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1.

Background and Aims

Natural selection and genetic drift are important evolutionary forces in determining genetic and phenotypic differentiation in plant populations. The extent to which these two distinct evolutionary forces affect locally adaptive quantitative traits has been well studied in common plant and animal species. However, we know less about how quantitative traits respond to selection pressures and drift in endangered species that have small population sizes and fragmented distributions. To address this question, this study assessed the relative strengths of selection and genetic drift in shaping population differentiation of phenotypic traits in Psilopeganum sinense, a naturally rare and recently endangered plant species.

Methods

Population differentiation at five quantitative traits (QST) obtained from a common garden experiment was compared with differentiation at putatively neutral microsatellite markers (FST) in seven populations of P. sinense. QST estimates were derived using a Bayesian hierarchical variance component method.

Key Results

Trait-specific QST values were equal to or lower than FST. Neutral genetic diversity was not correlated with quantitative genetic variation within the populations of P. sinense.

Conclusions

Despite the prevalent empirical evidence for QST > FST, the results instead suggest a definitive role of stabilizing selection and drift leading to phenotypic differentiation among small populations. Three traits exhibited a significantly lower QST relative to FST, suggesting that populations of P. sinense might have experienced stabilizing selection for the same optimal phenotypes despite large geographical distances between populations and habitat fragmentation. For the other two traits, QST estimates were of the same magnitude as FST, indicating that divergence in these traits could have been achieved by genetic drift alone. The lack of correlation between molecular marker and quantitative genetic variation suggests that sophisticated considerations are required for the inference of conservation measures of P. sinense from neutral genetic markers.  相似文献   

2.
The impact of directional selection on specific trait types in plant species, and how a species’ life history mediates this response to selection remains understudied. Discovering such interactions is however crucial for understanding the interplay between ecological and genetic processes underlying local adaptation in plants, and to evaluate a species’ evolutionary potential with respect to changing environments. Furthermore, it remains unclear whether the degree of adaptive differentiation generally increases with the geographical distance between plant populations. Here, we present a weighted mixed model based meta-analysis aimed at unraveling the potential interactions between plant trait types, life history characteristics and QST–FST comparisons, and assessing the effect of geographical scale on population differentiation. Based on 51 studies we found that QST values exceeded their corresponding FST values in 71.74 % out of 401 cases. Furthermore, different trait types were found to be differently susceptible to natural selection and the magnitude of QST–FST comparisons was mediated by a plant species’ life span. These findings may be closely related to the genetic architectures of trait types and life histories, with the proportion of large-effect genes likely shaping the response to natural selection. QST–FST values also increased with increasing distance between populations, pinpointing the combined effects of environmental differentiation and isolation by distance on the magnitude of population divergence. Finally, our model showed an inverse relationship between FST and QST–FST values, presumably resulting from isolation by distance, the exchange of advantageous alleles, or genetic correlations among traits.  相似文献   

3.
The Irrawaddy dolphin (Orcaella brevirostris) is an endangered, small cetacean species which is widely distributed in rivers, estuaries, and coastal waters throughout the tropical and subtropical Indo-Pacific. Despite the extensive distribution of this species, little is known of individual movements or genetic exchange among regions in Thailand. Here, we evaluate the genetic diversity and genetic structure of O. brevirostris in the eastern, northern and western Gulf of Thailand, and Andaman Sea. Although phylogenetic relationships and network analysis based on 15 haplotypes obtained from 32 individuals reveal no obvious divergence, significant genetic differentiation in mitochondrial DNA (overall FST = 0.226, P < 0.001; ΦST = 0.252, P < 0.001) is apparent among regions. Of 18 tested microsatellite loci, 10 are polymorphic and successfully characterized in 28 individuals, revealing significant genetic differentiation (overall FST = 0.077, P < 0.05) among the four sampling sites. Structure analysis reveals two inferred genetic clusters. Additionally, Mantel analysis demonstrates individual-by-individual genetic distances and geographic distances follow an isolation-by-distance model. We speculate that the significant genetic structure of O. brevirostris in Thailand is associated with a combination of geographical distribution patterns, environmental and anthropogenic factors, and local adaptations.  相似文献   

