Genetic variation and structure in the expanding moss Pogonatum dentatum (Polytrichaceae) in its area of origin and in a recently colonized area

Genetic variation in the expanding moss species Pogonatum dentatum was studied using intersimple sequence repeat (ISSR) markers. The genetic consequences of range expansion were studied by comparing source populations in a mountain area with populations from a recently colonized lowland area in Sweden. Indices of genetic variation show slightly lower number of alleles per locus in the lowlands and a similar gene diversity in both areas. Three of four lowland populations had evidence of a recently passed bottleneck. Considerably higher haplotype diversity was found in the recently colonized lowlands compared to source populations in the mountains. Patterns of allelic diversity suggest that P. dentatum experiences loss of genetic variation through founder effects and genetic drift when expanding its distribution range. Higher haplotypic diversity, less linkage disequilibrium, and fewer compatible loci indicate that sexual recombination is relatively more important in the lowlands compared to the mountains. A likely explanation is higher success of establishment from spores in the lowlands, while clonal propagation predominates in the mountains. Less genetic differentiation among lowland populations indicates more gene flow in the lowland area, involving more spores and/or fragments moving among populations.     

Population differentiation in Trinidadian guppies (Poecilia reticulata): patterns and problems

Populations of the guppy, Poecilia reficulata , in N. Trinidad exhibit marked population differentiation in allozyme frequencies. Here we investigate six further populations electrophoretically at 25 enzyme-coding loci to examine patterns in geographical structuring, genotypic distributions and genetic diversity. With one exception, possibly related to an experimental introduction, populations divided broadly (dendrogram of Nei's mean genetic identity, Ī) in accordance with proposed ancestral colonization. Most populations were in Hardy Weinberg equilibrium, though some significant deficits in heterozygotes were detected. Incorporating information from published data, markedly hjgher levels of genetic diversity (mean observed heterozygosity, _o) were recorded in lowland [_o= 0.0382 ± k0406 ( s.e .), n = 9] compared with upstream populations [_o= 0.0112 ± 00034 (S.E.), n = 9]. Patterns are discussed in relation to historical and present-day evolutionary forces.     

Relationships between Population Size, Genetic Diversity and Fitness Components in the Rare Plant Dictamnus albus in Central Germany

An increasing number of studies support the hypothesis that smaller populations face a higher risk of extinction, and declining population sizes are therefore one of the focal points in plant conservation. In small populations, loss of genetic diversity is often related to reduced reproductive fitness. For the rare Dictamnus albus in Central Germany, an earlier study had already confirmed a significant correlation between population size and genetic diversity. In order to assess whether these variables correlate with fitness components, plant height; flower, fruit and seed production; and germination were studied in a total of 11 populations of different size. In the seven populations that were sampled over two consecutive years, differences among populations and among years were tested using a Two-Way ANOVA. Co-linearity among variables was assessed using principal component analysis (PCA), followed by calculating correlations between ordination axes and both population size and genetic diversity. Plant height and flower number were uncorrelated to the other variables and, together with germination, did not show any correlation to either population size or genetic diversity. However, both size and genetic diversity of populations correlated significantly with other PCA axes that reflected reproductive components such as fruit number, seed number, seed fruit ratio, and seed mass. Our results support the idea that reproduction is hampered in small populations and raise concerns over the loss of genetic diversity in D. albus.     

Genetic differentiation in relation to seed weights in wild soybean species (Glycine soja Sieb. & Zucc.)

Seed weight is one of the most important botanical and phylogenetic characteristics. The study objective was to understand whether there is genetic difference in different seed weights of wild soybean (Glycine soja Sieb. & Zucc.). A total of 563 wild soybean samples, which belonged separately to genebank germplasm accessions (220 samples), one regional population samples (293 plants) and one natural population (150 plants), were analyzed using microsatellite markers. Of four size classes, the smallest seed size type had the highest coefficient of variation in seed weight; small and large seed types had relatively great genetic differences. In the national genebank germplasm accessions, genetic diversity gradually decreased from quantitatively dominant small and middling seed types to less frequent large seed types. In the regional and natural populations, generally, small to middling seed sizes had higher genetic diversity than the smallest and larger seed sizes. Cluster analysis revealed genetic differences in seed size traits. The semi-wild type (Glycine gracilis Skvortzow) was the most genetically differentiated from other seed sizes. However, it was also clearly shown that the phylogenic genetic differentiation among seed sizes was less than the genetic differentiation among geographical habitat populations in the wild soybean species.     

