The relationship between population genetic structure and pelagic larval duration in coral reef fishes on the Great Barrier Reef

Pelagic larval duration (PLD) is a commonly used proxy for dispersal potential in coral reef fishes. Here we examine the relationship between PLD, genetic structure and genetic variability in geographically widespread and ecological generalist species from one coral reef fish family (Pomacentridae) that differs in mean larval duration by more than a month. The genetic structure was estimated in eight species using a mitochondrial molecular marker (D-loop) and in a sub-set of five species using nuclear molecular markers (ISSRs). Estimates of genetic differentiation were similar among species with pelagic larvae, but differed between molecular markers. The mtDNA indicated no structure in all species except one, while the ISSR indicated some structure between the sampling locations in all species. We detected a relationship between PLD and genetic structure using both markers. These relationships, however, were caused by a single species, Acanthochromis polyacanthus, which differs from all the other species examined here in lacking a larval phase. With this species excluded, there was no relationship between PLD and genetic structure using either marker despite a range of PLDs of more than 20 days. Genetic diversities were generally high in all species and did not differ significantly among species and locations. Nucleotide diversity and total heterozygosity were negatively related to maximum PLD but again these relationships were caused by A. polyacanthus and disappeared when this species was excluded. These genetic patterns are consistent with moderate gene flow among well-connected locations and indicate that at this phylogenetic level (i.e., within family) the duration of the pelagic larval phase is unrelated to the patterns of genetic differentiation.     

A Hierarchical View of Genetic Structure in the Rare Annual Plant Clarkia springvillensis

Genetic structure at several spatial scales was examined in the rare California annual, Clarkia springvillensis . Using seven isozyme-encoding loci as genetic markers, we assessed the amount and distribution of genetic variation among three populations and eight subpopulations. Total genetic variation was lower than in species with similar life history traits but equivalent to that of other endemic plants. Spatial autocorrelation showed some evidence for very limited differentiation within subpopulations at a scale of 1–2 m. The subpopulations, separated by tens of meters, were found to be more differentiated from each other ( F _sp = 0.084) on average than were populations ( F,pt = 0.017). This local genetic differentiation was not correlated with physical distance between subpopulations. The low F_pt estimates suggest that substantial gene flow is occurring among populations. However, the lack of correlation between genetic and geographic distances and the significant differentiation of subpopulations suggest that genetic drift is occurring within populations. Therefore, we believe the apparent homogeneity of populations is due to each population's gene frequencies' being an average of several divergent subpopulations. If drift is causing differentiation within populations, it may eventually cause differentiation between populations. The importance of using a hierarchical approach to evaluating genetic structure is clear. Patterns occurring at one spatial scale may not be evident at others. One should not necessarily conclude that gene flow is substantial and that the risk of genetic erosion via drift is negligible just because differentiation between populations is small; the system may not be at equilibrium. This lesson is particularly important when recent changes in climate or land use are apparent.     

利用RAPD标记分析卧龙自然保护区不同海拔沙棘种群的遗传变异

从1800m到3400m五个海拔连续取样,用RAPD分子标记研究了卧龙自然保护区中国沙棘种群的遗传结构和遗传变异.用11条寡核苷酸引物,扩增得到151个重复性好的位点,其中143个多态位点,多态率达94.7%.在5个沙棘种群中,总遗传多样性值(HT)为0.289,B种群内的遗传多样性值为0.315,这完全符合沙棘这种多年生、远交的木本植物高遗传变异的特性.5个种群内遗传多样性随海拔升高呈低—高—低变异趋势,在2200m海拔处的B种群遗传多样性达最大值0.315,3400m海拔处的E种群则表现最小,仅0.098.5个种群间的遗传分化值GST=0.406,也即是说有40.6%的遗传变异存在于种群间,59.4%存在种群内.1800m海拔处的A种群与其他种群的明显分离是造成种群间遗传分化大的原因.UPGMA聚类图和PCoA散点图分别进一步确证了5个种群间关系和所有个体间的关系.最后,经过Mantel检测,遗传距离与海拔表现了明显的相关性(r=0.646,P=0.011).     

