Genetic Divergence and Signatures of Natural Selection in Marginal Populations of a Keystone,Long-Lived Conifer,Eastern White Pine (Pinus strobus) from Northern Ontario

Marginal populations are expected to provide the frontiers for adaptation, evolution and range shifts of plant species under the anticipated climate change conditions. Marginal populations are predicted to show genetic divergence from central populations due to their isolation, and divergent natural selection and genetic drift operating therein. Marginal populations are also expected to have lower genetic diversity and effective population size (N _e) and higher genetic differentiation than central populations. We tested these hypotheses using eastern white pine (Pinus strobus) as a model for keystone, long-lived widely-distributed plants. All 614 eastern white pine trees, in a complete census of two populations each of marginal old-growth, central old-growth, and central second-growth, were genotyped at 11 microsatellite loci. The central populations had significantly higher allelic and genotypic diversity, latent genetic potential (LGP) and N _e than the marginal populations. However, heterozygosity and fixation index were similar between them. The marginal populations were genetically diverged from the central populations. Model testing suggested predominant north to south gene flow in the study area with curtailed gene flow to northern marginal populations. Signatures of natural selection were detected at three loci in the marginal populations; two showing divergent selection with directional change in allele frequencies, and one balancing selection. Contrary to the general belief, no significant differences were observed in genetic diversity, differentiation, LGP, and N _e between old-growth and second-growth populations. Our study provides information on the dynamics of migration, genetic drift and selection in central versus marginal populations of a keystone long-lived plant species and has broad evolutionary, conservation and adaptation significance.     

Genetic diversity and structuring across the range of a widely distributed ladybird: focus on rear‐edge populations phenotypically divergent

Population genetics and phenotypic structures are often predicted to vary along the geographic range of a species. This phenomenon would be accentuated for species with large range areas, with discontinuities and marginal populations. We herein compare the genetic patterns of central populations of Coccinella septempunctata L. with those of two phenotypically differentiated populations considered as rear‐edge populations and subspecies based on phenotype (Algeria and Japan). According to the central‐marginal model and expected characteristics of rear‐edge populations, we hypothesize that these rear‐edge populations have (1) a reduced genetic diversity, resulting from their relative isolation over long periods of time, (2) a higher population genetic differentiation, explained by low contemporary gene flow levels, and (3) a relationship between genetic diversity characteristics and phenotypes, due to historical isolation and/or local adaptation. Based on genotyping of 28 populations for 18 microsatellite markers, several levels of regional genetic diversity and differentiation are observed between and within populations, according to their localization: low within‐population genetic diversity and higher genetic differentiation of rear‐edge populations. The genetic structuring clearly dissociates the Algerian and Eastern Asia populations from the others. Geographical patterns of genetic diversity and differentiation support the hypothesis of the central‐marginal model. The pattern observed is in agreement with the phenotypic structure across species range. A clear genetic break between populations of Algeria, the Eastern Asia, and the remaining populations is a dominant feature of the data. Differential local adaptations, absence of gene flow between marginal and central populations, and/or incapacity to mate after colonization, have contributed to their distinct genotypic and phenotypic characteristics.     

Rapid range expansion increases genetic differentiation while causing limited reduction in genetic diversity in a damselfly

Many ectothermic species are currently expanding their geographic range due to global warming. This can modify the population genetic diversity and structure of these species because of genetic drift during the colonization of new areas. Although the genetic signatures of historical range expansions have been investigated in an array of species, the genetic consequences of natural, contemporary range expansions have received little attention, with the only studies available focusing on range expansions along a narrow front. We investigate the genetic consequences of a natural range expansion in the Mediterranean damselfly Coenagrion scitulum, which is currently rapidly expanding along a broad front in different directions. We assessed genetic diversity and genetic structure using 12 microsatellite markers in five centrally located populations and five recently established populations at the edge of the geographic distribution. Our results suggest that, although a marginal significant decrease in the allelic richness was found in the edge populations, genetic diversity has been preserved during the range expansion of this species. Nevertheless, edge populations were genetically more differentiated compared with core populations, suggesting genetic drift during the range expansion. The smaller effective population sizes of the edge populations compared with central populations also suggest a contribution of genetic drift after colonization. We argue and document that range expansion along multiple axes of a broad expansion front generates little reduction in genetic diversity, yet stronger differentiation of the edge populations.     