Population genetic structure in Myrtus communis L. in a chronically fragmented landscape in the Mediterranean: can gene flow counteract habitat perturbation?

Ancient managed landscapes provide ideal opportunities to assess the consequences of habitat fragmentation on the patterns of genetic diversity and gene flow in long-lived plant species. Using amplified fragment length polymorphism (AFLP) and allozyme markers, we quantified seed-mediated gene flow and population genetic diversity and structure in 14 populations of Myrtus communis (myrtle), a common endozoochorous shrub species of forest patches in lowland agricultural Mediterranean areas. Overall, allozyme diversity for myrtle was low (P₉₅   =   25%; A   =   1.411; H_e = 0.085) compared to other known populations, and a significant portion of populations (57%) had lower levels of allelic diversity and/or heterozygosity than expected at random, as shown by simulated resampling of the whole diversity of the landscape. We found significant correlations between allozyme variability and population size and patch isolation, but no significant inbreeding in any population. Genetic differentiation among populations for both allozyme and AFLP markers was significant (Φ_ST = 0.144 and Φ_ST = 0.142, respectively) but an isolation-by-distance pattern was not detected. Assignment tests on AFLP data indicated a high immigration rate in the populations ( ca. 20–22%), likely through effective seed dispersal across the landscape by birds and mammals. Our results suggest that genetic isolation is not the automatic outcome of habitat destruction since substantial levels of seed-mediated gene flow are currently detectable. However, even moderate rates of gene flow seem insufficient in this long-lived species to counteract the genetic erosion and differentiation imposed by chronic habitat destruction.     

Extra-pair fertilization and effective population size in the song sparrow Melospiza melodia

The concept of effective population size (N_e) is used widely by conservation and evolutionary biologists as an indicator of the genetic state of populations, but its precision and relation to the census population size is often uncertain. Extra-pair fertilizations have the potential to bias estimates of N_e when they affect the number of breeders or their estimated reproductive success tallied from social pedigrees. We tested if the occurrence of extra-pair fertilizations influenced estimates of N_e in a resident population of song sparrows Melospiza melodia using four years of detailed behavioural and genetic data. Estimates of N_e based on social and genetic data were nearly identical and averaged c. 65% of the census population size over four years, despite that 28% of 471 independent young were sired outside of social pairs. Variance in male reproductive success also did not differ between estimates based on social and genetic data, indicating that extra-pair mating had little effect on the distribution of reproductive success in our study population. Our results show that the genetic assignment will not always be necessary to estimate N_e precisely.     

Population divergence in the amphicarpic species Amphicarpaea edgeworthii Benth. (Fabaceae): microsatellite markers and leaf morphology

Comparative analyses of the genetic differentiation in microsatellite markers ( F _ST) and leaf morphology characters ( Q _ST) of Amphicarpaea edgeworthii Benth. were conducted to gain insight into the roles of random processes and natural selection in the population divergence. Simple sequence repeat analyses on 498 individuals of 19 natural populations demonstrate that a significant genetic differentiation occurs among populations (mean F _ST = 0.578), and A. edgeworthii is a highly self-fertilized species (mean selfing rate s  = 0.989). The distribution pattern of genetic diversity in this species shows that central populations possess high genetic diversity (e.g. population WL with H _E = 0.673 and population JG with H _E = 0.663), whereas peripheral ones have a low H _E as in population JD (0.011). The morphological divergence of leaf shape was estimated by the elliptical Fourier analysis on the data from 11 natural and four common garden populations. Leaf morphology analyses indicate the morphological divergence does not show strong correlation with the genetic differentiation ( R  = 0.260, P  = 0.069). By comparing the 95% confidence interval of Q _ST with that of F _ST, Q _ST values for five out of 12 quantitative traits are significantly higher than the average F _ST value over eight microsatellite loci. The comparison of F _ST and Q _ST suggests that two kinds of traits can be driven by different evolutionary forces, and the population divergence in leaf morphology is shaped by local selections.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 505–516.     