Latitudinal variability in spatial genetic structure in the invasive ascidian, Styela plicata

Increases in temperature can shorten planktonic larval durations, so that higher temperatures may reduce dispersal distances for many marine animals. To test this prediction, we first quantified how minimum time to settlement is shortened at higher temperatures for the ascidian Styela plicata. Second, using latitude as a correlate for ocean temperature and spatial genetic structure as a proxy for dispersal, we tested for a negative correlation between latitude and spatial genetic structure within populations, as measured by anonymous DNA markers. Spatial genetic structure was variable among latitudes, with significant structure at low and intermediate latitudes (high and medium temperatures) and there was no genetic structure within high-latitude (low temperature) populations. In addition, we found consistently high genetic diversity across all Australian populations, showing no evidence for recent local bottlenecks associated S. plicata’s history as an invasive species. There was, however, significant genetic differentiation between all populations indicating limited ongoing gene flow.     

Morphometric and electrophoretic analysis of 13 populations of Anatolian black pine in Turkey

The genetic variation in populations of Anatolian black pine (Pinus nigra Arn. subsp. pallasiana (L.) Holmboe.), one of the species covering large areas in Turkey, was investigated. Open pollinated seeds were collected from 13 populations in a natural distribution range. Six characters of seeds (length, width, ratio of length to width, weight/1000 seeds) and seedling characters (cotyledon number and hypocotyls height) and two enzyme systems viz. leucine aminopeptidase (LAP) and glutamate oxaloacetate transaminase, (GOT) were investigated. Significant differences were detected among the populations for the morphological characters. In addition, isozyme patterns of two enzyme systems revealed that LAP has two loci (one with 2 alleles and the other with 3), while GOT has three loci (two with 3 alleles and the third one with 2 alleles). Polymorphic loci were 74% on the average. The mean number of alleles per loci was 1.94 and expected heterozygosity was 19%. The mean total genetic diversity was calculated as 0.203; the mean gene diversity within populations was determined as 0.188, and the average between subpopulations diversity was 0.016. The relative magnitude of genetic differentiation among subpopulations was measured as 0.074 indicating that only 7.4% of the total genetic diversity was there between populations. Average genetic distance was 0.093 according to Gregorius. Nei's genetic distance was 0.022.     

Genetic Diversity, Population Size, and Fitness in Central and Peripheral Populations of a Rare Plant Lychnis viscaria

Abstract: Genetic diversity is expected to decrease in small and isolated populations as a consequence of bottlenecks, founder effects, inbreeding, and genetic drift. The genetics and ecology of the rare perennial plant Lychnis viscaria (Caryophyllaceae) were studied in both peripheral and central populations within its distribution area. We aimed to investigate the overall level of genetic diversity, its spatial distribution, and possible differences between peripheral and central populations by examining several populations with electrophoresis. Our results showed that the level of genetic diversity varied substantially among populations (  H _exp = 0.000–0.116) and that the total level of genetic diversity (mean H _exp = 0.056) was low compared to that of other species with similar life-history attributes. The peripheral populations of L. viscaria had less genetic variation (mean H _exp = 0.034) than the central ones (0.114). Analysis of genetic structure suggested limited gene flow (mean F _ST = 0.430) and high differentiation among populations, emphasizing the role of genetic drift (  N _e m = 0.33). Isolation was even higher than expected based on the physical distance among populations. We also focused on the association between population size and genetic diversity and possible effects on fitness of these factors. Population size was positively correlated with genetic diversity. Population size and genetic diversity, however, were not associated with fitness components such as germination rate, seedling mass, or seed yield. There were no differences in the measured fitness components between peripheral and central populations. Even though small and peripheral populations had lower levels of genetic variation, they were as viable as larger populations, which emphasizes their potential value for conservation.     

Influence of contrasting larval developmental types upon the population-genetic structure of cheilostome bryozoans

For many sedentary or sessile benthic marine invertebrates the planktonic duration of the larval stage is believed to be a key determinant of the magnitude of genetic differences between populations. An obvious dichotomy in dispersal potential exists within cheilostome bryozoans that develop from either (1) a cyphonautes larva that spends several weeks in the plankton or (2) a brooded coronate larva that settles soon after release from the adult colony. This study characterises the pattern of variation at allozyme loci among British populations of four species of bryozoan—two species with cyphonautes and two with coronate larvae. There is some variation in the estimates of genetic differentiation among similarly separated populations that may be a consequence of non-equilibrium genetic conditions arising from sporadic migration, possibly through dispersal by rafting on macroalgae by mature colonies. Despite this, however, both the level of genetic differentiation between populations and the pattern of migrant exchange correlate with the larval developmental mode. Bryozoan species that brood coronate larvae show higher levels of genetic heterogeneity between populations and significant isolation by distance genetic structure while, by contrast, distance has little or no effect upon the amount of genetic differentiation among populations of bryozoans with cyphonautes larvae. For cheilostome bryozoans, therefore, it appears that genetic differentiation between populations is directly associated with the type of larval development. These data are discussed also with respect to levels of gene diversity and the geological pattern of cheilostome bryozoan species diversity.     