Genetic diversity of populations of a rare species of Dictamnus gymnostylis Stev. in Bashkir Cis-Urals

We examined genetic diversity in populations of Dictamnus gymnostylis Stev., a rare species growing in the Bashkir Cis-Urals, based on the analysis of 8 gene-enzyme systems and detected a fairly high level of intraspecific genetic diversity and population differentiation. We determined the average number of alleles per locus (A) as 1.57; the portion of polymorphic loci (P ₉₅) as 0.508; the observed heterozygosity (H _o) as 0.139; and the expected heterozygosity (H _e) as 0.169. Of the total genetic diversity, 88.3% stems from variability within populations, and 11.7% is due to variation among populations. The average value for the Nei’s genetic distance (D) constituted 0.028.     

Genetic diversity of nine faba bean (Vicia faba L.) populations revealed by isozyme markers

Seven isozyme systems (Sod, 6-Pgd, Me, Est, Skdh, Fdh and Gdh) representing nine loci were used to study the genetic diversity of nine faba bean populations. Seven loci revealed polymorphic bands and showed the same quaternary structure as that found in several species. They revealed a high number of phenotypes. Indeed, from 3 to 9 phenotypes per locus were investigated in this study. The percentage of polymorphic loci (P = 59.3 %) was higher than that mentioned in the autogamous species (P = 20.3 %) and less than the optimum (P=96 %) indicated for allogamous plants. Total genetic diversity (H _T) and within population genetic diversity (H _S) were estimated with the isozyme markers. The contribution of among population genetic diversity (D _ST) to total genetic diversity was 22%. Enzyme markers pointed out an average inbreeding level for whole population (F _IT) and within population (F _IS). Within population genetic diversity represents 78% of total diversity. Intra-population genetic diversity (H _S = 0.206) was ranged with the respect of allogamous species and was clearly higher than that of among population genetic diversity (D _ST = 0.057) indicating an out-crossing predominance in the studied populations. The expected heterozygosity was higher than that observed heterozygosity at the allogamous species was confirmed in this study. Although, the mean estimated gene flow was less than 1(Nm=0.814), the dendrogram based on Nei’s genetic distance of the 9 populations using UPGMA method showed some genetic drift between populations.     

Large‐scale patterns in genetic variation,gene flow and differentiation in five species of European Coenagrionid damselfly provide mixed support for the central‐marginal hypothesis

Recently, an increased effort has been directed towards understanding the distribution of genetic variation within and between populations, particularly at central and marginal areas of a species’ distribution. Much of this research is centred on the central‐marginal hypothesis, which posits that populations at range margins are sparse, small and genetically diminished compared to those at the centre of a species’ distribution range. We tested predictions derived from the central‐marginal hypothesis for the distribution of genetic variation and population differentiation in five European Coenagrionid damselfly species. We screened genetic variation (microsatellites) in populations sampled in the centre and margins of the species’ latitudinal ranges, assessed genetic diversity (H_S) in the populations and the distribution of this genetic diversity between populations (F_ST). We further assessed genetic substructure and migration with Bayesian assignment methods, and tested for significant associations between genetic substructure and bioclimatic and spatial (altitude and latitude) variables, using general linearized models. We found no general adherence to the central‐marginal hypothesis; instead we found that other factors such as historical or current ecological factors often better explain the patterns uncovered. This was illustrated in Coenagrion mercuriale whose colonisation history and behaviour most likely led to the observation of a high genetic diversity in the south and lower genetic diversity with increasing latitude, and in C. armatum and C. pulchellum whose patterns of low genetic diversity coupled with the weakest genetic differentiation at one of their range margins suggested, respectively, possible range shifts and recent, strong selection pressure.     

