GENE DISPERSAL AND SPATIAL GENETIC STRUCTURE

Spatial autocorrelation statistics have been studied in theoretical population genetic models and widely used in experimental studies of spatial structure in many plant and animal populations. However, the statistical properties of spatial autocorrelation statistics have remained uncharacterized. Little is known about how values of spatial autocorrelation statistics in population samples depend on the level of dispersal and scheme of sampling. In this paper, we characterize the statistical properties of join-count spatial autocorrelation statistics for population genetic surveys under various conditions of dispersal and sampling. The results indicate generally high statistical power. These results can provide a method to estimate gene dispersal based on standing spatial patterns of genetic variation observed within populations.     

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.     

THE INFLUENCE OF INTRASPECIFIC VARIATION IN DISPERSAL STRATEGIES ON THE GENETIC STRUCTURE OF PLANTHOPPER POPULATIONS

The hypothesis that levels of gene flow among populations are correlated with dispersal ability has typically been tested by comparing gene flow among species that differ in dispersal abilities, an approach that potentially confounds dispersal ability with other species-specific differences. In this study, we take advantage of geographic variation in the dispersal strategies of two wing-dimorphic planthopper species, Prokelisia marginata and P. dolus, to examine for the first time whether levels of gene flow among populations are correlated with intraspecific variation in dispersal ability. We found that in both of these coastal salt marsh–inhabiting species, population-genetic subdivision, as assessed using allozyme electrophoresis, parallels geographic variation in the proportion of flight-capable adults (macropters) in a population; in regions where levels of macroptery are high, population genetic subdivision is less than in regions where levels of macroptery are low. We found no evidence that geographic variation in dispersal capability influences the degree to which gene flow declines with distance in either species. Thus, both species provided evidence that intraspecific variation in dispersal strategies influences the genetic structure of populations, and that this effect is manifested in population-genetic structure at the scale of large, coastal regions, rather than in genetic isolation by distance within a region. This conclusion was supported by interspecific comparisons revealing that: (1) population-genetic structure (G_ST) of the two Prokelisia species correlated negatively with the mean proportion of flight-capable adults within a region; and (2) there was no evidence that the degree of isolation by distance increased with decreasing dispersal capability. Populations of the relatively sedentary P. dolus clustered by geographic region (using Nei's distances), but this was not the case for the more mobile P. marginata. Furthermore, gene flow among the two major regions we surveyed (Atlantic and Gulf Coasts) has been substantial in P. marginata, but relatively less in P. dolus. The results for P. marginata suggest that differences in the dispersal strategies of Atlantic and Gulf Coast populations occur despite extensive gene flow. We argue that gene flow is biased from Atlantic to Gulf Coast populations, indicating that selection favoring a reduction in flight capability must be intense along the Gulf. Together, the results of this study provide the first rigorous evidence of a negative relationship within a species between dispersal ability and the genetic structure of populations. Furthermore, regional variation in dispersal ability is apparently maintained by selective differences that outweigh high levels of gene flow among regions.     

SPATIAL SCALE OF GENETIC STRUCTURE AND AN INDIRECT ESTIMATE OF GENE FLOW IN EELGRASS,ZOSTERA MARINA

In this study, the first investigation of population structure in an aquatic angiosperm, I show that populations of a marine angiosperm (eelgrass, Zostera marina) are genetically differentiated at a number of spatial scales. I find also that there is no correspondence between geographic and genetic distances separating subpopulations, an increasingly common result in spatially stratified studies of genetic structure in marine invertebrates. F-statistics, calculated for two years from electrophoretic variation at five polymorphic allozyme loci, indicate significant genetic differentiation among sampling quadrats within each of two bays (θ = 0.064-0.208), between tide zones within a bay (θ = 0.025-0.157) and between bays (θ = 0.079). Spatial autocorrelation analysis was used to explore genetic differentiation at smaller spatial scales; estimated patch sizes (within which genetic individuals are randomly associated) indicated no appeciable genetic structure at scales less than 20 m × 20 m. Calculated values of F-statistics were a function of the spatial scale from which samples were drawn: increasing the size of the “subpopulation” included in calculation of fixation indices for the same “total” sample resulted in an increase in the magnitude of f (e.g., from 0.092 to 0.181) and a decrease in θ (e.g., from 0.186 to 0.025). On the basis of the best estimate of the spatial scale of subpopulations, the effective number of migrants per generation (N_em) ranges from 1.1 to 2.8. Genetic consequences of the disturbance regime in the eelgrass habitat sampled were extreme variation between years in the allele richness and proportion of heterozygotes in a sample and a positive relationship between the extinction probability of patches and the genetic variance among them. The changes in F-statistics as a function of sampling scale and the observation that θ among sampled quadrats was positively associated with the probability of extinction among quadrats indicated that indirect estimates of gene flow (N_em) calculated from θ should be cautiously interpreted in populations that may not yet be in drift-migration equilibrium.     