4.
This study aimed to assess the population genetic structure of a widespread Neotropical tree species, Simarouba amara, at local, regional and continental spatial scales. We used five microsatellite loci to examine genetic variation in 14 natural populations (N?=?478 individuals) of this vertebrate-dispersed rain forest tree species in Panama, Ecuador, and French Guiana. Estimates of genetic differentiation (F st and R st) were significant among all but one population pair and global differentiation was moderate (F st?=?0.25, R st?=?0.33) with 94% of genetic variation ascribed to differences among three main geographic regions (Central America, Western Ecuador, Amazon basin). There was no evidence of isolation by distance within regions. Allele-size mutations contributed significantly (R ST > F ST) to the divergences between cis- and trans-Andean populations, highlighting the role of the northern Andean cordilleras as an important geographic barrier for this species.  相似文献   

5.
This study of population structure of two intertidal monodontine species: Diloma nigerrima and Diloma subrostrata, revealed the level of genetic connectedness among populations. Despite their markedly different geographic ranges (D. nigerrima is a geographically widespread species, inhabiting both Chile and New Zealand, including its subantarctic islands, whereas D. subrostrata is endemic to New Zealand), both species are believed to possess short-lived lecithotrophic larval stages. Polymorphic DNA microsatellite sequences were used to reveal the level of genetic connectedness among populations, thus inferring the two species' relative effective dispersal abilities. For each species F statistics, AMOVA values and the strength of the relationship between geographic and genetic distance were calculated. We observed a higher within-species level of genetic variation (ΦST = 0.099 vs. ΦST = 0.016) and a higher proportion of variance (11.15% vs. 0%) among populations of D. nigerrima than of D. subrostrata. A larger fraction of significant FST values was observed among D. nigerrima population pairs (65%) than among D. subrostrata population pairs (33%). Significant correlation between genetic and geographic distance was observed for D. nigerrima but not for D. subrostrata, but this relationship was not consistent among pairwise D. nigerrima population comparisons and PCA analysis confirms that, for each species, population structure does not follow a consistent pattern of increasing with geographic distance. The lack of population structure among D. subrostrata populations is probably due to its ubiquitous distribution, meaning little opportunity exists for genetic structure. D. nigerrima, by contrast has a patchier distribution, which allows for greater opportunities for genetic differentiation to occur. We argue that, despite the probable short larval stage in this species, the lack of geographical pattern in the genetic structure found in D. nigerrima is best explained by occasional dispersal over relatively short distances around the coast of New Zealand, over longer distances from New Zealand to the subantarctic islands and even across the Pacific Ocean from New Zealand to Chile.  相似文献   

6.
Aim Natural and human‐induced differences in frugivore assemblages can influence the seed dispersal distances of trees. An important issue in seed dispersal systems is to understand whether differences in seed dispersal distances also affect the genetic structure of mature trees. One possible approach to test for a relationship between seed dispersal and the genetic structure of mature trees is to compare the genetic structure of two closely related tree species between two biogeographical regions that differ in frugivore assemblages and seed dispersal distances. Previous studies on two Commiphora species revealed that Commiphora guillauminii in Madagascar has a much lower seed dispersal distance than Commiphora harveyi in South Africa. We tested whether the lower seed dispersal distance might have caused decreased gene flow, resulting in a stronger genetic structure in Madagascar than in South Africa. Location Madagascar and South Africa. Methods Using amplified fragment length polymorphism markers we investigated the genetic structure of 134 trees in Madagascar and 158 trees in South Africa at a local and a regional spatial scale. Results In concordance with our hypothesis, kinship analysis suggests that gene flow was restricted mostly to 3 km in Madagascar and to 30 km in South Africa. At the local spatial scale, the genetic differentiation among groups of trees within sample sites was marginally significantly higher in Madagascar (FST = 0.069) than in South Africa (FST = 0.021). However, at a regional spatial scale genetic differentiation was lower in Madagascar (FST = 0.053) than in South Africa (FST = 0.163). Main conclusions Our results show that lower seed dispersal distances of trees were linked to higher genetic differentiation of trees only at a local spatial scale. This suggests that seed dispersal affects the genetic population structure of trees at a local, but not at a regional, spatial scale.  相似文献   

