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1.
The Earth's evolutionary history is threatened by species loss in the current sixth mass extinction event in Earth's history. Such extinction events not only eliminate species but also their unique evolutionary histories. Here we review the expected loss of Earth's evolutionary history quantified by phylogenetic diversity (PD) and evolutionary distinctiveness (ED) at risk. Due to the general paucity of data, global evolutionary history losses have been predicted for only a few groups, such as mammals, birds, amphibians, plants, corals and fishes. Among these groups, there is now empirical support that extinction threats are clustered on the phylogeny; however this is not always a sufficient condition to cause higher loss of phylogenetic diversity in comparison to a scenario of random extinctions. Extinctions of the most evolutionarily distinct species and the shape of phylogenetic trees are additional factors that can elevate losses of evolutionary history. Consequently, impacts of species extinctions differ among groups and regions, and even if global losses are low within large groups, losses can be high among subgroups or within some regions. Further, we show that PD and ED are poorly protected by current conservation practices. While evolutionary history can be indirectly protected by current conservation schemes, optimizing its preservation requires integrating phylogenetic indices with those that capture rarity and extinction risk. Measures based on PD and ED could bring solutions to conservation issues, however they are still rarely used in practice, probably because the reasons to protect evolutionary history are not clear for practitioners or due to a lack of data. However, important advances have been made in the availability of phylogenetic trees and methods for their construction, as well as assessments of extinction risk. Some challenges remain, and looking forward, research should prioritize the assessment of expected PD and ED loss for more taxonomic groups and test the assumption that preserving ED and PD also protects rare species and ecosystem services. Such research will be useful to inform and guide the conservation of Earth's biodiversity and the services it provides.  相似文献   

2.
重要物种优先保护种群的确定   总被引:10,自引:0,他引:10  
由于同一物种不同种群的重要性不同、用于物种保护的资金有限以及保护与发展经济之间的矛盾,因此对于重要物种(尤其是濒危种类以及农作物和驯化动物的野生近缘种)需要确定保护什么以及保护哪儿。目前确定优先保护种群的方法主要有3类,分别为基于遗传变异、基于遗传差异性和基于遗传贡献率的方法。基于遗传变异的方法主要是根据遗传变异程度(尤其是等位基因多样性)来确定优先保护的顺序,但忽略了种群之间的遗传差异性,这容易使得存在于遗传变异程度较低的种群中的特有等位基因得不到有效保护。而基于遗传差异性的方法(如确定进化显著单元)则是从遗传分化程度的角度考虑优先性,即独特性越强的种群越具有保护价值。基于遗传贡献率的方法由于综合考虑了遗传多样性和差异性,最适合于确定哪些种群需要优先保护。我国开展此类研究十分必要。  相似文献   

3.
Tropical islands are species foundries, formed either as a by-product of volcanism, when previously submerged seabed is thrust upwards by tectonics, or when a peninsula is isolated by rising sea level. After colonisation, the geographical isolation and niche vacancies provide the competitive impetus for an evolutionary radiation of distinct species-island endemics. Yet the very attributes which promote speciation in evolutionary time also leave island endemics highly vulnerable to recent and rapid impacts by modern people. Indeed, the majority of documented human-driven extinctions have been exacted upon island endemics. The causes include over-exploitation, invasive species brought by people and destruction of island’s naturally constrained habitats. Imminent threats include inundation by rising sea levels and other adaptive pressures related to anthropogenic global warming. We review recent work which underscores the susceptibility of island endemics to the drivers of global change, and suggest a methodological framework under which, we argue, the science and mitigation of island extinctions can be most productively advanced.  相似文献   

4.
The amphibian decline and extinction crisis demands urgent action to prevent further large numbers of species extinctions. Lists of priority species for conservation, based on a combination of species' threat status and unique contribution to phylogenetic diversity, are one tool for the direction and catalyzation of conservation action. We describe the construction of a near-complete species-level phylogeny of 5713 amphibian species, which we use to create a list of evolutionarily distinct and globally endangered species (EDGE list) for the entire class Amphibia. We present sensitivity analyses to test the robustness of our priority list to uncertainty in species' phylogenetic position and threat status. We find that both sources of uncertainty have only minor impacts on our 'top 100' list of priority species, indicating the robustness of the approach. By contrast, our analyses suggest that a large number of Data Deficient species are likely to be high priorities for conservation action from the perspective of their contribution to the evolutionary history.  相似文献   