DEPENDENCE OF GENE FLOW ON GEOGRAPHIC DISTANCE IN TWO SOLITARY CORALS WITH DIFFERENT LARVAL DISPERSAL CAPABILITIES

When the level of gene flow among populations depends upon the geographic distance separating them, genetic differentiation is relatively enhanced. Although the larval dispersal capabilities of marine organisms generally correlate with inferred levels of average gene flow, the effect of different modes of larval development on the association between gene flow and geographic distance remains unknown. In this paper, I examined the relationship between gene flow and distance in two co-occurring solitary corals. Balanophyllia elegans broods large, nonfeeding planulae that generally crawl only short distances from their place of birth before settling. In contrast, Paracyathus stearnsii free-spawns and produces small planktonic larvae presumably capable of broad dispersal by oceanic currents. I calculated F-statistics using genetic variation at six (P. stearnsii) or seven (B. elegans) polymorphic allozyme loci revealed by starch gel electrophoresis, and used these F-statistics to infer levels of gene flow. Average levels of gene flow among twelve Californian localities agreed with previous studies: the species with planktonic, feeding larvae was less genetically subdivided than the brooding species. In addition, geographic isolation between populations appeared to affect gene flow between populations in very different ways in the two species. In the brooding B. elegans, gene flow declined with increasing separation, and distance explained 31% of the variation in gene flow. In the planktonically dispersed P. stearnsii distance of separation between populations at the scale studied (10–1000 km) explained only 1% of the variation in gene flow between populations. The mechanisms generating geographic genetic differentiation in species with different modes of larval development should vary fundamentally as a result of these qualitative differences in the dependence of gene flow on distance.     

EVOLUTION OF GENETIC INTEGRATION BETWEEN DISPERSAL AND COLONIZATION ABILITY IN A BIRD

Discrete behavioral strategies comprise a suite of traits closely integrated in their expression with consistent natural selection for such coexpression leading to developmental and genetic integration of their components. However, behavioral traits are often also selected to respond rapidly to changing environments, which should both favor their context-dependent expression and inhibit evolution of genetic integration with other, less flexible traits. Here we use a multigeneration pedigree and long-term data on lifetime fitness to test whether behaviors comprising distinct dispersal strategies of western bluebirds—a species in which the propensity to disperse is functionally integrated with aggressive behavior—are genetically correlated. We further investigated whether selection favors flexibility in the expression of aggression in relation to current social context. We found a significant genetic correlation between aggression and dispersal that is concordant with consistent selection for coexpression of these behaviors. To a limited extent, individuals modified their aggression to match their mate; however, we found no fitness consequences on such adjustments. These results introduce a novel way of viewing behavioral strategies, where flexibility of behavior, while often aiding an organism's fit in its current environment, may be limited and thereby enable integration with less flexible traits.     

FINE-SCALE GENETIC STRUCTURE OF A TURKEY OAK FOREST

Theoretical models and computer simulations of the genetic structure of a continuous population predict the existence of patches of highly inbred individuals when gene flow within the population is limited. A map of the three genotypes of a two-allele locus is expected to exhibit patches of homozygotes embedded in a matrix of heterozygotes, when gene flow is limited. A search for such patch structure was made on a 160 × 160 m plot within a continuous 60+ ha old-growth stand of Quercus laevis (turkey oak). Approximately 3400 trees were genotyped for 9 polymorphic loci using starch-gel electrophoresis, and the genetic structure was analyzed with spatial autocorrelation (both nominal and interval), hierarchical F statistics, and number-of-alleles-in-common. Adults (diameter at breast height > 0) and juveniles were analyzed separately but showed similar structure. While no distinct patch structure was found, a greater degree of relatedness was observed on a scale of 5 m–10 m than at greater distances, probably because of the limited acorn dispersal from maternal trees and a small amount of cloning by root sprouts. A computer simulation of a 10,000 tree forest breeding for 10,000 yr indicates that the effective neighborhood sizes (of randomly drawn seed- and pollen-donors) are both in excess of 440 individuals. The model thus cannot distinguish the observed data from panmictic mating.     