7.
In landscape genetics, isolation-by-distance (IBD) is regarded as a baseline pattern that is obtained without additional effects of landscape elements on gene flow. However, the configuration of suitable habitat patches determines deme topology, which in turn should affect rates of gene flow. IBD patterns can be characterized either by monotonically increasing pairwise genetic differentiation (for example, FST) with increasing interdeme geographic distance (case-I pattern) or by monotonically increasing pairwise genetic differentiation up to a certain geographical distance beyond which no correlation is detectable anymore (case-IV pattern). We investigated if landscape configuration influenced the rate at which a case-IV pattern changed to a case-I pattern. We also determined at what interdeme distance the highest correlation was measured between genetic differentiation and geographic distance and whether this distance corresponded to the maximum migration distance. We set up a population genetic simulation study and assessed the development of IBD patterns for several habitat configurations and maximum migration distances. We show that the rate and likelihood of the transition of case-IV to case-I FST–distance relationships was strongly influenced by habitat configuration and maximum migration distance. We also found that the maximum correlation between genetic differentiation and geographic distance was not related to the maximum migration distance and was measured across all deme pairs in a case-I pattern and, for a case-IV pattern, at the distance where the FST–distance curve flattens out. We argue that in landscape genetics, separate analyses should be performed to either assess IBD or the landscape effects on gene flow.  相似文献   

8.
Population genetic studies of ectomycorrhizal fungi at scales larger than 100 km are still relatively rare with highly variable results. In this study, we determined the population genetic structure of the ectomycorrhizal basidiomycete Suillus spraguei over distances up to 600 km in northeastern USA forests. S. spraguei associates exclusively with five-needled pines and only with white pine (Pinus strobus) in the eastern USA. We used six microsatellite loci to assess the genetic structure between eight sites sampled in the Adirondack Park of New York and seven sites sampled in other forests of New York, Pennsylvania, and Massachusetts. Except for one site, little to no genetic differentiation was detected in pairwise comparisons of the sites (F ST?=?0 to 0.05). Only one site was moderately differentiated from most other sites (F ST?=?0.02 to 0.15). The Mantel test showed no significant correlation between genetic and geographic distances (isolation by distance; R 2?=?0.003, P?=?0.3). The STRUCTURE analysis also supported the presence of a single cluster (K?=?1).  相似文献   

9.
Limited dispersal is commonly used to explain differences in diversification rates. An obvious but unexplored factor affecting dispersal is the mode of locomotion used by animals. Whether individuals walk, swim or fly can dictate the type and severity of geographical barriers to dispersal, and determine the general range over which genetic differentiation might occur. We collated information on locomotion mode and genetic differentiation (FST) among vertebrate populations from over 400 published articles. Our results showed that vertebrate species that walk tend to have higher genetic differentiation among populations than species that swim or fly. Within species that swim, vertebrates in freshwater systems have higher genetic differentiation than those in marine systems, which is consistent with the higher number of species in freshwater environments. These results show that locomotion mode can impact gene flow among populations, supporting at a broad‐scale what has previously been proposed at smaller taxonomical scales.  相似文献   

10.
Genetic variation was assessed in Atlantic wolffish, Anarhichas lupus, across the North Atlantic Ocean using microsatellite and amplified fragment length polymorphism (AFLP) markers. Despite unusual life history attributes such as large benthic eggs, large larvae, a limited pelagic stage and relatively sedentary adults, which suggest potential for strong population structure, range‐wide FST values were comparable to other marine fishes (≤0.035). Nevertheless, both significant genetic differentiation among regions and isolation by distance were observed, suggesting limited dispersal in this species. AFLP loci, evaluated on a subset of samples, revealed slightly higher FST values, but similar patterns of differentiation and isolation‐by‐distance estimates, compared to microsatellites. The genetic structure of Atlantic wolffish has likely been shaped by its post‐glacial history of recolonization, North Atlantic current patterns and continuity of habitat on continental shelves.  相似文献   

11.
Populus trichocarpa is an ecologically important tree across western North America. We used a large population sample of 498 accessions over a wide geographical area genotyped with a 34K Populus SNP array to quantify geographical patterns of genetic variation in this species (landscape genomics). We present evidence that three processes contribute to the observed patterns: (1) introgression from the sister species P. balsamifera, (2) isolation by distance (IBD), and (3) natural selection. Introgression was detected only at the margins of the species’ distribution. IBD was significant across the sampled area as a whole, but no evidence of restricted gene flow was detected in a core of drainages from southern British Columbia (BC). We identified a large number of FST outliers. Gene Ontology analyses revealed that FST outliers are overrepresented in genes involved in circadian rhythm and response to red/far‐red light when the entire dataset is considered, whereas in southern BC heat response genes are overrepresented. We also identified strong correlations between geoclimate variables and allele frequencies at FST outlier loci that provide clues regarding the selective pressures acting at these loci.  相似文献   