Genetic Bottlenecks Driven by Population Disconnection

Abstract: Connectivity among populations plays a crucial role in maintaining genetic variation at a local scale, especially in small populations affected strongly by genetic drift. The negative consequences of population disconnection on allelic richness and gene diversity (heterozygosity) are well recognized and empirically established. It is not well recognized, however, that a sudden drop in local effective population size induced by such disconnection produces a temporary disequilibrium in allelic frequency distributions that is akin to the genetic signature of a demographic bottleneck. To document this effect, we used individual‐based simulations and empirical data on allelic richness and gene diversity in six pairs of isolated versus well‐connected (core) populations of European tree frogs. In our simulations, population disconnection depressed allelic richness more than heterozygosity and thus resulted in a temporary excess in gene diversity relative to mutation drift equilibrium (i.e., signature of a genetic bottleneck). We observed a similar excess in gene diversity in isolated populations of tree frogs. Our results show that population disconnection can create a genetic bottleneck in the absence of demographic collapse.     

Importance of dispersal routes that minimize open‐ocean movement to the genetic structure of island populations

Islands present a unique scenario in conservation biology, offering refuge yet imposing limitations on insular populations. The Kimberley region of northwestern Australia has more than 2500 islands that have recently come into focus as substantial conservation resources. It is therefore of great interest for managers to understand the driving forces of genetic structure of species within these island archipelagos. We used the ubiquitous bar‐shouldered skink (Ctenotus inornatus) as a model species to represent the influence of landscape factors on genetic structure across the Kimberley islands. On 41 islands and 4 mainland locations in a remote area of Australia, we genotyped individuals across 18 nuclear (microsatellite) markers. Measures of genetic differentiation and diversity were used in two complementary analyses. We used circuit theory and Mantel tests to examine the influence of the landscape matrix on population connectivity and linear regression and model selection based on Akaike's information criterion to investigate landscape controls on genetic diversity. Genetic differentiation between islands was best predicted with circuit‐theory models that accounted for the large difference in resistance to dispersal between land and ocean. In contrast, straight‐line distances were unrelated to either resistance distances or genetic differentiation. Instead, connectivity was determined by island‐hopping routes that allow organisms to minimize the distance of difficult ocean passages. Island populations of C. inornatus retained varying degrees of genetic diversity (N_A = 1.83 – 7.39), but it was greatest on islands closer to the mainland, in terms of resistance‐distance units. In contrast, genetic diversity was unrelated to island size. Our results highlight the potential for islands to contribute to both theoretical and applied conservation, provide strong evidence of the driving forces of population structure within undisturbed landscapes, and identify the islands most valuable for conservation based on their contributions to gene flow and genetic diversity.     

Roads,Interrupted Dispersal,and Genetic Diversity in Timber Rattlesnakes

Abstract: Anthropogenic habitat modification often creates barriers to animal movement, transforming formerly contiguous habitat into a patchwork of habitat islands with low connectivity. Roadways are a feature of most landscapes that can act as barriers or filters to migration among local populations. Even small and recently constructed roads can have a significant impact on population genetic structure of some species, but not others. We developed a research approach that combines fine‐scale molecular genetics with behavioral and ecological data to understand the impacts of roads on population structure and connectivity. We used microsatellite markers to characterize genetic variation within and among populations of timber rattlesnakes (Crotalus horridus) occupying communal hibernacula (dens) in regions bisected by roadways. We examined the impact of roads on seasonal migration, genetic diversity, and gene flow among populations. Snakes in hibernacula isolated by roads had significantly lower genetic diversity and higher genetic differentiation than snakes in hibernacula in contiguous habitat. Genetic‐assignment analyses revealed that interruption to seasonal migration was the mechanism underlying these patterns. Our results underscore the sizeable impact of roads on this species, despite their relatively recent construction at our study sites (7 to 10 generations of rattlesnakes), the utility of population genetics for studies of road ecology, and the need for mitigating effects of roads.     