GENETIC FINGERPRINT-INFERRED POPULATION SUBDIVISION AND SPATIAL GENETIC TESTS FOR ISOLATION BY DISTANCE AND ADAPTATION IN THE COASTAL PLANT LIMONIUM CAROLINIANUM

This paper examines two wild populations of Limonium carolinianum for population genetic subdivision and spatial patterns of genetic variation in an attempt to simultaneously test for both the action of local adaptation to tidal gradients and isolation by distance (IBD). A VNTR (variable number of tandem repeats) genetic “fingerprinting” marker was used to infer relatedness among mapped plants in two populations. Band sharing within and between populations estimated F'_ST, an approximate measure of F_ST. Regression models were used to analyze the relationship between band sharing and spatial separation in tidal elevation and horizontal distance, as well as the relationship of fecundity differences with band sharing and spatial distance. Populations differed in band size frequency distributions and mean number of bands per profile and, therefore, likely differed in effective population size. F'_ST was estimated at 0.0678 and was significantly greater than F'_ST among randomly constructed subpopulations. Band sharing decreased 0.13% per meter in one population but showed no significant relation to distance in the other. In the population with significant IBD band sharing increased with increasingly different tidal elevation, contrary to an adaptive hypothesis, possibly due to directional gene flow or drift. Deme sizes were approximately 25 meters and greater than 100 meters, spanning larger areas than the entire environmental gradient. Fecundity differences were not associated with spatial parameters or band sharing. Unequal potential maternal fecundity measured as variance in number of seeds per maternal family was a significant source of genetic sampling variance. The VNTR marker employed is capable of detecting adaptation as identity by descent in ecological time and is an appropriate method for estimating the net evolutionary fate of polygenic traits. The results show that the net balance between selection along an environmental gradient and the effects of IBD and unequal maternal fecundity favor genetic differentiation by random processes in populations of Limonium.     

GENETIC CONSEQUENCES OF SEED DISPERSAL IN THREE SYMPATRIC FOREST HERBS. I. HIERARCHICAL POPULATION-GENETIC STRUCTURE

To examine the effects of seed dispersal on spatial genetic structure, we compare three sympatric species of forest herbs in the family Apiaceae whose fruits differ widely in morphological adaptations for animal-attached dispersal. Cryptotaenia canadensis has smooth fruits that are gravity dispersed, whereas Osmorhiza claytonii and Sanicula odorata fruits have appendages that facilitate their attachment to animals. The relative seed-dispersal ability among species, measured as their ability to remain attached to mammal fur, is ranked Sanicula > Osmorhiza > Cryptotaenia. We use a nested hierarchical sampling design to analyze genetic structure at spatial scales ranging from a few meters to hundreds of kilometers. Genetic differentiation among population subdivisions, estimated by average genetic distance and hierarchical F-statistics, has an inverse relationship with dispersal ability such that Cryptotaenia > Osmorhiza > Sanicula. In each species, genetic differentiation increases with distance among population subdivisions. Stochastic variation in gene flow, arising from seed dispersal by attachment to animals, may partly explain the weak relationship between pairwise spatial and genetic distance among populations and heterogeneity in estimates of single locus F-statistics. A hierarchical island model of gene flow is invoked to describe the effects of seed dispersal on population genetic structure. Seed dispersal is the predominant factor affecting variation in gene flow among these ecologically similar, taxonomically related species.     