12.
Studies linking genetic structure in amphibian species with ecological characteristics have focused on large differences in dispersal capabilities. Here, we test whether two species with similar dispersal potential but subtle differences in other ecological characteristics also exhibit strong differences in genetic structure in the same landscape. We examined eight microsatellites in marbled salamanders (Ambystoma opacum) from 29 seasonal ponds and spotted salamanders (Ambystoma maculatum) from 19 seasonal ponds in a single geographic region in west-central Massachusetts. Despite overall similarity in ecological characteristics of spotted and marbled salamanders, we observed clear differences in the genetic structure of these two species. For marbled salamanders, we observed strong overall genetic differentiation (F ST = 0.091, F′ ST = 0.375), three population-level clusters of populations (K = 3), a strong pattern of isolation by distance (r = 0.58), and marked variation in family-level structure (from 1 to 23 full-sibling families per site). For spotted salamanders, overall genetic differentiation was weaker (F ST = 0.025, F′ ST = 0.102), there was no evidence of population-level clustering (K = 1), the pattern of isolation by distance (r = 0.17) was much weaker compared to marbled salamanders, and there was less variation in family-level structure (from 10 to 36 full-sibling families per site). We suspect that a combination of breeding site fidelity, effective population size, and generation interval is responsible for these marked differences. Our results suggest that marbled salamanders, compared to spotted salamanders, are more sensitive to fragmentation from various land-use activities and would be less likely to recolonize extirpated sites on an ecologically and conservation-relevant time frame.  相似文献   

13.
Xu J H  Wang L L  Xue H L  Wang Y S  Xu L X 《农业工程》2009,29(5):314-319
In order to verify the effect of social behavior and geographical isolation on the genetic structure of the Himalayan marmot (Marmota himalayana) population, we examined the genetic diversity of Himalayan marmots alongside the Qinghai–Tibet Railway using microsatellite markers. Eight microsatellite loci were used to examine 120 animals of 4 populations: Ulan (U), Delhi (D), Tuotuohe (T) and Ando (A). The results show that: (1) Himalayan marmots alongside the Qinghai–Tibet Railway are highly genetically diversified. The allele number (Na), effective allele number (Ne), observed heterozygosity (Ho), Nei’s expected heterozygosity (He) and polymorphism information content (PIC) of the total Himalayan marmot population were 4.75, 3.0332, 0.6990, 0.6672, 0.6102, respectively. (2) Himalayan marmots may be able to avoid inbreeding by a mechanism that will prevent the genetic diversity reduction caused by their social lifestyle. Heterozygote excess was observed at most loci. The inbreeding coefficients within the subpopulation (FIS), in the total population (FIT), the differentiation index of population (FST), and the gene flow (Nm) were ?0.2265, ?0.0477, 0.1458, and 1.4646, respectively. (3) The genetic differentiation of the Himalayan marmot population was in accordance with Wright’s “isolation by distance” theory. The Mantel test indicates that the correlation between genetic distance and geographic distance was significant (P < 0.05, r = 0.698). (4) Each of the four geographical populations had moderate differentiation. Both geographic distance and isolation could affect the population genetic structure of the Himalayan marmot. The maximum gene flow (3.5915), the smallest genetic differentiation index (0.0651), the lowest genetic distance (0.0700) and the highest genetic identity (0.9526) were all between the Ulan population and Delhi populations. (5) The cluster analysis, based on Nei’s standard genetic distance, showed that the populations of Delhi and Ulan were first merged in a cluster, and then Tuotuohe population was merged in the clustering. The Ando population was the last element in the clustering.  相似文献   

