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1.
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   

2.
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   

3.
Reviews that summarize the genetic diversity of plant species in relation to their life history and ecological traits show that forest trees have more genetic diversity at population and species levels than annuals or herbaceous perennials. In addition, among-population genetic differentiation is significantly lower in trees than in most herbaceous perennials and annuals. Possible reasons for these differences between trees and herbaceous perennials and annuals have not been discussed critically. Several traits, such as high rates of outcrossing, long-distance pollen and seed dispersal, large effective population sizes (Ne), arborescent stature, low population density, longevity, overlapping generations, and occurrence in late successional communities, may make trees less sensitive to genetic bottlenecks and more resistant to habitat fragmentation or climate change. We recommend that guidelines for genetic conservation strategies be designed differently for tree species versus other types of plant species. Because most tree species fit an LH scenario (low [L] genetic differentiation and high [H] genetic diversity), tree seeds could be sourced from a few populations distributed across the species’ range. For the in situ conservation of trees, translocation is a viable option to increase Ne. In contrast, rare herbaceous understory species are frequently HL (high differentiation and low diversity) species. Under the HL scenario, seeds should be taken from many populations with high genetic diversity. In situ conservation efforts for herbaceous plants should focus on protecting habitats because the typically small populations of these species are vulnerable to the loss of genetic diversity. The robust allozyme genetic diversity databases could be used to develop conservation strategies for species lacking genetic information. As a case study of reforestation with several tree species in denuded areas on the Korean Peninsula, we recommend the selection of local genotypes as suitable sources to prevent adverse effects and to insure the successful restoration in the long term.  相似文献   

4.
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   

5.
Abstract: Zieria prostrata (Rutaceae) is known from only four headlands within a 3-km stretch of coastline in New South Wales, Australia. The species was presumed to have occurred at a headland 24 km south of its present range. We used random amplified polymorphic DNA analysis to assess patterns of genetic variation within and among the extant populations. The analysis also included an individual reputedly rescued from the now extinct population. A high level of population divergence was revealed by principal coordinate analysis and an analysis of molecular variance (AMOVA; 37% among populations). Our genetic findings provide implications for the conservation management of the species. First, the loss of any one population would lead to a severe loss of genetic variation. Second, an adequate ex situ collection must sample the full range of genetic diversity from all populations. Third, the consequences of mixing populations may be an important conservation consideration if further translocations proceed. Fourth, the individual apparently sampled prior to its population extinction is genetically similar to individuals from one of the extant sites. This degree of similarity was unexpected and, after further investigation, led to the conclusion that prior existence of the species at the site is doubtful. Subsequently, a planned reintroduction program was abandoned. So far, of these four management implications, only the last has had a direct management outcome. Those implications that failed to lead to practical management outcomes did so because the same management recommendations could be obtained without genetic research. Clearly, the challenge for more effective conservation is to identify those cases in which genetic studies are likely to produce practical outcomes for conservation managers. This may be best achieved by assessing the outcomes of genetic studies already conducted.  相似文献   

6.
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.  相似文献   

7.
Translocations and the Preservation of Allelic Diversity   总被引:3,自引:0,他引:3  
Translocation is a tool commonly used for the conservation of threatened and endangered fish species. Despite extensive use, the biological implications of translocation remain poorly understood. Of particular interest is the effect of translocation on genetic variability. Maintenance of genetic variability in these "refuge" populations is assumed to be important for both short- and long-term success. We examined allozyme variability at 16 loci for western mosquitofish ( Gambusia affinis ) populations with known histories of introduction. Refuge populations had significantly lower levels of heterozygosity. Refuge populations also had considerably lower levels of allelic diversity than parental populations. All losses were of relatively rare alleles (frequency less than 0.1 in parental population). These losses were probably due to an undocumented bottleneck early in the introduction history. These results were surprising because the initial transplant involved 900 fish and because mosquitofish have numerous reproductive traits that should minimize the effects of bottlenecks on genetic diversity. A literature review revealed that genetic variability is often reduced in refuge populations and that such reductions typically involve the loss of alleles. We suggest that translocated populations be examined periodically for losses of genetic variability.  相似文献   