Spatial and temporal genetic homogeneity in the Arctic surfclam (<Emphasis Type="Italic">Mactromeris polynyma</Emphasis>)

Commercially harvested marine bivalve populations show a broad range of population-genetic patterns that may be driven by planktonic larval dispersal (gene flow) or by historical (genetic drift) and ecological processes (selection). We characterized microsatellite genetic variation among populations and year classes of the commercially harvested Arctic surfclam, Mactromeris polynyma, in order to test the relative significance of gene flow and drift on three spatial scales: within commercially harvested populations in the northwest Atlantic; among Atlantic populations; and between the Atlantic and Pacific oceans. We found small nonsignificant genetic subdivision among eight populations from the northwest Atlantic (F _ST = 0.002). All of these Atlantic populations were highly significantly differentiated from a northeast Pacific population (F _ST = 0.087); all populations showed high inbreeding coefficients (F _IS = 0.432). We tested one likely source of heterozygote deficits by aging individual clams and exploring genetic variation among age classes within populations (a temporal Wahlund effect). Populations showed strikingly different patterns of age structure, but we found little differentiation among age classes. In one case, we were able to analyze genetic diversity between age classes older or younger than the advent of intensive commercial harvesting. The results generally suggest spatially broad and temporally persistent genetic homogeneity of these bivalves. We discuss the implications of the results for the biology and management of surfclam populations. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Genetic diversity and genetic structure in the brown alga Halidrys dioica (Fucales: Cystoseiraceae) in Southern California

Genetic diversity and genetic structure in a population of the brown seaweed Halidrys dioica Gardner were evaluated in five sites in southern California, USA, in 1991, using isozyme electrophoresis. H. dioica is relatively long-lived and has an outcrossing mating system and floating reproductive fronds with the potential for longdistance gamete dispersal. Because these characteristics are hypothetically important in determining genetic diversity and structure, we predicted that genetic diversity would be high and genetic structure would be exhibited only at relatively large geographic scales in H. dioica populations. The data were consistent with the prediction: genetic diversity (% polymorphic loci, no. of alleles/locus, average expected heterozygosity) was high compared to that of other seaweed species. Genetic structure (Wright's F statistics, Nei's genetic distance, point-pattern analysis of alleles) was low within and among distinct rocky reefs over 4 km of coast but high in subpopulations separated by 90 km. Life-history characteristics may be useful predictors of genetic diversity and structure in seaweed populations, but information on selection regimes, long-distance dispersal, and the extent of clonal propagation, for example, are critically lacking.     

麻疯树种质资源遗传多样性的ISSR分析

采用ISSR分子标记技术对大戟科属麻疯树9个自然居群的遗传多样性水平和遗传结构进行了研究.用10个引物对9个居群共135个样品进行了扩增,共获得169条清晰的扩增位点,其中多态性位点164个.POPGENE分析结果表明,麻疯树居群具有丰富的遗传多样水平(多态百分率PPB=97.04%,Neis遗传多样性H=0.2357,Shannon信息指数I=0.3760).AMOVA分析表明,9个自然居群间遗传多样性出现了较大的遗传分化(Gst=0.5398)可能与其有限的基因流(Nm=0.4667)和遗传漂变有关;而居群内有限的遗传分化可能是由于麻疯树自交、近交占主导方式的繁育方式有关.并进行了麻疯树居群的聚类分析.图3表4参19     

Genetic Structure of Fragmented Populations of the Endangered Daisy Rutidosis leptorrhynchoides