14.
Many marine gastropods are sedentary as adults but have planktonic larvae which can potentially be dispersed over large distances. Consequently larval transport is expected to play a prominent role in facilitating gene flow and determining population structure. The spotted babylon (Babylonia areolata) is a dioecious species possessing an approximately two week planktonic larval stage. We analyzed the population structure of the spotted babylon using amplified fragment length polymorphism (AFLP). One hundred and sixteen AFLP loci were analyzed in 63 individuals from three populations and revealed a high level of genetic diversity, with all individuals harboring a unique banding pattern. AMOVA results and an assignment test revealed that population differentiation was present. PCoA, pairwise FST and UPGMA tree all revealed that gene flow might be present only on a small geographic scale (around 160 km), but, over a large distance (around 1000 km), only reduced gene flow occurred. A mantel test indicated a highly significant positive correlation between genetic differentiation and geographical distance.  相似文献   

15.
Knowledge of genetic structure and patterns of connectivity is valuable for implementation of effective conservation management. The arid zone of Australia contains a rich biodiversity, however this has come under threat due to activities such as altered fire regimes, grazing and the introduction of feral herbivores and predators. Suitable habitats for many species can be separated by vast distances, and despite an apparent lack of current geographical barriers to dispersal, habitat specialisation, which is exhibited by many desert species, may limit connectivity throughout this expansive region. We characterised the genetic structure and differentiation of the great desert skink (Liopholis kintorei), which has a patchy, but widespread distribution in the western region of the Australian arid zone. As a species of cultural importance to local Aboriginal groups and nationally listed as Vulnerable, it is a conservation priority for numerous land managers in central Australia. Analysis of mitochondrial ND4 sequence data and ten nuclear microsatellite loci across six sampling localities through the distribution of L. kintorei revealed considerable differentiation among sites, with mitochondrial FST and microsatellite F′ST ranging from 0.047-0.938 and 0.257-0.440, respectively. The extent of differentiation suggests three main regions that should be managed separately, in particular the southeastern locality of Uluru. Current genetic delineation of these regions should be maintained if future intervention such as translocation or captive breeding is to be undertaken.  相似文献   

16.
We investigated the gene flow of the common marine diatom, Skeletonema marinoi, in Scandinavian waters and tested the null hypothesis of panmixia. Sediment samples were collected from the Danish Straits, Kattegat and Skagerrak. Individual strains were established from germinated resting stages. A total of 350 individuals were genotyped by eight microsatellite markers. Conventional F-statistics showed significant differentiation between the samples. We therefore investigated whether the genetic structure could be explained using genetic models based on isolation by distance (IBD) or by oceanographic connectivity. Patterns of oceanographic circulation are seasonally dependent and therefore we estimated how well local oceanographic connectivity explains gene flow month by month. We found no significant relationship between genetic differentiation and geographical distance. Instead, the genetic structure of this dominant marine primary producer is best explained by local oceanographic connectivity promoting gene flow in a primarily south to north direction throughout the year. Oceanographic data were consistent with the significant FST values between several pairs of samples. Because even a small amount of genetic exchange prevents the accumulation of genetic differences in F-statistics, we hypothesize that local retention at each sample site, possibly as resting stages, is an important component in explaining the observed genetic structure.  相似文献   

17.
Environmental variables that are correlated with depth have been suggested to be among the major forces underlying speciation in the deep sea. This study incorporated phylogenetics and ecological niche models (ENM) to examine whether congeneric species of Callogorgia (Octocorallia: Primnoidae) occupy different ecological niches across the continental slope of the Gulf of Mexico (GoM) and whether this niche divergence could be important in the evolution of these closely related species. Callogorgia americana americana, Callogorgia americana delta and Callogorgia gracilis were documented at 13 sites in the GoM (250–1000 m) from specimen collections and extensive video observations. On a first order, these species were separated by depth, with C. gracilis occurring at the shallowest sites, C. a. americana at mid‐depths and C. a. delta at the deepest sites. Callogorgia a. delta was associated with areas of increased seep activity, whereas C. gracilis and C. a. americana were associated with narrow, yet warmer, temperature ranges and did not occur near cold seeps. ENM background and identity tests revealed little to no overlap in ecological niches between species. Temporal calibration of the phylogeny revealed the formation of the Isthmus of Panama was a vicariance event that may explain some of the patterns of speciation within this genus. These results elucidate the potential mechanisms for speciation in the deep sea, emphasizing both bathymetric speciation and vicariance events in the evolution of a genus across multiple regions.  相似文献   