Comparison of quantitative and molecular genetic variation of native vs. invasive populations of purple loosestrife (Lythrum salicaria L., Lythraceae)

Study of adaptive evolutionary changes in populations of invasive species can be advanced through the joint application of quantitative and population genetic methods. Using purple loosestrife as a model system, we investigated the relative roles of natural selection, genetic drift and gene flow in the invasive process by contrasting phenotypical and neutral genetic differentiation among native European and invasive North American populations ( Q _ST −  F _ST analysis). Our results indicate that invasive and native populations harbour comparable levels of amplified fragment length polymorphism variation, a pattern consistent with multiple independent introductions from a diverse European gene pool. However, it was observed that the genetic variation reduced during subsequent invasion, perhaps by founder effects and genetic drift. Comparison of genetically based quantitative trait differentiation ( Q _ST) with its expectation under neutrality ( F _ST) revealed no evidence of disruptive selection ( Q _ST >  F _ST) or stabilizing selection ( Q _ST <  F _ST). One exception was found for only one trait (the number of stems) showing significant sign of stabilizing selection across all populations. This suggests that there are difficulties in distinguishing the effects of nonadaptive population processes and natural selection. Multiple introductions of purple loosestrife may have created a genetic mixture from diverse source populations and increased population genetic diversity, but its link to the adaptive differentiation of invasive North American populations needs further research.     

Genetic Variation and Clonal Diversity of the Two Divergent Types of Clonal Populations of Leymus chinensis Tzvel on the Song Liao Steppe in the West of Northeastern China

The genetic variation and clonal diversity of two divergent types (grey-green and yellow-green) of clonal populations of Leymus chinensis Tzvel at 14 loci were compared. Total gene diversity (HT) and the coefficient of genetic differentiation (GST) were all higher for the yellow-green type (HT = 0.270; GST =0.186) than for the grey-green type (HT = 0.250; GST = 0.157) of L. chinensis. Rare alleles usually occurred as heterozygotes rather than homozygotes and significant deviations from Hardy-Weinberg equilibrium were found only at a few loci. This indicated that these two types of populations were mainly out-crossing. Clonal diversity, evenness of clones, and mean clone size were not significantly different between the two types. We found that differences between the clone size and genetic variation of the yellow-green type of populations occurred with different climate and habitat population groups. However, for the grey-green type of populations, these genetic variations decreased under conditions of different climate and habitat population groups.     

Widespread gene flow and high genetic variability in populations of water voles Arvicola terrestris in patchy habitats

Theory predicts that the impact of gene flow on the genetic structure of populations in patchy habitats depends on its scale and the demographic attributes of demes (e.g. local colony sizes and timing of reproduction), but empirical evidence is scarce. We inferred the impact of gene flow on genetic structure among populations of water voles Arvicola terrestris that differed in average colony sizes, population turnover and degree of patchiness. Colonies typically consisted of few reproducing adults and several juveniles. Twelve polymorphic microsatellite DNA loci were examined. Levels of individual genetic variability in all areas were high ( H _O= 0.69–0.78). Assignments of juveniles to parents revealed frequent dispersal over long distances. The populations showed negative F _IS values among juveniles, F _IS values around zero among adults, high F _ST values among colonies for juveniles, and moderate, often insignificant, F _ST values for parents. We inferred that excess heterozygosity within colonies reflected the few individuals dispersing from a large area to form discrete breeding colonies. Thus pre-breeding dispersal followed by rapid reproduction results in a seasonal increase in differentiation due to local family groups. Genetic variation was as high in low-density populations in patchy habitats as in populations in continuous habitats used for comparison. In contrast to most theoretical predictions, we found that populations living in patchy habitats can maintain high levels of genetic variability when only a few adults contribute to breeding in each colony, when the variance of reproductive success among colonies is likely to be low, and when dispersal between colonies exceeds nearest-neighbour distances.     

