNAIL ALBINARIA CORRUGATA (PULMONATA: CLAUSILIIDAE)

The amount of gene flow among local populations partly determines the relative importance of genetic drift and natural selection in the differentiation of such populations. Land snails, because of their limited powers for dispersal, may be particularly likely to show such differentiation. In this study, we directly estimate gene flow in Albinaria corrugata, a sedentary, rock-dwelling gastropod from Crete, by mark-recapture studies. In the same area, 23 samples were taken and studied electrophoretically for six polymorphic enzyme loci. The field studies indicate that the population structure corresponds closely to the stepping-stone model: demes are present on limestone boulders that are a few meters apart, and dispersal takes place mainly between adjacent demes. Average deme size (N) is estimated at 29 breeding individuals and the proportion of migrants per generation at 0.195 (Nm = 5.7). We find no reason to assume long-distance dispersal, apart from dispersal along occasional stretches of suitable habitat. Genetic subdivision of the population, as derived from F_ST values, corresponds to the direct estimate only at the lowest spatial level (distance between sample sites < 10 m), where values for Nm of 5.4 and 17.6 were obtained. In contrast, at the larger spatial scales, F_ST values give gene-flow estimates that are incompatible with the expected amount of gene flow at these scales. We explain these discrepancies by arguing that gene flow is in fact extremely limited, making correct estimates of Nm from F_ST impossible at the larger spatial scales. In view of these low levels of gene flow, it is concluded that both genetic drift and natural selection may play important roles in the genetic differentiation of this species, even at the lowest spatial scales.     

Abundant genetic diversity of the wild rice Zizania latifolia in central China revealed by microsatellites

Wild rice Zizania latifolia is a perennial emergent aquatic plant widely distributed across China. Wild populations of Z. latifolia are important to aquatic ecosystems and are valuable genetic resources for breeding. However, they have been faced with significant habitat losses in recent decades. For 10 nuclear microsatellites, high levels of genetic diversity (H_E = 0.572–0.636) were found across seven surveyed populations from central China. The main factors responsible for that were its long life history and predominant outcrossing reproductive system. Low genetic differentiation among populations was found based on Wright's F_ST = 0.098. Similarly, AMOVA analysis showed only 7.73% of the total molecular variation was attributed to inter‐population differentiation. The weak population structure of Z. latifolia could be due to high gene flow mediated by water or birds (Nm = 2.30, M = 2.18). Importantly, most populations exhibited mutation‐drift disequilibrium, suggesting a recent population decline. Based on the results, wild populations of Z. latifolia are expected to lose genetic diversity and increase genetic structure in future generations. Therefore, conservation management is urgently needed to maintain the genetic resources of Z. latifolia.     

Gene flow in malaria vector Anopheles stephensi (Diptera: Culicidae) across the Aravalli Hills in North-west India

The population structure of An. stephensi in North-west India was studied to assess the impact of the Aravalli Hills, as a barrier to gene flow using microsatellite markers. Large and significant genetic differentiation was found along the sides of, as well as across, the Aravalli Hills as the mean F_ST and R_ST on west vs. east of the Aravalli Hills were 0.213, 0.112 and 0.179, 0.056, respectively. Similarly, across the hills, mean values of F_ST and R_ST were 0.100 and 0.094, respectively. Genetic diversity on both sides did not vary significantly. The F_ST values were more sensitive than R_ST values, indicating that genetic drift might have caused genetic differentiation between populations. A positive correlation (r = 0.0149 and 0.157, respective to F_ST and R_ST) was found between genetic differentiations and geographic distances irrespective of the hills. Low level of gene flow was found along both sides (Nm = 0.92 and 0.14; west vs. east of Aravalli Hills, respectively) as compared to across the Aravalli Hills (Nm = 2.25). It was found that the Aravalli Hills are not working as an effective barrier to gene flow for An. Stephensi, maybe because of the low average height and discontinuous hills, however, the distance is playing a major role for differentiation between populations due to active mode of dispersal of An. stephensi mosquitoes which have a short flight range. All this information should help draw the strategies for genetic control of mosquitoes using transgenic mosquitoes.