Genetic Diversity and Population Structure in Chinese Indigenous Poplar (Populus simonii) Populations Using Microsatellite Markers

Populus simonii Carr. is an important ecological and commercial breeding species in northern China; however, human interference during the last few centuries has led to the reduction and fragmentation of natural populations. To evaluate genetic diversity and differentiation within and among existing populations, we used 20 microsatellite markers to examine the genetic variation and structure of 16 natural populations. Our results indicated that the level of genetic diversity differed among populations, with average number of alleles per locus (AR) and expected heterozygosity (H _e) ranging from 3.7 to 6.11 and 0.589 to 0.731, respectively. A marginal population from Qilian in the Qinghai–Tibetan Plateau showed the highest values (AR?=?6.11, H _e?=?0.731), and the Zhangjiakou and Yishui populations showed the lowest values (AR?=?4.08, H _e?=?0.589 and AR?=?3.7, H _e?=?0.604). The inbreeding coefficient (F _IS) values for all populations were positive, which indicated an excess of homozygotes. The microsatellites allowed the identification of a significant subpopulation structure (K?=?3), consistent with an isolation by distance model for P. simonii populations. Additionally, molecular variance analysis revealed that 14.2 % of the variation resided among populations, and 85.8 % could be attributed to variation within populations. These data provide valuable information for natural resource conservation and for optimization of breeding programs in the immediate future.     

Rangewide Genetic Diversity in Natural Populations of Chinese Pine (Pinus tabulaeformis)

Thirteen natural populations from throughout the range of the Chinese pine (Pinus tabulaeformis Carr.) were examined using inter-simple sequence repeat markers to characterize the genetic structure at the species level and to compare the extent and distribution of genetic variation among central, intermediate, and marginal populations. Although the total genetic variation in the Chinese pine was mainly maintained within populations, the genetic differentiation among populations was significant (P < 0.001). The genetic divergence was significantly correlated with geographic distance (P < 0.05). Genetic diversity tended to decrease from the central to intermediate and marginal populations. The marginal populations had significantly lower intrapopulation genetic diversity than central populations (P < 0.05). Cluster analysis based on Nei’s unbiased genetic distances confirmed the difference among four central populations and the rest. Both historical and contemporary factors may have played key roles in shaping the spatial genetic structure of this species.     

Phylogeography and Conservation Genetics of the Common Wall Lizard,Podarcis muralis,on Islands at Its Northern Range

Populations at range limits are often characterized by lower genetic diversity, increased genetic isolation and differentiation relative to populations at the core of geographical ranges. Furthermore, it is increasingly recognized that populations situated at range limits might be the result of human introductions rather than natural dispersal. It is therefore important to document the origin and genetic diversity of marginal populations to establish conservation priorities. In this study, we investigate the phylogeography and genetic structure of peripheral populations of the common European wall lizard, Podarcis muralis, on Jersey (Channel Islands, UK) and in the Chausey archipelago. We sequenced a fragment of the mitochondrial cytochrome b gene in 200 individuals of P. muralis to infer the phylogeography of the island populations using Bayesian approaches. We also genotyped 484 individuals from 21 populations at 10 polymorphic microsatellite loci to evaluate the genetic structure and diversity of island and mainland (Western France) populations. We detected four unique haplotypes in the island populations that formed a sub-clade within the Western France clade. There was a significant reduction in genetic diversity (H_O, H_E and A_R) of the island populations in relation to the mainland. The small fragmented island populations at the northern range margin of the common wall lizard distribution are most likely native, with genetic differentiation reflecting isolation following sea level increase approximately 7000 BP. Genetic diversity is lower on islands than in marginal populations on the mainland, potentially as a result of early founder effects or long-term isolation. The combination of restriction to specific localities and an inability to expand their range into adjacent suitable locations might make the island populations more vulnerable to extinction.     

PHENOTYPIC DIFFERENTIATION AT SOUTHERN LIMIT BORDERS: THE CASE STUDY OF TWO FUCOID MACROALGAL SPECIES WITH DIFFERENT LIFE‐HISTORY TRAITS1