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.     

Temporal effective size estimates of a managed walleye Sander vitreus population and implications for genetic-based management

The goal of this research was to use the long-term fishery data set and DNA from archived scales of walleye Sander vitreus in Escanaba Lake, WI, U.S.A., to improve the understanding of the underlying mechanism(s) influencing genetic diversity in naturally recruiting populations. The introduced population of S. vitreus in Escanaba Lake has a low mean effective population size ( N _E) between 124·6 and 185·5 despite a mean census size ( N _C) of 4659 ( N _E/ N _C c. 0·04), suggesting an accelerated rate of genetic drift between 1952 and 2002. These values are smaller than the median N _E range of several studies suggesting typical N _E/ N _C ratios of 0·11–0·16 in a wide range of taxa. N _E increased steadily during the past two sampled decades (1992 and 2002) and was consistent with a lowering of the variance in S. vitreus reproductive success, possibly linked to a large, sustained exploitation (mean 28%) rate. Variance in reproductive success is one of the most important factors influencing N _E in species, like S. vitreus , which have a potential for large fecundities and large juvenile mortalities (type III survivorship). The N _B estimates across six sequential cohorts (age classes of S. vitreus , assayed from 1994 to 1999) was consistent with estimates of N _E reported for 1992–2002. These results, coupled with in-depth census and exploitation data, show that the genetic characteristics of Escanaba Lake S. vitreus have changed substantially and that management activities, such as supplemental stocking and harvest practices, have profoundly influenced the genetic dynamics of S. vitreus in this lake.     

Spatio-temporal population genetics of the Danish pine marten (Martes martes)

A spatio-temporal study of genetic variation in the Danish pine marten ( Martes martes ) populations from the Jutland peninsula and from the island of Sealand was performed using 11 microsatellite markers. Samples obtained from 1892 to 2003 were subdivided into historical (prior to 1970) and recent (from 1970) groups. As compared with the historical samples, there was a significant loss of genetic variation in the recent Jutland population, but not in Sealand. Effective population sizes were estimated using Bayesian-based software (TMVP). Historical effective population sizes were 5897 (90% highest probability density, HPD, limits: 1502–6849) in Jutland and 1300 (90% HPD limits: 224–5929) in Sealand, whereas recent effective population sizes were 14.7 (90% HPD limits: 10.9–23.5) in Jutland and 802 (90% HPD limits: 51.8–5510) in Sealand. Significant genetic differentiation ( F _ST) was found between the two historical samples, between the two recent samples, and between the historical and the recent sample in Jutland; whereas the F _ST value between the historical and the recent sample in Sealand was not significant. The significant genetic differentiation between the historical and the recent samples indicates changes in the genetic compositions over time, and the higher F _ST values between the two recent samples, as compared with the two historical samples, indicates that the populations in Sealand and Jutland have drifted apart within a short time span. No deviation from Hardy–Weinberg equilibrium was found within populations, indicating no further substructuring.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 457–464.     

Allozyme Diversity in Populations of Cymbidium goeringii (Orchidaceae)