GENETIC POPULATION STRUCTURE AND LEVELS OF GENE FLOW IN THE STREAM DWELLING WATERSTRIDER,AQUARIUS (=GERRIS) REMIGIS (HEMIPTERA: GERRIDAE)

Gene flow, in combination with selection and drift, determines levels of differentiation among local populations. In this study we estimate gene flow in a stream dwelling, flightless waterstrider, Aquarius remigis. Twenty-eight Aquarius remigis populations from Quebec, Ontario, New Brunswick, Iowa, North Carolina, and California were genetically characterized at 15 loci using starch gel electrophoresis. Sampling over two years was designed for a hierarchical analysis of population structure incorporating variation among sites within streams, streams within watersheds, watersheds within regions, and regions within North America. Hierarchical F statistics indicated that only sites within streams maintained enough gene flow to prevent differentiation through drift (Nm = 27.5). Above the level of sites within streams gene flow is highly restricted (Nm ≤ 0.5) and no correlation is found between genetic and geographic distances. This agrees well with direct estimates of gene flow based on mark and recapture data, yielding an N_e of approximately 170 individuals. Previous assignment of subspecific status to Californian A. remigis is not supported by genetic distances between those populations and other populations in North America. Previous suggestion of specific status for south-eastern A. remigis is supported by genetic distances between North Carolina populations and other populations in North America, and a high proportion of region specific alleles in the North Carolina populations. However, because of the high degree of morphological and genetic variability throughout the range of this species, the assignment of specific or subspecific status to parts of the range may be premature.     

EXAMINING THE FULL EFFECTS OF LANDSCAPE HETEROGENEITY ON SPATIAL GENETIC VARIATION: A MULTIPLE MATRIX REGRESSION APPROACH FOR QUANTIFYING GEOGRAPHIC AND ECOLOGICAL ISOLATION

Understanding the effects of landscape heterogeneity on spatial genetic variation is a primary goal of landscape genetics. Ecological and geographic variables can contribute to genetic structure through geographic isolation, in which geographic barriers and distances restrict gene flow, and ecological isolation, in which gene flow among populations inhabiting different environments is limited by selection against dispersers moving between them. Although methods have been developed to study geographic isolation in detail, ecological isolation has received much less attention, partly because disentangling the effects of these mechanisms is inherently difficult. Here, I describe a novel approach for quantifying the effects of geographic and ecological isolation using multiple matrix regression with randomization. I explored the parameter space over which this method is effective using a series of individual‐based simulations and found that it accurately describes the effects of geographic and ecological isolation over a wide range of conditions. I also applied this method to a set of real‐world datasets to show that ecological isolation is an often overlooked but important contributor to patterns of spatial genetic variation and to demonstrate how this analysis can provide new insights into how landscapes contribute to the evolution of genetic variation in nature.     

THE INFLUENCE OF DENSITY AND SEX ON PATTERNS OF FINE-SCALE GENETIC STRUCTURE

Natal philopatry is expected to limit gene flow and give rise to fine-scale spatial genetic structure (SGS). The banner-tailed kangaroo rat ( Dipodomys spectabilis ) is unusual among mammals because both sexes are philopatric. This provides an opportunity to study patterns of local SGS faced by philopatric and dispersing animals. We evaluated SGS using spatial genetic autocorrelation in two D. spectabilis populations (Rucker and Portal) over a 14-year temporal series that covered low, medium, and high population densities. Significantly positive autocorrelation values exist up to 800 m at Rucker and 400 m at Portal. Density was negatively associated with SGS (low >medium >high), and suggests that increases in density are accompanied by greater spatial overlap of kin clusters. With regard to sex-bias, we find a small but significant increase in the SGS level of males over females, which matches the greater dispersal distances observed in females. We observed variation in SGS over the ecological time scale of this study, indicating genetic structure is temporally labile. Our study is the first temporal exploration of the influence of density and sex on spatial genetic autocorrelation in vertebrate populations. Because few organisms maintain discreet kin clusters, we predict that density will be negatively associated with SGS in other species.     

Spatial correlations at different spatial scales are themselves highly correlated in isolation by distance processes

Although many properties of spatial autocorrelation statistics are well characterized, virtually nothing is known about possible correlations among values at different spatial scales, which ultimately would influence how inferences about spatial genetics are made at multiple spatial scales. This article reports the results of stochastic space-time simulations of isolation by distance processes, having a very wide range of amounts of dispersal for plants or animals, and analyses of the correlations among Moran's I-statistics for different mutually exclusive distance classes. In general, the stochastic correlations are extremely large (>0.90); however, the correlations bear a complex relationship with level of dispersal, spatial scale and spatial lag between distance classes. The correlations are so large that any existing or conceived statistical method that employs more than one distance class (or spatial scale) should not ignore them. This result also suggests that gains in statistical power via increasing sample size are limited, and that increasing numbers of assayed loci generally should be preferred. To the extent that sampling error for real data sets can be treated as white noise, it should be possible to account for stochastic correlations in formulating more precise statistical methods. Further, while the current results are for isolation by distance processes, they provide some guidance for some more complex stochastic space-time processes of landscape genetics. Moreover, the results hold for several popular measures other than Moran's I. In addition, in the results, the signal to noise ratios strongly decreased with distance, which also has several implications for optimal statistical methods using correlations at multiple spatial scales.     