Marginal populations are often geographically isolated, smaller, and more fragmented than central populations and may frequently have to face suboptimal local environmental conditions. Persistence of these populations frequently involves the development of adaptive traits at phenotypic and genetic levels. We compared population structure and demographic variables in two fucoid macroalgal species contrasting in patterns of genetic diversity and phenotypic plasticity at their southern distribution limit with a more central location. Models were Ascophyllum nodosum (L.) Le Jol. (whose extreme longevity and generation overlap may buffer genetic loss by drift) and Fucus serratus L. (with low genetic diversity at southern margins). At edge locations, both species exhibited trends in life‐history traits compatible with population persistence but by using different mechanisms. Marginal populations of A. nodosum had higher reproductive output in spite of similar mortality rates at all life stages, making edge populations denser and with smaller individuals. In F. serratus, rather than demographic changes, marginal populations differed in habitat, occurring restricted to a narrower vertical habitat range. We conclude that persistence of both A. nodosum and F. serratus at the southern‐edge locations depends on different strategies. Marginal population persistence in A. nodosum relies on a differentiation in life‐history traits, whereas F. serratus, putatively poorer in evolvability potential, is restricted to a narrower vertical range at border locations. These results contribute to the general understanding of mechanisms that lead to population persistence at distributional limits and to predict population resilience under a scenario of environmental change.     

Genetic diversity and parentage assignment in Dojo loach,Misgurnus anguillicaudatus based on microsatellite markers

In this study, genetic diversity and structure of three Misgurnus anguillicaudatus populations from three different geographical locations in China (Hunan, Hubei and Henan province) were investigated using microsatellite markers. High level of genetic diversity of all three populations was revealed by expected heterozygosity (H_e), observed heterozygosity (H_o) and allele number. Significant genetic differentiations were found between all pairs of populations. The efficiency of eight microsatellite markers in parentage assignment of 540 progeny from twenty full-sib families was evaluated. Simulation based on allele frequency data demonstrated that probabilities of exclusion per locus range from 0.313 to 0.825 when no parent information is available and 0.504 to 0.904 when one parent is known. The assignment success rate based on the real data using eight markers was 96.85%. This study indicates that these M. anguillicaudatus resources are valuable genetic and breeding material for aquaculture and the microsatellite markers will be useful for investigation of genetic background and molecular marker-assisted selective breeding in this species.     

Extremely low genetic diversity and extensive genetic admixture at the northern range margins of Bombax ceiba

Understanding the evolutionary processes in species at the margins of their range is of great significance, because marginal populations may harbor local adaptations and will initiate further expansion in response to changes in the environment. Here we examined genetic variation in two nuclear genes and one chloroplast intergenic spacer in 13 northern marginal populations and one geographically central population of Bombax ceiba, a tree distributed mainly in tropical regions. Our results revealed an extremely low level of genetic diversity in each population at the northern margin of its range and strong genetic differentiation between southern China and South Asia. Cultivated and natural populations showed no significant differences in genetic variability. Genetic admixture in a nuclear gene was detected in 10 of the 13 populations at the northern margin of their range. The founder effect, in which a small number of individuals colonize the northern margins of its range, may explain the extremely low genetic diversity. During the establishment of new populations, different source populations may mix and undergo further genetic drift and differentiation. This study indicates that patterns of genetic diversity in tropical species at the margin of their range may also be severely influenced by founder effects.     

Range-edge genetic diversity: locally poor extant southern patches maintain a regionally diverse hotspot in the seagrass Zostera marina

Refugial populations at the rear edge are predicted to contain higher genetic diversity than those resulting from expansion, such as in post-glacial recolonizations. However, peripheral populations are also predicted to have decreased diversity compared to the centre of a species' distribution. We aim to test these predictions by comparing genetic diversity in populations at the limits of distribution of the seagrass Zostera marina, with populations in the species' previously described central diversity 'hotspot'. Zostera marina populations show decreased allelic richness, heterozygosity and genotypic richness in both the 'rear' edge and the 'leading' edge compared to the diversity 'hotspot' in the North Sea/Baltic region. However, when populations are pooled, genetic diversity at the southern range is as high as in the North Sea/Baltic region while the 'leading edge' remains low in genetic diversity. The decreased genetic diversity in these southern Iberian populations compared to more central populations is possibly the effect of drift because of small effective population size, as a result of reduced habitat, low sexual reproduction and low gene flow. However, when considering the whole southern edge of distribution rather than per population, diversity is as high as in the central 'hotspot' in the North Sea/Baltic region. We conclude that diversity patterns assessed per population can mask the real regional richness that is typical of rear edge populations, which have played a key role in the species biogeographical history and as marginal diversity hotspots have very high conservation value.     

