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1.
The breeding system of a rare Great Lakes endemic, the lakeside daisy ( Hymenoxys acaulis var. glabra ), was investigated when plants from a remnant Illinois population produced no seeds for over 15 years. To determine if the Lakeside daisy was self-incompatible, 20 plants from two populations, Illinois and Ohio, were selfed and outcrossed. Seed/ovule ratios were compared among the different treatments and the location of the incompatibility reaction was identified. Lakeside daisy was found to be self-incompatible (sporophytic). The last Illinois population was effectively extinct because the remaining plants belonged to the same mating type ( N e = 1) and only produced seeds when outcrossed to the Ohio plants. Cross-incompatibility was also observed among Ohio plants, suggesting that within large populations, compatible mating types may be rare locally. In addition, inbreeding depression (lower seed/ovule ratios in inbred than in outcrosses) was observed after one generation of inbreeding. Small populations of self-incompatible species are vulnerable to extinction if the number of self-incompatibility alleles, either as a result of a bottleneck or of genetic drift, falls below tbe number needed for the breeding system to function. Recovery protocols based on these genetic considerations were developed and implemented in 1988 when Lakeside daisy populations were established at three Illinois nature preserves.  相似文献   

2.
Abstract:  Many populations have recovered from severe bottlenecks either naturally or through intensive conservation management. In the past, however, few conservation programs have monitored the genetic health of recovering populations. We conducted a conservation genetic assessment of a small, reintroduced population of Mauritius Kestrel ( Falco punctatus ) to determine whether genetic deterioration has occurred since its reintroduction. We used pedigree analysis that partially accounted for individuals of unknown origin to document that (1) inbreeding occurred frequently (2.6% increase per generation; N eI= 18.9), (2) 25% of breeding pairs were composed of either closely or moderately related individuals, (3) genetic diversity has been lost from the population (1.6% loss per generation; N eV= 32.1) less rapidly than the corresponding increase in inbreeding, and (4) ignoring the contribution of unknown individuals to a pedigree will bias the metrics derived from that pedigree, ultimately obscuring the prevailing genetic dynamics. The rates of inbreeding and loss of genetic variation in the subpopulation of Mauritius Kestrel we examined were extreme and among the highest yet documented in a wild vertebrate population. Thus, genetic deterioration may affect this population's long-term viability. Remedial conservation strategies are needed to reduce the impact of inbreeding and loss of genetic variation in this species. We suggest that schemes to monitor genetic variation after reintroduction should be an integral component of endangered species recovery programs.  相似文献   

3.
Abstract:  The area of Caricion davallianae alliance in Switzerland has been considerably reduced and fragmented during the last 150 years. We assessed the genetic variability, inbreeding level, and among-population differentiation of two common habitat-specific plant species, Carex davalliana SM. and Succisa pratensis Moench, in 18 Caricion davallianae fen meadows subjected to fragmentation. We used a spatial field design of fen systems (six systems total), each consisting of one large habitat island and two small habitat islands. We used allozyme electrophoresis to derive standard genetic parameters ( A, P, HO, HE, FIS, FST ). In Carex we identified a consistently lower A in isolated habitat islands; furthermore, HE was lower in small habitat islands than in large habitat islands. In Succisa we identified a lower HO in small habitat islands than in larger ones. Small habitat islands were marginally significantly differentiated (  FST ) from large islands for Succisa . For both species, no effects were evident for FIS ; therefore, we argue that genetic drift rather than inbreeding is the main cause of the observed differences. The genetic structure of Carex and Succisa in small habitat islands differed from that in large habitat islands, but differences were small. It appears that the observed differences in genetic variability among fen meadows correspond to observed differences in fitness and demographic traits. We show that habitat fragmentation affects not only the rare species in an ecosystem but also reduces the survival probabilities of common species. One of the main goals of conservation should be to mitigate fragmentation of natural habitats in order to increase population sizes and connectivity.  相似文献   

4.
Abstract: We used microsatellite DNA markers to investigate the maintenance of genetic diversity within and between samples of subpopulations (spanning five captive-bred generations) of the haplochromine cichlid Prognathochromis perrieri . The subpopulations are maintained as part of the Lake Victoria Cichlid species survival plan. Changes in the frequencies of 24 alleles, over four polymorphic loci, were used to estimate effective population size (   N e   ). Point estimates of N e ranged from 2.5 to 7.7 individuals and were significantly smaller than the actual census size (   N obs  ) for all subpopulations (32–243 individuals per generation), with the corresponding conservative N e   /  N obs ratios ranging from 0.01 to 0.12. Approximately 19% of the initial alleles were lost within the first four generations of captive breeding. Between-generation comparisons of expected heterozygosity showed significant losses ranging from 6% to 12% per generation. Seven private alleles were observed in the last sampled generation of four subpopulations, and analysis of population structure by F ST indicated that approximately 33% of the total genetic diversity is maintained between the subpopulations from different institutions. To reduce the loss of genetic variation, we recommend that offspring production be equalized by periodically removing dominant males, which will encourage reproduction by additional males. Consideration should also be given to encouraging more institutions to maintain populations, because a significant fraction of the genetic variation exists as among-population differences resulting from random differentiation among subpopulations.  相似文献   