8.
Isozyme, randomly amplified polymorphic DNA, and ribosomal DNA analyses were used to evaluate the genetic distinctness and diversity of the endangered island endemic Malacothamnus fasciculatus var. nesioticus (Santa Cruz Island bush mallow). Analysis of amplified DNA profiles and electrophoretic data indicates that var. nesioticus is genetically distinct from other varieties within the species. No isozyme or amplified DNA variation was detected among individuals of nesioticus populations, but restriction site analysis of ribosomal DNA revealed the presence of three closely related genotypes within one of the nesioticus populations. Apparent clonal structuring was found in both nesioticus populations and in one mainland population, var. nuttallli . Results from these analyses provide the background necessary for the design of conservation efforts with recovery goals that insure the long-term survival of Malacothamnus fasciculatus var. nesioticus .  相似文献   

9.
Abstract: Conservation of rare plant species often involves small, local populations dispersed within apparently suitable habitat. We used enzyme electrophoresis to study genotypic diversity in 32 populations of Calystegia collina , a self-incompatible clonal plant endemic to serpentine substrate in northern California's Coast Range. Genotypic diversity within a population was a good predictor of seed set by randomly marked flowers. Ramets from the most abundant genotypes in a population were most likely to produce flowers, but flowers from abundant genotypes were less likely to produce seed capsules than were flowers from rarer genotypes. These results are consistent with previous findings that reproductive success in C. collina is limited by the availability of compatible pollen. On small serpentine outcrops supporting only one or two populations, C. collina did not show reduced genotypic diversity or heterozygosity compared with populations on large outcrops supporting many populations. Instead, genotypic diversity and outcrop size had strong but independent influences on reproductive success. Although large serpentine outcrops contain more populations and provide better conditions for flower and seed production, significant diversity of unique genotypes clearly exists in isolated serpentine outcrops. Our findings suggest that plant conservation strategies must take into account the natural distribution of populations. The effects of habitat fragmentation on C. collina and other plant species that occur naturally in small, discrete patches may be unlike those that have been documented in more recently fragmented species.  相似文献   

10.
Maintenance of biodiversity through seed banks and botanical gardens, where the wealth of species’ genetic variation may be preserved ex situ, is a major goal of conservation. However, challenges can persist in optimizing ex situ collections if trade-offs exist among cost, effort, and conserving species evolutionary potential, particularly when genetic data are not available. We evaluated the genetic consequences of population preservation informed by geographic (isolation by distance [IBD]) and environmental (isolation by environment [IBE]) distance for ex situ collections for which population provenance is available. We used 19 genetic and genomic data sets from 15 plant species to assess the proportion of population genetic differentiation explained by geographic and environmental factors and to simulate ex situ collections prioritizing source populations based on pairwise geographic distance, environmental distance, or both. Specifically, we tested the impact prioritizing sampling based on these distances may have on the capture of neutral, functional, or putatively adaptive genetic diversity and differentiation. Individually, IBD and IBE explained limited population genetic differences across all 3 genetic marker classes (IBD, 10–16%; IBE, 1–5.5%). Together, they explained a substantial proportion of population genetic differences for functional (45%) and adaptive (71%) variation. Simulated ex situ collections revealed that inclusion of IBD, IBE, or both increased allelic diversity and genetic differentiation captured among populations, particularly for loci that may be important for adaptation. Thus, prioritizing population collections based on environmental and geographic distance data can optimize genetic variation captured ex situ. For the vast majority of plant species for which there is no genetic information, these data are invaluable to conservation because they can guide preservation of genetic variation needed to maintain evolutionary potential within collections.  相似文献   

11.
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   

12.
Pathogen-driven declines in animal populations are increasingly regarded as a major conservation issue. The Tasmanian devil (Sarcophilus harrisii) is threatened with extinction by devil facial tumor disease, a unique transmissible cancer. The disease is transmitted through direct transfer of tumor cells, which is possible because the genetic diversity of Tasmanian devils is low, particularly in the major histocompatibility complex genes of the immune system. The far northwest of Tasmania now holds the last remaining disease-free wild devil populations. The recent discovery of unique major histocompatibility complex genotypes in the northwestern region of Tasmania has raised the possibility that some animals may be resilient to the disease. We examined the differences in the epidemiology and population effects of devil facial tumor disease at 3 well-studied affected sites in eastern Tasmania and 1 in western Tasmania (West Pencil Pine). In contrast to the 3 eastern sites, there has been no rapid increase in disease prevalence or evidence of population decline at West Pencil Pine. Moreover, this is the only onsite at which the population age structure has remained unaltered 4 years after the first detection of disease. The most plausible explanations for the substantial differences in population effects and epidemiology of the disease between eastern and western sites are geographic differences in genotypes or phenotypes of devils and functional differences between tumor strains in the 2 regions. We suggest that conservation efforts focus on identifying whether either or both these explanations are correct and then, if resistance alleles exist, to attempt to spread the resistant alleles into affected populations. Such assisted selection has rarely been attempted for the management of wildlife diseases, but it may be widely applicable.  相似文献   