Abstract: The endangered grassland daisy Rutidosis leptorrhynchoides has been subject to severe habitat destruction and fragmentation over the past century. Using allozyme markers, we examined the genetic diversity and structure of 16 fragmented populations. The species had high genetic variation compared to other plant species, and both polymorphism and allelic richness showed strong positive relationships with log reproductive population size, reflecting a loss of rare alleles (frequency of q < 0.1) in smaller populations. Fixation coefficients were positively related to size, due either to a lack of rare homozygotes in small populations or to Wahlund effects (owing to spatial genetic structure) in large ones. Neither gene diversity nor heterozygosity was related to population size, and other population parameters such as density, spatial contagion, and isolation had no apparent effect on genetic variation. Genetic divergence among populations was low , despite a large north-to-south break in the species' current distribution. To preserve maximum genetic variation, conservation strategies should aim to maintain the five populations larger than 5000 reproductive plants, all of which occur in the north of the range, as well as the largest southern population of 626 plants at Truganina. Only one of these is currently under formal protection. High heterozygosity in smaller populations suggests that they are unlikely to be suffering from inbreeding depression and so are also valuable for conservation. Erosion of allelic richness at self-incompatibility loci, however, may limit the reproductive capacity of populations numbering less than 20 flowering plants.     

Mitochondrial DNA variation and population genetic structure of the blue crab<Emphasis Type="Italic"> Callinectes sapidus</Emphasis> in the eastern United States

The genetic population structure of red grouper, Epinephelus morio (Valenciennes), and scamp, Mycteroperca phenax Jordan and Swain, from the southeastern U.S. Atlantic coast and the Gulf of Mexico was examined using nuclear microsatellite DNA markers in order to test the null hypothesis of panmixia throughout this range. Physical and biological data indicate that relatively isolated populations of these fish exist. Genetic variation was assessed at four microsatellite loci in red grouper and six loci in scamp. The fish were collected on different dates between 1991 and 2001. The microsatellite loci were highly polymorphic, with an average expected heterozygosity of 0.75 in red grouper and 0.68 in scamp. Heterozygote deficiencies (significant deviations from Hardy–Weinberg equilibrium, HWE) were found at two of four loci in all red grouper samples except the eastern Gulf of Mexico, and for all red grouper combined. In contrast, all loci conformed to HWE in the separate scamp samples. Minimal genetic differences distinguished southeastern U.S. Atlantic or Mexican red grouper from other localities, and no indication of genetic differentiation was observed in scamp. This large-scale genetic homogeneity may be attributed to ongoing gene flow and/or historical contact between present-day populations. For management purposes, genetic homogeneity does not necessarily imply a single stock. Because larval dispersal may be sufficient to homogenize gene frequencies but not to replenish depleted stocks, other data must be considered in the management of these species.Electronic Supplementary Material Supplementary material is available in the online version of this article at .Communicated by J.P. Grassle, New Brunswick     

入侵植物加拿大一枝黄花和本地一枝黄花的遗传多样性比较

为了阐明加拿大一枝黄花成功入侵的机制,利用简单序列重复区间标记(ISSR)方法对加拿大一枝黄花和本地一枝黄花的遗传多样性进行比较研究。从100条引物中筛选出12条引物用于PCR扩增,利用POPGEN32软件对2种一枝黄花进行遗传多样性分析。结果表明,加拿大一枝黄花在物种水平上的多态位点百分率为95.19%,Nei’s基因多样性指数为0.308 5,Shannon’s信息指数为0.415 8;本地一枝黄花在物种水平上的多态位点百分率(89.80%)、Nei’s基因多样性指数(0.249 1)和Shannon’s信息指数(0.383 4)都比加拿大一枝黄花小。加拿大一枝黄花和本地一枝黄花居群间遗传分化系数分别为0.118 2和0.131 3,居群内变异分别为0.881 8和0.868 7,表明2个物种居群间的遗传分化不明显,遗传一致度高,且主要的遗传变异存在于居群内。入侵植物加拿大一枝黄花具有较高遗传多样性,且高于本地一枝黄花,这可能是加拿大一枝黄花成功入侵的原因之一。     

Local variation of within-host clonal diversity coupled with genetic homogeneity in a marine trematode

Despite their ubiquity and importance to intertidal ecosystems, information is currently lacking regarding the genetic diversity of trematode parasites within coastal organisms and the distribution of their genetic variation among intertidal habitats. In this study, we quantified the clonal diversity of the coastal marine trematode Maritrema novaezealandensis within Zeacumantus subcarinatus snail hosts from three coastal bays in Otago Harbour, New Zealand, using five microsatellite loci to determine if differences exist in the frequency of occurrence of multi-clone infections. In addition, we examined gene flow among M. novaezealandensis collected from the three bays. The frequency of mixed-clone infections varied fourfold among bays and no genetic differentiation was detected among intertidal bays. Across the coastal bays studied, M. novaezealandensis comprises a single population that is potentially infecting multiple Z. subcarinatus populations with varying life history traits.     