5.
Abstract:  To study the relative importance of inbreeding depression and the loss of adaptive diversity in determining the extinction risk of small populations, we carried out an experiment in which we crossed and self-fertilized founder plants from a single, large population of shore campion ( Silene littorea Brot.). We used the seeds these plants produced to colonize 18 new locations within the distribution area of the species. The reintroduced populations were of three kinds: inbred and genetically homogeneous, each made up of selfed seed from a single plant; inbred and mixed, made up of a mixture of selfed seeds from all founder plants; and outbred and mixed, made up of a mixture of seeds obtained in outcrosses between the founders. We compared the inbred homogeneous populations with the inbred mixed to measure the effect of genetic diversity among individuals and the inbred mixed with the outbred mixed to measure the effect of inbreeding. Reintroduction success was seriously limited by inbreeding, whereas it was not affected by genetic diversity. This observation and the nonsignificant interaction between family and reintroduction location for individual plant characters suggest that the fixation of overall deleterious genes causing inbreeding depression posed a more serious threat to the short-term survival of the populations than the loss of genes involved in genotype and environment interactions. Thus, reintroduction success was related to adaptive diversity. Preventing such fixation might be the most important consideration in the genetic management and conservation of shore campion populations.  相似文献   

6.
Abstract:  Endangered species are commonly found in several (partially) isolated populations dispersed on different fragments of a habitat, natural reserve, or zoo. A certain level of connectivity among such populations is essential for maintaining genetic variation within and between populations to allow local and global adaptation and for preventing inbreeding depression. A rule of thumb widely accepted by the conservation community is that one migrant per generation (OMPG) into a population is the appropriate level of gene flow. This rule is based on Wright's study of his island model under a long list of simplifying assumptions. I examined the robustness of the OMPG rule to the violation of each of the many assumptions, quantifying the effect with population genetics theory. I showed that, when interpreted as one effective migrant per generation, OMPG is generally valid for real populations departing from the ideal model in the discrepancies of actual (  N ) and effective (  Ne  ) population sizes and actual ( m ) and effective ( me  ) migration rates. I also addressed the issue of converting the effective number of migrants (  Me= Neme  ) into the actual number of migrants ( M = Nm  ) of a certain age and sex. In particular, Ne < N , a case common for natural populations, did not necessarily require M > Me to maintain a certain level of differentiation among populations. Rather, translating the elusive Me into the manageable M depends on the specific causes (e.g., biased sex ratio, reproductive skew) that lead to Ne < N .  相似文献   

7.
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.  相似文献   

8.
Abstract: Populations of the endangered Rutidosis leptorrhynchoides (Asteraceae) have been reestablished in conservation reserves to reduce their risk of extinction in the wild. The reproductive success (number of seeds per inflorescence, percent seed set, germinability) of five small reestablished populations (at 5–10 years after establishment) was compared to that of two large natural remnant populations from which they were derived. Initial growth rates under glasshouse conditions were compared for seedlings derived from seed from remnant and reestablished populations. Seed set per inflorescence in all five reestablished populations was equal to or greater than seed set in remnant population plants. The resulting seed was as germinable, and in some cases more germinable, than seed derived from remnant populations; the seedlings then grew as large or larger than remnant population seedlings grown under glasshouse conditions. Fitness reductions in small reestablished populations of R. leptorrhynchoides are not evident at the reproductive stage. Hence, these populations have some potential to maintain natural regeneration processes and might therefore positively contribute to the conservation of this species.  相似文献   