13.
Abstract: Swayne's hartebeest ( Alcelaphus buselaphus swaynei ) is an endangered antelope that survives in four or five relict populations in Ethiopia. We examined the two main populations (Senkele and Nechisar) for mitochondrial (D-loop) and nuclear (microsatellite) variability in order to measure levels of genetic variation within the subspecies and degree of differentiation between populations. For comparison, we examined samples from a large population of red hartebeest ( Alcelaphus buselaphus caama ). Both swaynei and caama exhibited high levels of variation. There was significant differentiation between the populations of swaynei at Senkele and Nechisar, and gene diversity in Nechisar, the smaller of the two populations, was significantly lower than that in Senkele. Many mitochondrial haplotypes and microsatellite alleles present at high frequencies among the Senkele individuals were missing in Nechisar, suggesting that the translocation of animals from Senkele undertaken in 1974 did not contribute notably to the gene pool in Nechisar. Subsamples taken from Senkele in 1988 and 1995 showed a significant change in allele frequencies, a change that probably can be attributed to a massive population decline during this period. We recommend that both populations be protected in situ to maintain as much as possible of the diversity that exists within the taxon and that a breeding program be established. In spite of the earlier unsuccessful attempt, we argue that translocation of animals for enhancement of population size as well as genetic variation in Nechisar should be considered.  相似文献   

14.
Ellers J  Rog S  Braam C  Berg MP 《Ecology》2011,92(8):1605-1615
Increases in biodiversity can result from an increase in species richness, as well as from a higher genetic diversity within species. Intraspecific genetic diversity, measured as the number of genotypes, can enhance plant primary productivity and have cascading effects at higher trophic levels, such as an increase in herbivore and predator richness. The positive effects of genotypic mixtures are not only determined by additive effects, but also by interactions among genotypes, such as facilitation or inhibition. However, so far there has been no effort to predict the extent of such effects. In this study, we address the question of whether the magnitude of the effect of genotype number on population performance can be explained by the extent of dissimilarity in key traits among genotypes in a mixture. We examine the relative contribution of genotype number and phenotypic dissimilarity among genotypes to population performance of the soil arthropod, Orchesella cincta. Nearly homogeneous genotypes were created from inbred isofemale lines. Phenotypic dissimilarity among genotypes was assessed in terms of three life-history traits that are associated with population growth rate, i.e., egg size, egg development time, and juvenile growth rate. A microcosm experiment with genotype mixtures consisting of one, two, four, and eight genotypes, showed that genotypic richness strongly increased population size and biomass production and was associated with greater net diversity effects. Most importantly, there was a positive log-linear relationship between phenotypic dissimilarity in a mixture and the net diversity effects for juvenile population size and total biomass. In other words, the degree of phenotypic dissimilarity among genotypes determined the magnitude of the genotypic richness effect, although this relationship leveled off at higher values of phenotypic dissimilarity. Although the exact mechanisms responsible for these effects are currently unknown, similar advantages of trait dissimilarity have been found among species. Hence, to better understand population performance, genotype number and phenotypic dissimilarity should be considered collectively.  相似文献   

15.
The Butte County meadowfoam ( Limnanthes floccosa subsp, californica ), an endangered annual endemic to vernal pools in Butte County, California, is of agronomic interest as a sperm whale oil substitute. Because it is threatened by rapid development of the community of Chico, a field and genetic survey was required to guide a conservation program. Eight new populations were discovered, bringing the total known to eleven. Population sizes varied from 220 to 45,689 plants; mean seed set per flower, a bioassay of site quality, ranged from 0.82 to 2.56 among populations. The array of associated taxa, an indicator of appropriate habitat was fairly uniform across populations. Based on allozyme and morphometric data no introgression between L. floccosa subsp, californica and L. alba subsp, alba at sympatric sites was found. Populations were remarkably monomorphic, probably due to past population bottlenecks in conjunction with high selfing rates. Electrophoretic analysis of 28 isozyme loci revealed that 96% of total genetic diversity was distributed among populations. The migration rate, Nm , calculated from the mean frequency of private alleles, ( P (1)), and from the proportion of genetic variation distributed between populations, Gst , estimated exchange between local populations of one diploid individual every 50 to 100 generations, respectively. Mean genetic identity among populations was 0.91 ± 0.068. Grouping of populations on the basis of genetic distance identified two distinct populations and three clusters of populations that deserve high priority for preservation because they are likely to have high frequencies of locally adapted alleles. The low migration rates and substantial differentiation among populations suggest a conservation plan that emphasizes preservation of as many populations as possible, at possible expense of numbers of individuals.  相似文献   