Abstract: Using 14 allozyme loci, we investigated levels of genetic diversity within populations, and degree of genetic divergence among 24 populations of Cymbidium goeringii (Orchidaceae) in Korea and Japan. Cymbidium goeringii maintains high levels of genetic diversity both at population (mean expected heterozygosity, H _e = 0.238) and species levels (0.260). Means of H _e found in 24 populations were not significantly different from each other. About 90 % of the total variation in the species is common to all populations (mean G _ST = 0.108). No unique allele was found in any population. The indirect estimate of gene flow based on the mean G _ST was high ( Nm = 2.06). Nei's genetic identities for pairs of populations had high values (mean = 0.974 [SD = 0.013]). The Mantel-Z test showed a significant correlation between genetic distance and geographic distance. However, the mean G _ST value between 17 populations in Korea and seven Japanese populations was relatively low (0.029), even though the land connection between the southern Korean peninsula and southern Japanese archipelagos has not existed since the middle Pleistocene. Large numbers of small seeds of C. goeringii might travel long distances by wind from populations to populations both in Korea and Japan, increasing genetic diversity within populations and maintaining low genetic differentiation among populations.     

Stability of genetic structure and effective population size inferred from temporal changes of microsatellite DNA polymorphisms in the land snail Helix aspersa (Gastropoda: Helicidae)

Temporal evolution of genetic variability may have far-reaching consequences for a diverse array of evolutionary processes. Within the polders of the Bay of Mont-Saint-Michel (France), populations of the land snail Helix aspersa are characterized by a metapopulation structure with occasional extinction processes resulting from farming practices. A temporal survey of genetic structure in H . aspersa was carried out using variability at four microsatellite loci, in ten populations sampled two years apart. Levels of within-population genetic variation, as measured by allelic richness, H _e or F _is , did not change over time and similar levels of population differentiation were demonstrated for both sampling years. The extent of genetic differentiation between temporal samples of the same population established (i) a stable structure for six populations, and (ii) substantial genetic changes for four populations. Using classical F -statistics and a maximum likelihood method, estimates of the effective population size ( N _e) illustrated a mixture of stable populations with high N _e, and unstable populations characterized by very small N _e estimates (of 5–11 individuals). Owing to human disturbances, intermittent gene flow and genetic drift are likely to be the predominant evolutionary processes shaping the observed genetic structure. However, the practice of multiple matings and sperm storage is likely to provide a reservoir of variability, minimizing the eroding genetic effects of population size reduction and increasing the effective population size.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 89–102.     

Gene flow and effective population size in Lepanthes (Orchidaceae): a case for genetic drift

Genetic drift can play an important role in population differentiation, particularly when effective population sizes are small and gene flow is limited. Such conditions are suspected to be common in the species-rich Orchidaceae. We investigated the likelihood of genetic drift in natural populations of three endemic species of Lepanthes (Orchidaceae) from Puerto Rico. We estimated effective population size, Ne, using three ecologically based methods. Two of the three estimates were based on variance in reproductive potential and the third was based on coalescence time. All estimates of Ne were usually <40% of the standing population size, resulting in values of <20 individuals per population. Based on starch gel electrophoresis of isozymes, Nm estimates suggest restricted gene flow among populations in the range of one or less successful migrant per generation. Genetic differentiation among populations is expected under such conditions from random genetic drift. Indeed we observed high genetic differentiation among populations (L. rubripetala, F_ST> G_ST, 6; θ.248, 0.266, 0.293; L. rupestris, 0.148, 0.169, 0.138; L. eltoroensis, 0.251, 0.219, 0.218, respectively). Genetic drift is likely to be important for population differentiation in Lepanthes as a result of small effective population sizes and restricted gene flow.     

Population genetic structure and levels of inbreeding depression in the Mediterranean island endemic Cyclamen creticum (Primulaceae)

The variation and evolution of reproductive traits in island plants have much attention from conservation and evolutionary biologists. However, plants on islands in the Mediterranean region have very little attention. In the present study, we examine the floral biology and mating system of Cyclamen creticum , a diploid perennial herb endemic to Crete and Karpathos. Our purpose is to quantify (1) variation and covariation of floral traits related to the mating system, (2) the ability of the species to self in the absence of pollinators and its relative performance on selfing and outcrossing and (3) generic diversity within and among populations. Pollen/ovule ratios were indicative of a xenogamous species. A controlled pollination experiment showed that the species is self-compatible but is unable to set seed, in the absence of pollinators, probably due to stigma-anther separation. A multiplicative estimate of inbreeding depression based on fruit maturation, seed number and percentage seed germination gave δ= 0.38 Population genetic diversity was high, 54.76% polymorphic loci, a mean of 1.78 alleles per locus and a mean observed heterozygosity of 0.053. F -statistics nevertheless indicated high inbreeding rates (mean F ^is= 0.748) in natural populations, and low levels of population differentiation (mean F^is= 0.168). C. creticum thus appears to have a mixed-mating system with high levels of (pollinator) mediated inbreeding (either by facilitated selfing, geitonogamy or biparental inbreeding) in natural populations.     