Population structure and genetic diversity of Dysosma versipellis (Berberidaceae), a rare endemic from China

Dysosma versipellis (Berberidaceae) is an endangered and endemic species in China. To provide scientific foundation for formulating conservation strategies, we sampled six extant populations of this species and assessed the levels and patterns of genetic diversity using ISSR markers (11 primers). Of 144 bands detected 57.64% were polymorphic, but on average only 20.72% were polymorphic within populations. Our results revealed a low level of intraspecific genetic diversity (at population level: H_pop = 0.082, H_B = 0.177, SI = 0.1194; at species level: H_pop = 0.207, H_B = 0.378, SI = 0.3069). A high level of genetic differentiations among populations was detected based on Nei's genetic diversity analysis (60%), AMOVA analysis (65%), and Bayesian analysis (53%). The low levels of heterozygosity and high genetic differentiation observed in D. versipellis may be the consequence of low rate of natural recruitment, clonal growth, gene drift, and habitat fragmentation. Based on this, we suggest that in situ conservation be an important and practical measure for maintaining the genetic diversity of this species. Ex situ conservation should sample from different populations across the distribution range of the species to conserve high genetic diversity.     

植物边缘种群遗传多样性研究进展

边缘种群指地理分布边缘可检测到的一定数量的同种个体集合, 准确评价其遗传多样性对于理解第四纪冰期后气候变化对物种边缘扩展或收缩、遗传资源保护与利用以及物种形成等有重要意义。该文探讨了维持植物边缘种群遗传多样性的进化机制, 分析交配系统对物种边缘及其遗传多样性的影响, 比较了边缘与中心种群遗传多样性的差异及其形成的生态与进化过程, 并探讨了边缘种群遗传多样性与其所在的群落物种多样性的关系及理论基础。该文提出今后研究的重点是应用全基因组序列或转录组基因序列研究前缘-后缘种群之间或边缘-中心种群之间的适应性差异, 边缘种群与所在群落其他物种之间相互作用的分子机制, 深入解析边缘种群对环境的适应及边缘种群遗传多样性与群落物种多样性关系的生态与进化过程。     

Genetic diversity and population structure of the ark shell Scapharca broughtonii along the coast of China based on microsatellites

As a commercially important species in East Asia, the natural resources of Scapharca broughtonii have been suffering from severe population decline across its main habitats. In China, recovery efforts for S. broughtonii are in progress. To provide scientific bases for fisheries management and conservation program, genetic diversity and population structure of seven wild populations of S. broughtonii from the northern China coast was assessed using seven microsatellite loci in this study. High genetic diversity was present in all the seven populations, as observed in mean allelic richness per locus (11.3–12.5), and average expected heterozygosity (0.835–0.867). No significant difference in allelic richness or expected heterozygosity was observed among the seven populations. Pairwise F_ST estimates and NJ tree topologies based on D_C distances indicated that the seven populations fell into two groups, showing a clear division between the populations from the south and north of the Shandong Peninsula. Genetic differentiation was further analyzed using AMOVA and assignation tests. Genetic barrier analysis using Monmonier algorithm also confirmed that the Shandong Peninsula was the putative barrier separating the northern and southern populations. In addition, marine currents probably play an important role in high gene flow among three populations from the same marine gyre.     

Population genetic structure of the noxious weed Amaranthus retroflexus in Central Europe

Genetic variation was assessed in a range of populations of Amaranthus retroflexus using isoenzyme analysis. Population genetic diversity was measured by evaluating patterns of variation at six putatively neutral isoenzyme loci (comprising 24 putative alleles) within and among 20 populations of A. retroflexus collected in different habitats: ruderal habitats, cereal fields and hop gardens. Amaranthus retroflexus is a noxious weed of North American origin that infests various crops. Overall, A. retroflexus displayed moderate levels of genetic diversity in comparison with other herbaceous plants. The percentage of polymorphic loci was 50.0%, with mean values of 2.01, 0.142 and 0.227 for the average number of alleles per polymorphic locus (A), observed heterozygosity (H_o) and expected heterozygosity (H_e), respectively. A discrepancy between observed and expected heterozygosity and significant differences from H-W expectation indicate that there is an excess of homozygotes in many populations. As a result, there is strong evidence of inbreeding within populations (F_IS = 0.382) and significant population differentiation (F_ST = 0.270). Even though the species is partly autogamous, inbreeding does not lead to strong inbreeding depression resulting from self-pollination, as inbreeding has no effect on the success of the species in today's countryside. Moreover, allele frequencies detected in agricultural habitats (i.e., cereal fields and hop gardens) differed from those detected in populations collected from ruderal habitats, which is probably caused by systematic application of herbicides in agricultural ecosystems.     