18.
Aim To characterize the genetic structure and diversity of Pinus cembra L. populations native to two disjunct geographical areas, the Alps and the Carpathians, and to evaluate the rate of genetic differentiation among populations. Location The Swiss Alps and the Carpathians. Methods We screened 28 populations at three paternally inherited chloroplast simple sequence repeats (cpSSRs) for length variation in their mononucleotide repeats. Statistical analysis assessed haplotypic variation and fixation indices. Hierarchical analysis of molecular variance (AMOVA), Mantel test, spatial analysis of molecular variance (SAMOVA) and barrier analyses were applied to evaluate the geographical partitioning of genetic diversity across the species’ range. Results Haplotypic diversity was generally high throughout the natural range of P. cembra, with the mean value substantially higher in the Carpathians (H = 0.53) than in the Alps (H = 0.35). The isolated Carpathian populations showed the highest haplotype diversity among the populations originating from the High Tatras (Velka Studena Dolina) and South Carpathians (Retezat Mountains). AMOVA revealed that only 3% of the total genetic variation derived from genetic differentiation between the two mountain ranges. Differentiation among Carpathian populations was higher (FST = 0.19) than among Alpine populations (FST = 0.04). Low, but significant, correlation was found between the geographical and genetic distances among pairs of populations (r = 0.286, P < 0.001). SAMOVA results revealed no evident geographical structure of populations. barrier analysis showed the strongest differentiation in the eastern part of the species’ range, i.e. in the Carpathians. Main conclusions The populations of P. cembra within the two parts of the species’ range still share many cpDNA haplotypes, suggesting a common gene pool conserved from a previously large, continuous distribution range. Carpathian populations have maintained high haplotypic variation, even higher than Alpine populations, despite their small population sizes and spatial isolation. Based on our results, we emphasize the importance of the Carpathian populations of Swiss stone pine for conservation. These populations comprise private haplotypes and they may represent a particular legacy of the species’ evolutionary history.  相似文献   

19.
The analysis of geographical patterns in population divergence has always been a powerful way to infer microevolutionary processes involved in population differentiation, and several approaches have been used to investigate such patterns. Most frequently, multivariate spatial patterns of population differentiation are analyzed by computing pairwise genetic distances or FST (or related statistics, such as ?ST from AMOVA), which are then correlated with geographical distances or landscape features. However, when calculating distances, especially based on presence-absence of alleles in local populations, there would be a confounding effect of allelic richness differences in the population differentiation. Moreover, the relative magnitude of these components and their spatial patterns can help identifying microevolutionary processes driving population differentiation. Here we show how recent methodological advances in ecological community analyses that allows partitioning dissimilarity into turnover (turnover) and richness differences, or nestedness-resultant dissimilarity, can be applied to allelic variation data, using an endemic Cerrado tree (Dipteryx alata) as a case study. Individuals from 15 local populations were genotyped for eight microsatellite loci, and pairwise dissimilarities were computed based on presence-absence of alleles. The turnover of alleles among populations represented 69?% of variation in dissimilarity, but only the richness difference component shows a clear spatial structure, appearing as a westward decrease of allelic richness. We show that decoupling richness difference and turnover components of allelic variation reveals more clearly how similarity among populations reflects geographical patterns in allelic diversity that can be interpreted in respect to historical range expansion in the species.  相似文献   

20.
Six south Florida populations of the endangered red-cockaded woodpecker (Picoides borealis) were sampled to examine genetic diversity and population structure in the southernmost portion of the species' range relative to 14 previously sampled populations from throughout the species range. Random amplified polymorphic DNA (RAPD) analyses were used to evaluate the populations (n= 161 individuals, 13 primers, one band/primer). Results suggested that south Florida populations have significant among-population genetic differentiation (FST= 0.17, P < 0.000), although gene flow may be adequate to offset drift (Nm= 1.26). Comparison of Florida populations with others sampled indicated differentiation was less in Florida (FST for all populations = 0.21). Cluster analyses of all 20 populations did not reflect complete geographical predictions, although clustering of distant populations resulted in a significant correlation between genetic distance and geographical distance. Overall, results suggest populations in south Florida, similar to the remainder of the species, have low genetic diversity and high population fragmentation. Exact clustering of distant populations supports the ability of RAPDs to differentiate populations accurately. Our results further support past management recommendations that translocations of birds among geographically proximate populations is preferable to movement of birds between distant populations.  相似文献   

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