Microbial diversity--insights from population genetics

Although many environmental microbial populations are large and genetically diverse, both the level of diversity and the extent to which it is ecologically relevant remain enigmatic. Because the effective (or long-term) population size, N_e, is one of the parameters that determines population genetic diversity, tests and simulations that assume selectively neutral mutations may help to identify the processes that have shaped microbial diversity. Using ecologically important genes, tests of selective neutrality suggest that adaptive as well as non-adaptive types of selection act and that departure from neutrality may be widespread or restricted to small groups of genotypes. Population genetic simulations using population sizes between 10³ and 10⁷ suggest extremely high levels of microbial diversity in environments that sustain large populations. However, census and effective population sizes may differ considerably, and because we know nothing of the evolutionary history of environmental microbial populations, we also have no idea what N_e of environmental populations is. On the one hand, this reflects our ignorance of the microbial world. On the other hand, the tests and simulations illustrate interactions between microbial diversity and microbial population genetics that should inform our thinking in microbial ecology. Because of the different views on microbial diversity across these disciplines, such interactions are crucial if we are to understand the role of genes in microbial communities.     

Populations do not become less genetically diverse or more differentiated towards the northern limit of the geographical range in clonal Vaccinium stamineum (Ericaceae)

Geographically peripheral populations are expected to exhibit lower genetic diversity and higher differentiation than central populations because of their smaller size and greater spatial isolation. In plants, a shift from sexual to clonal asexual reproduction may further reduce diversity and increase differentiation. Here, these predictions were tested by assaying 36 inter-simple sequence repeat (ISSR) polymorphisms in 21 populations of the woody, clonal plant Vaccinium stamineum in eastern North America, from the range center to its northern limit where it has 'threatened' status. Populations decline in frequency, but not size or sexual reproductive output, across the range. Within-population diversity did not decline towards range margins. Modest genetic differentiation among populations increased slightly towards range margins and in small populations with high clonal propagation and low seed production, although none of these trends was significant. Low seed production and high clonal propagation were not associated with large-scale clonal spread. By combining demographic and genetic data, this study determined that increased population isolation, rather than reduced population size, can account for the weak increase in genetic differentiation at range margins.     

Marked genetic divergence revealed by allozymes among populations of the guppy Poecilia reticulata (Poeciliidae), in Trinidad

Populations of the guppy Poecilia reticulata from six locations in N. Trinidad were examined by starch gel electrophoresis to estimate their degree of genetic divergence. Variability at seven enzyme-coding loci demonstrated that populations differed markedly in allele frequencies with some allelic substitution between sites (for 23 loci, Nei's mean genetic identity, Ī ranged from 0.869–1.00; the coefficient of gene differentiation, G _ST= 0.086; and the absolute differentiation between populations, _m= 0.044). There was a good correspondence between degree of physical isolation and extent of genetic differentiation, as exemplified by the partitioning of gene diversity (Nei's gene diversity analysis = 66% between river basins; 32% within river basins; 2% within rivers), though there was considerable variability in the contribution of individual loci. Most populations were in Hardy-Weinberg equilibrium, and any significant deviations were due exclusively to heterozygote deficiencies. The patterns of population differentiation are discussed in relation to probable past geological and historical events, and present-day evolutionary forces. Notable among these are previous continental land connections, periodic immigration from N.E. South America, and post-colonization events including differential predation, sexual selection, apparent restricted vagility and small effective population sizes.