Genetic diversity and population structure of the endangered and medically important Rheum tanguticum (Polygonaceae) revealed by SSR Markers

Rheum tanguticum (Polygonaceae), an endangered plant, is endemic to the Qinghai-Tibetan Plateau. A total of 114 individual of R. tanguticum from 10 geographically separate populations were analyzed using seven pairs simple sequence repeats (SSR) markers. 102 alleles were recorded, with an average of 14.6 alleles per locus (ranging from 13 to 17) and the expected heterozygosity (H_e) ranged from 0.384 to 0.515 (average 0.459). The genetic differentiation between populations was relatively high (F_st = 0.249); the gene flow (N_m = 0.754), however, was limited, which suggested that around 21.18% of the total genetic variations occurred between populations. Our results revealed high levels of genetic variations within and between populations. The endangered status of this species is probably due to harvesting of the wild populations, rather than a lack of the genetic diversity. Anthropologic effects as well as other factors may, together, have shaped the genetic structure of this species.     

High genetic structure of the Cozumel Harvest mice,a critically endangered island endemic: conservation implications

We assessed the genetic structure and diversity of Reithrodontomys spectabilis, a critically endangered, endemic rodent from Cozumel Island, México. A total of 90 individuals were trapped from September 2001 to January 2005. Microsatellite data analysis revealed high genetic diversity values: a total of 113 alleles (average 12.5 per locus), H _o  = 0.78, H _e  = 0.80. These high values can be related to Cozumel’s size (478 km²) and extensive native vegetation cover, factors that could be promoting a suitable population size, high heterozygosity and the persistence of rare alleles in the species, as well as some long-term movement of individuals between sampling localities. A strong genetic structure was also observed, with at least four genetic groups, associated with a pattern of isolation by distance. We found a strong allelic and genetic differentiation shown between localities, with negligible recent gene flow and low inbreeding coefficients. The species life history and ecological characteristics—being nocturnal, semi-terrestrial, a good tree climber, having lunar phobia and significant edge effect—are likely affecting its genetic structure and differentiation. The high genetic diversity and population structure award R. spectabilis a significant conservation value. Our results can serve as a basis for future research and conservation of the species, particularly considering the problems the island is facing from habitat perturbation, urbanization and introduction of exotic species. In view of the structure and genetic variability observed, it is essential to establish and reinforce protected areas and management programs for the conservation of the endemic and endangered Cozumel Harvest mice.     

Genetic diversity and local population structure in Ambrosina bassii (Araceae,Ambrosineae), a Mediterranean relict species

The effects of habitat fragmentation on the genetic structure of Ambrosina bassii are analyzed. The species, whose reproductive biology is mostly unknown, is the only representative of its genus and tribe and it is endemic to the central Mediterranean area. The selected study area was the island of Sicily, in which wild populations show a wide morphological variability and ecological amplitude. Patterns of within- and among-population genetic diversity in eleven Sicilian populations, occurring in six disjunct areas, were examined by means of allozyme electrophoresis. High levels of genetic diversity were found as shown by the mean expected heterozygosity (H_e = 0.263), the percentage of polymorphic loci (P₉₅ = 65.3), the mean number of alleles per locus (A = 2.0). Genetic differentiation between populations was relatively low (mean F_ST = 0.091 and Nm = 1.98). A very weak correlation exists between genetic distances and geographic distances between populations. Despite its restricted and fragmented geographical range, A. bassii showed (i) high levels of genetic diversity, mainly within populations; (ii) no genetic differentiation between populations and morphotypes.