9.
Hong Kong once supported more than 109 species of wild orchids, of which approximately 30% were endemic. Most of the local wild orchids have now become rare or endangered. I conducted a comparative study of genetic diversity in two closely related terrestrial orchids, an allotetraploid, Spiranthes hongkongensis , and its diploid progenitor, S. sinensis , to assess the effects of the population bottleneck associated with the origin of the polyploid and to investigate the relationships between number of breeding individuals, mating system, and level of isozyme variation in their populations. Nearly complete genetic uniformity was observed both within and among populations of S. hongkongensis . In contrast, S. sinensis had high levels of genetic variation for all of the genetic parameters examined. Regression analysis of population size and several components of genetic diversity in S. sinensis revealed that, among various measures of within-population variation, the proportion of polymorphic loci ( P ) and average number of alleles per locus ( A ) or per polymorphic locus ( A p ) were the most sensitive to population size ( R 2 = 0.942, p = 0.001; R 2 = 0.932, p = 0.002; and R 2 = 0.923, p = 0.002 respectively). The highly negative correlation ( r = −0.999, p < 0.01) between population size and the mean frequency of private alleles in pairwise population comparisons, p (1), indicated that population size may also be used to predict the extent of population differentiation caused by random genetic drift. Conservation of genetic diversity in S. sinensis could be maximized by protecting several of both large and small populations, whereas fewer populations may be needed to achieve this goal for S. hongkongensis.  相似文献   

10.
Introduced Birds and the Fate of Hawaiian Rainforests   总被引:3,自引:0,他引:3  
Abstract:  The Hawaiian Islands have lost nearly all their native seed dispersers, but have gained many frugivorous birds and fleshy-fruited plants through introductions. Introduced birds may not only aid invasions of exotic plants but also may be the sole dispersers of native plants. We assessed seed dispersal at the ecotone between native- and exotic-dominated forests and quantified bird diets, seed rain from defecated seeds, and plant distributions. Introduced birds were the primary dispersers of native seeds into exotic-dominated forests, which may have enabled six native understory plant species to become reestablished. Some native plant species are now as common in exotic forest understory as they are in native forest. Introduced birds also dispersed seeds of two exotic plants into native forest, but dispersal was localized or establishment minimal. Seed rain of bird-dispersed seeds was extensive in both forests, totaling 724 seeds of 9 native species and 2 exotics with over 85% of the seeds coming from native plants. Without suitable native dispersers, most common understory plants in Hawaiian rainforests now depend on introduced birds for dispersal, and these introduced species may actually facilitate perpetuation, and perhaps in some cases restoration, of native forests. We emphasize, however, that restoration of native forests by seed dispersal from introduced birds, as seen in this study, depends on the existence of native forests to provide a source of seeds and protection from the effects of ungulates. Our results further suggest that aggressive control of patches of non-native plants within otherwise native-dominated forests may be an important and effective conservation strategy.  相似文献   

11.
Abstract:  We assessed spatial and temporal patterns of genetic diversity to evaluate effects of river fragmentation on remnant populations of the federally endangered Rio Grande silvery minnow ( Hybognathus amarus ). Analysis of microsatellite and mitochondrial DNA detected little spatial genetic structure over the current geographic range, consistent with high gene flow despite fragmentation by dams. Maximum-likelihood analysis of temporal genetic data indicated, however, that present-day effective population size ( NeV ) of the largest extant population of this species was 78 and the ratio of effective size to adult numbers ( NeV/N ) was ∼ 0.001 during the study period (1999 to 2001). Coalescent-based analytical methods provided an estimate of historical (river fragmentation was completed in 1975) effective size ( NeI  ) that ranged between 105 and 106. We propose that disparity between contemporary and historical estimates of Ne and low contemporary Ne/N result from recent changes in demography related to river fragmentation. Rio Grande silvery minnows produce pelagic eggs and larvae subject to downstream transport through diversion dams. This life-history feature results in heavy losses of yearly reproductive effort to emigration and mortality, and extremely large variance in reproductive success among individuals and spawning localities. Interaction of pelagic early life history and river fragmentation has altered demographic and genetic dynamics of remnant populations and reduced Ne to critically low values over ecological time.  相似文献   

12.
Abstract: We reviewed the history and conservation of Sophora toromiro , a species that has been extinct in the wild since 1960 but has survived as scattered individuals in botanic gardens and private collections. The short-term conservation of S. toromiro is dependent on the management of surviving ex situ stocks. This is being achieved through international collaboration by a working group established to coordinate the conservation management of the species. Molecular evidence indicates that the species retains greater genetic variability than expected. The greatest amount of genetic variability was located in specimens outside botanic garden collections. No unmodified natural habitat survives on Rapa Nui ( Easter Island); so opportunities to establish a viable wild population are limited. Evidence from past reintroductions indicates that the best short-term opportunity for the species is through conventional horticultural management in botanic gardens and traditional farm plots on Rapa Nui. Some extinct-in-the-wild taxa (sensu World Conservation Union 1994), such as the Toromiro, retain genetic variability, and appropriate reintroduction sites exist. These taxa represent valid priorities for conservation management.  相似文献   