Gene flow among vestimentiferan tube worm (Riftia pachyptila) populations from hydrothermal vents of the eastern Pacific

Allozyme data are presented for six discrete populations of the giant hydrothermal vent tube worm Riftia pachyptila Jones, 1981 collected throughout the species' known range along mid-ocean spreading ridges of the eastern Pacific Ocean. Contrary to an earlier report, levels of genetic variation are relatively high in this species. Estimates of gene flow based on F-statistics revealed that dispersal throughout the surveyed region is sufficiently high to counter random processes that would lead to losses of genetic diversity and significant population differentiation. R. pachyptila, like other species of tube worms, displays considerable morphologic variation among populations, but this diversity is not reflected in allozyme variation. Vestimentifera, in general, appear to show extensive phenotypic plasticity. In the light of the available genetic data, caution is warranted when making inferences about the taxonomic status of collections based on morphological variation alone. A general decrease in estimated rates of gene flow between geographically more distant populations supports the hypothesis that dispersal in this species follows a stepping-stone model, with exchange between neighboring populations in great excess of long-distance dispersal. High levels of gene flow have been recorded in a variety of vent fauna and may be a prerequisite for success of species found in the ephemeral habitats associated with regions of sea-floor hydrothermal activity.     

High genetic diversities and complex genetic structure in an Indo-Pacific tropical reef fish (Chlorurus sordidus): evidence of an unstable evolutionary past?

Historical sea level fluctuations have influenced the genetic structure and evolutionary history of marine species and examining widespread species across their species ranges may elucidate some of these effects. Chlorurus sordidus is a common and widespread parrotfish found on coral reefs throughout the Indo-central Pacific. We used phylogenetic, phylogeographic, and cladistic analyses to examine the genetic composition and population structure of this species across most of its latitudinal range limits. We sequenced 354 bp of the mitochondrial control region I in 185 individuals from nine populations. Populations of C. sordidus displayed high levels of genetic diversity, similar to those recorded for widespread pelagic fish species, but much greater nucleotide diversity values than those previously recorded for other demersal reef fishes. Both phylogenetic and phylogeographic analyses detected strong genetic subdivision at the largest spatial scale (i.e. among oceans). The Pacific Ocean was characterised by weak population genetic structure. Separation of the Hawaiian location from other Pacific and West Indian Ocean sites was evident in phylogenetic analyses, but not from analysis of molecular variance. NCA and isolation-by-distance tests suggested that the genetic structure of this species was the result of multiple contemporary and historical processes, including long-distance colonisation and range expansion arising from fluctuating sea levels, limited current gene flow, and isolation by distance. This pattern is to be expected when historically fragmented populations come into secondary contact. We suggest the patterns of population genetic structure recorded in C. sordidus are caused by large local population sizes, high gene flow, and a recent history of repeated fragmentation and remixing of populations resulting from fluctuating sea levels.Communicated by M.S. Johnson, Crawley     

Conservation genetics of <Emphasis Type="Italic">Lumnitzera littorea</Emphasis> (Combretaceae), an endangered mangrove,from the Indo-West Pacific

Mangrove forests, with their ecological significance and economic benefits, are vital inter-tidal wetland ecosystems. Lumnitzera littorea (Combreataceae) is a non-viviparous mangrove distributed in tropical Asia and North Australia. Due to natural and human impacts, populations of this species have been isolated, fragmented, and highly disturbed. In China, L. littorea is an endangered species, restricted to small regions of Hainan Island. The genetic composition of five populations of this species from the Indo-West Pacific (South China, Malay Peninsula, Sri Lanka, North Australia) was assessed using inter simple sequence repeat (ISSR) makers. At the species level, expected mean heterozygosity (He) was 0.240 with 75.6% of loci polymorphic (P). However, genetic variation was much lower at the population level (P = 37.1%, He = 0.118). A high coefficient of gene differentiation (Gst = 0.515) and low level of gene flow (Nm = 0.470) indicated significant genetic differentiation among populations. AMOVA also indicated that more than half the total variation (58.4%) was partitioned among populations. The high degree of differentiation observed among populations emphasizes the need for appropriate conservation measures that incorporate additional populations into protected areas, and achieve the restoration of separate, degraded populations.