16.
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   

17.
Abstract:  A joint demographic and population genetics stage-based model for a subdivided population was applied to Gentiana pneumonanthe , an early successional perennial herb, at a regional (metapopulation) scale. We used numerical simulations to determine the optimal frequency of habitat disturbance (sod cutting) and the intensity of gene flow among populations of G. pneumonanthe to manage both population viability and genetic diversity in this species. The simulations showed that even small populations that initially had near-equal allele frequencies could, if managed properly through sod cutting every 6 to 7 years, sustain their high genetic variation over the long run without gene flow. The more the allele frequencies in the small populations are skewed, however, the higher the probability that in the absence of gene flow, some alleles will be lost and within-population genetic variation will decrease even under proper management. This implies that although local population dynamics should be the major target for management, regional dynamics become important when habitat fragmentation and decreased population size lead to the loss of local genetic diversity. The recommended strategy to improve genetic composition of small populations is the introduction of seeds or seedlings of nonlocal origin.  相似文献   

18.
温度是导致水稻(Oryza sativa L.)遗传分化的一个重要环境因素,而海拔是导致环境温度差异的一个重要的地理因素。水稻的雄配子即花粉粒的活性极易受环境温度的影响。研究用来自云南2 670 m海拔的粳稻地方品种小麻谷与具籼稻细胞质背景的籼粳交后代品系南34正反交产生的F1,然后将其分布种植在海拔高低相差达1 800 m的4个海拔点并与育性稳定的1个滇Ⅰ型细胞质不育系杂交产生了8个测交群体,用SSR标记检测发现海拔因素和细胞质背景都对测交群体的遗传分化产生了影响。结果表明:同一海拔产生的正反交群体(合系42A//小麻谷/南34)间的遗传多样性具有比较明显的差异。正交组合F1(小麻谷×南34)在1 860 m的海拔产生的群体的遗传多样性最高,在1 600 m的海拔产生的群体的遗传多样性最低。在反交组合F1产生的群体中,群体的遗传多样性随产生群体海拔(400 m海拔除外)的升高而降低。利用Nei′s遗传距离评价各群体间的遗传分化,正反交组合F1产生测交群体间的遗传分化均高于同一组合在不同海拔下产生的测交群体之间的遗传分化,同一组合在4个海拔产生的测交群体间的遗传分化总体趋势是随海拔差异的增加而增大。研究证实了环境温度差异及细胞质背景导致F1产生的雄配子基因型发生选择,导致雄配子基因型的遗传比例偏离孟德尔规律,影响其后代群体的遗传多样性和遗传分化。  相似文献   

19.
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.  相似文献   

20.
Abstract:  The Mesa Central of Mexico is of special conservation interest due to its high richness of freshwater fish species, of which the goodeines are one of the most representative groups. Through an integrated approach, we determined conservation priorities for goodeine populations. We based our recommendations on the genetic diversity (variation in five microsatellite DNA loci) in 10 populations of Zoogoneticus quitzeoensis and on an analysis of ecological (e.g., presence of exotic species), social (e.g., political situation), and environmental (e.g., pollution) information for 52 historical occurrence points for species in the genus Zoogoneticus . Patterns of genetic erosion and genetic diversity indices were closely associated with human impact. Recent bottleneck events were most evident in the populations from remnants of the lakes drained at the beginning of the twentieth century. We identified seven operational conservation units (OCUs), all of which should be conserved because they contain unique portions of the total variation of the species. Special attention needs to be given to increase genetic variability, recover population sizes, and reestablish contact among populations within OCUs. It is imperative to create an integrative and effective approach for the recovery and conservation of the freshwater fish diversity of Central Mexico that is based on social and natural sciences.  相似文献   

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