13.
We analyzed the amount and distribution of genetic variation in Baptisia arachnifera Duncan to develop a sampling strategy for ex situ research. Baptisia arachnifera is an endangered plant species endemic to the coastal plain of Georgia (U.S.) where all populations are within 16 km of each other. A reduction in numbers of individuals has been observed during the last 50 years. Baptisia arachnifera was polymorphic at 24% of the 37 loci examined with an average of 1.32 alleles per locus. The genetic diversity index was relatively low ( He = 0.097) as expected for endemic species. Populations were in Hardy-Weinberg equilibrium, suggesting that the species is outcrossing. Consistent with this conclusion is the observation that the majority (approximately 90%) of the genetic variation present in the species is found within individual populations. Indirect evidence of gene flow between populations was detected (   Nm = 2.35). The close proximity of the populations and the recent reduction in population sizes suggest that the populations surveyed may be fragments of a once more continuous gene pool. Based on the observed distribution of genetic diversity among populations (GST = 0.096), sampling two populations would capture 99% of the allozyme diversity surveyed. Allozyme data were used to determine which 2 of the 10 populations surveyed should be sampled to maximize the ex situ conservation of genetic diversity. Although the paper-producing companies that own most of the land where Baptisia arachnifera occurs are modifying their harvesting techniques, the species could become extinct without more effective management and preservation efforts.  相似文献   

14.
Most of the old-growth redwood ( Sequoia sempervirens ) in California has been cut; regenerating forests will probably never resemble those that were harvested, and what old growth remains on private land occurs in small, isolated remnant patches. The landscapes in which these stands occur differ so markedly from their original condition that their value as habitat to many species of wildlife, including bats, is unknown. Previous research in unfragmented redwood forests demonstrated that bats use basal hollows in old-growth redwoods as roosts. We sought to determine whether bats use similar old-growth trees as roosts when they occur in small, remnant patches of isolated old growth on commercial forest land. We compared bat occurrence in remnant and contiguous stands by collecting guano in traps suspended in hollows and by monitoring flight activity with ultrasonic bat detectors. Hollows in trees within the remnant stands had significantly more guano deposited per tree than the trees within the contiguous forest. The mean numbers of bat passes per night were statistically indistinguishable between the two treatments, although mean flight activity in the remnant stands was greater than in the contiguous forest. Bats frequently used basal hollows in small (<5 ha) stands of remnant old growth, which may be due to the closer proximity of remnants to stream courses, to their greater interface with edge where foraging success may be greater, or to the fact that the lower density of hollow-bearing trees in remnants than in contiguous forest favored greater use per tree. Significant use of small, residual old-growth redwood provides reason to maintain these remnants in managed landscapes as potentially important habitat for forest bats.  相似文献   

15.
Abstract:  Many tropical forests have been converted for agri- or silviculture or a combination of both (agroforestry). Conservation at a landscape scale requires an understanding of the distribution and abundance of native biodiversity in these converted natural ecosystems, of which the knowledge is especially poor for African agroecosystems. We compared species density and species composition of four plant groups (trees and shrubs, epiphytic vascular plants, mosses, and liverworts) among three arboreal land-use types in southwestern Ethiopia (montane rainforest fragments, shade-tree coffee home gardens, and exotic tree plantations). Species density was significantly higher in forests than in coffee home gardens for all plant groups and in exotic tree plantations for all groups except mosses. Home gardens had more vascular epiphytic species than plantations, whereas the reverse was true for mosses and liverworts. The species composition of the forest plots was sometimes more similar to home-garden plots than plantation plots and sometimes vice versa. Fifteen forest plots had, however, cumulatively more species than a random selection of 15 nonforest (coffee home garden and plantation) plots, even if the 2 plot types complemented each other in terms of habitats for forest plants. Tree plantations dominated by Eucalyptus had many small trees and shrubs in common with forests, whereas plantations with Cupressus were important substrates for forests mosses and liverworts. Our results illustrate the importance of undisturbed forests habitats for conservation of species at a landscape scale and that different human-made land-use types may complement each other in their capacity as additional habitats for forest species.  相似文献   

16.
We examined genetic diversity in 464 individuals of the monotypic lily Harperocallis flava in its two habitats (seepage bogs and a roadside right-of-way) and five populations of a co-occurring related lily, Tofieldia racemosa. The endangered H. flava, endemic to the Apalachicola lowlands of the Florida panhandle, was monomorphic for the 22 loci scored. In contrast, T. racemosa had a high proportion of polymorphic loci ( Ps = 68.2%; Pp = 47.7%) with moderate genetic diversity (   Hes = 0.134; Hep = 0.114). Estimated gene flow was moderately high ( Nm = 2.07) for T. racemosa, with most (93%) of the total genetic diversity found within populations. Despite the low level of genetic divergence, some isolation by distance was detected among T. racemosa populations. Harperocallis flava and other species without discernable genetic variation pose special problems for conservation biologists because genetic criteria are not available for the development of ex situ and in situ conservation and management strategies.  相似文献   