PATTERNS OF ALLOZYME VARIATION IN DIPLOID AND TETRAPLOID CENTAUREA JACEA AT DIFFERENT SPATIAL SCALES

Abstract.— The extent and spatial patterns of genetic variation at allozyme markers were investigated within and between diploid and autotetraploid knapweeds (Centaurea jacea L. sensu lato, Asteraceae) at contrasted geographic scales: (1) among populations sampled from a diploid‐tetraploid contact zone in the northeastern part of the Belgian Ardennes, and (2) within mixed populations from that zone where diploids and tetraploids coexist. Our data were also compared with a published dataset by Sommer (1990) describing allozyme variation in separate diploid and tetraploid knapweeds populations collected throughout Europe. Genetic diversity was higher in tetraploids. In the Belgian Ardennes and within the mixed populations, both cytotypes had similar levels of spatial genetic structure, they were genetically differentiated, and their distributions of allele frequencies were not spatially correlated. In contrast, at the European scale, diploids and tetraploids did not show differentiated gene pools and presented a strong correlation between their patterns of spatial genetic variation. Numerical simulations showed that the striking difference in patterns observed at small and large geographic scales could be accounted for by a combination of (1) isolation by distance within cytotypes; and (2) partial reproductive barriers between cytotypes and/or recurrent formation of tetraploids. We suggest that this may explain the difficulty of the taxonomic treatment of knapweeds and of polyploid complexes in general.     

Hierarchical population structure and genetic diversity of lake trout (Salvelinus namaycush) in a dendritic system in Northern Labrador

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DISCORDANT DISTRIBUTION OF POPULATIONS AND GENETIC VARIATION IN A SEA STAR WITH HIGH DISPERSAL POTENTIAL

Patiria miniata, a broadcast‐spawning sea star species with high dispersal potential, has a geographic range in the intertidal zone of the northeast Pacific Ocean from Alaska to California that is characterized by a large range gap in Washington and Oregon. We analyzed spatial genetic variation across the P. miniata range using multilocus sequence data (mtDNA, nuclear introns) and multilocus genotype data (microsatellites). We found a strong phylogeographic break at Queen Charlotte Sound in British Columbia that was not in the location predicted by the geographical distribution of the populations. However, this population genetic discontinuity does correspond to previously described phylogeographic breaks in other species. Northern populations from Alaska and Haida Gwaii were strongly differentiated from all southern populations from Vancouver Island and California. Populations from Vancouver Island and California were undifferentiated with evidence of high gene flow or very recent separation across the range disjunction between them. The surprising and discordant spatial distribution of populations and alleles suggests that historical vicariance (possibly caused by glaciations) and contemporary dispersal barriers (possibly caused by oceanographic conditions) both shape population genetic structure in this species.     

SPATIAL GENETIC STRUCTURE OF CLONAL AND SEXUAL REPRODUCTION IN POPULATIONS OF ADENOPHORA GRANDIFLORA (CAMPANULACEAE)

The spatial distribution of clonal versus sexual reproduction in plant populations should generally have differing effects on the levels of biparental inbreeding and the apparent selfing rate, produced via mating by proximity through limited pollen dispersal. We used allozyme loci, join-count statistics, and Moran's spatial autocorrelation statistics to separate the spatial genetic structure caused by clonal reproduction from that maintained in sexually reproduced individuals in two populations of Adenophora grandiflora, a perennial herb. Join-count statistics showed that there were statistically significant clustering of clonal genotypes within distances less than 4 m. Both the entire populations and the sets of sexually reproduced individuals exhibited significant spatial autocorrelation at less than about 12 m, and the sexually reproduced individuals are substantially structured in an isolation-by-distance manner, consistent with a neighborhood size of about 50.