17.
Abstract: The endangered Hawaiian monk seal breeds at six locations in the northwestern Hawaiian Islands. To determine whether significant genetic differentiation exists among these sites, we used microsatellite loci to examine the monk seal population structure at the five largest breeding colonies. Of 27 loci isolated from other seal species, only 3 were polymorphic in an initial screening of one individual from each breeding site. Only two alleles were found at each of these 3 loci in samples of 46–108 individuals. This extremely low variation is consistent with other measures of genetic variability in this species and is probably the result of a recent severe population bottleneck, combined with a long-term history of small population sizes. Although the smallest monk seal subpopulation in this study ( Kure Atoll) showed some evidence of heterozygote deficit, possibly indicative of inbreeding, the next smallest ( Pearl and Hermes Reef) had an apparent excess of heterozygous individuals. Genetic differentiation was detected between the two subpopulations at extreme ends of the range ( Kure and French Frigate Shoals). This trend was significant only at the microsatellite locus for which we had the largest sample size ( Hg6.3: R ST = 0.206, p = 0.002; allelic goodness of fit G h = 15.412, p < 0.005). French Frigate Shoals is the source population for translocated animals that have been released primarily at Kure Atoll. Differentiation between these sites consisted of allele frequency differences (with the same allele predominant in each location at all three loci), rather than the preservation of alternative alleles. Although the translocations have had positive demographic effects, we recommend continued genetic monitoring of both the source and recipient populations because translocated individuals are now entering the breeding population.  相似文献   

18.
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.  相似文献   

19.
Abstract: To assess the genetic consequences for a Neotropical tree of the loss of its main seed disperser, we compared the genetic structure of Inga ingoides in a site where the spider monkey (Ateles paniscus) was abundant and a site where it had been eliminated by subsistence hunting. Gene flow should be reduced in the site where the spider monkey is absent, and there should be a corresponding subpopulation differentiation of seedlings within the spatial range of the movements of these primates in the absence of between-site differences in allelic frequencies. At the microhabitat (  family) scale, seedlings growing under parent plants should be genetically more related in the absence of the spider monkey than in its presence. Subpopulation differentiation was smaller where the spider monkey was present (  four loci, FST = 0.011) than where it was absent (  four loci, FST = 0.053) for the first year of study, but not for the second year (three loci, FST = 0.005 vs. 0.003). The number of alleles in common among seedlings growing under parent plants was smaller in the presence of the spider monkey than in its absence, showing family genetic structure in the first generation for both years of study ( Mann-Whitney, z = −2.17, p = 0.03 and z = −2.72, p = 0.006 for 1996 and 1997, respectively). This family genetic structure in the first generation should accelerate the development of population genetic structure. Development of genetic structure might result in demographic changes, one of which would be a fitness reduction if the species were self-incompatible, as suggested for Inga by available evidence. Large birds and mammals are the main targets of subsistence hunting in the Neotropics. Extinction of seed-dispersing frugivores may result in pronounced changes in the demographic and genetic structure of tree species in Neotropical forests.  相似文献   

20.
Genetic variation was examined in Helonias bullata , a threatened perennial plant species that occurs in isolated wetland habitats. Fifteen populations representing the species' geographic range were sampled. Genetic diversity was low for the species ( H es = 0.053) as well as within populations ( H ep = 0.029). Of the 33 allozyme loci examined, 11 (33%) were polymorphic, while on average only 12.8% (4) of the loci were polymorphic within populations. The number of alleles per polymorphic locus was 2.36 for the species and averaged 2.09 across populations. For every genetic parameter calculated, variation in H. bullata was lower than that typically found for narrowly distributed plant species. The lowest levels of genetic diversity were found in northern areas that were colonized following the last glacial epoch. The number of genotypes detected per population ranged from three to 21, with a mean of 13 for this clonally reproducing species. We found a relatively high proportion of total genetic diversity (30.6%) among populations and a significant correlation (p < 0.002) between genetic distance and geographic distance. Genetic drift phenomena appear to play a major role in the population genetics of this species. Anomalously, several populations that appeared most limited in size and vigor were genetically most variable, perhaps because they represent older, relictual populations. Life-history characteristics of H. bullata coupled with low levels of genetic diversity and the degradation and disappearance of wetlands threaten the existence of this species.  相似文献   

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