5.
Island biogeography is the study of the spatio-temporal distribution of species, communities, assemblages or ecosystems on islands and other isolated habitats. Island diversity is structured by five classes of process: dispersal, establishment, biotic interactions, extinction and evolution. Classical approaches in island biogeography focused on species richness as the deterministic outcome of these processes. This has proved fruitful, but species traits can potentially offer new biological insights into the processes by which island life assembles and why some species perform better at colonising and persisting on islands. Functional traits refer to morphological and phenological characteristics of an organism or species that can be linked to its ecological strategy and that scale up from individual plants to properties of communities and ecosystems. A baseline hypothesis is for traits and ecological strategies of island species to show similar patterns as a matched mainland environment. However, strong dispersal, environmental and biotic-interaction filters as well as stochasticity associated with insularity modify this baseline. Clades that do colonise often embark on distinct ecological and evolutionary pathways, some because of distinctive evolutionary forces on islands, and some because of the opportunities offered by freedom from competitors or herbivores or the absence of mutualists. Functional traits are expected to be shaped by these processes. Here, we review and discuss the potential for integrating functional traits into island biogeography. While we focus on plants, the general considerations and concepts may be extended to other groups of organisms. We evaluate how functional traits on islands relate to core principles of species dispersal, establishment, extinction, reproduction, biotic interactions, evolution and conservation. We formulate existing knowledge as 33 working hypotheses. Some of these are grounded on firm empirical evidence, others provide opportunities for future research. We organise our hypotheses under five overarching sections. Section A focuses on plant functional traits enabling species dispersal to islands. Section B discusses how traits help to predict species establishment, successional trajectories and natural extinctions on islands. Section C reviews how traits indicate species biotic interactions and reproduction strategies and which traits promote intra-island dispersal. Section D discusses how evolution on islands leads to predictable changes in trait values and which traits are most susceptible to change. Section E debates how functional ecology can be used to study multiple drivers of global change on islands and to formulate effective conservation measures. Islands have a justified reputation as research models. They illuminate the forces operating within mainland communities by showing what happens when those forces are released or changed. We believe that the lens of functional ecology can shed more light on these forces than research approaches that do not consider functional differences among species.  相似文献   

6.
Urbanization, one of the most important anthropogenic impacts on Earth, is rapidly expanding worldwide. This expansion of urban land‐covered areas is known to significantly reduce different components of biodiversity. However, the global evidence for this effect is mainly focused on a single diversity measure (species richness) with a few local or regional studies also supporting reductions in functional diversity. We have used birds, an important ecological group that has been used as surrogate for other animals, to investigate the hypothesis that urbanization reduces the global taxonomical and/or evolutionary diversity. We have also explored whether there is evidence supporting that urban bird communities are evolutionarily homogenized worldwide in comparison with nonurban ones by means of using evolutionary distinctiveness (how unique are the species) of bird communities. To our knowledge, this is the first attempt to quantify the effect of urbanization in more than one single diversity measure as well as the first time to look for associations between urbanization and phylogenetic diversity at a large spatial scale. Our findings show a strong and globally consistent reduction in taxonomic diversity in urban areas, which is also synchronized with the evolutionary homogenization of urban bird communities. Despite our general patterns, we found some regional differences in the intensity of the effect of cities on bird species richness or evolutionary distinctiveness, suggesting that conservation efforts should be adapted locally. Our findings might be useful for conservationists and policymakers to minimize the impact of urban development on Earth's biodiversity and help design more realistic conservation strategies.  相似文献   

7.
Establishment of conservation priorities for primates is a particular concern in the island archipelagos of Southeast Asia, where rates of habitat destruction are among the highest in the world. Conservation programs require knowledge of taxonomic diversity to ensure success. The Philippine tarsier is a flagship species that promotes environmental awareness and a thriving ecotourism economy in the Philippines. However, assessment of its conservation status has been impeded by taxonomic uncertainty, a paucity of field studies, and a lack of vouchered specimens and genetic samples available for study in biodiversity repositories. Consequently, conservation priorities are unclear. In this study we use mitochondrial and nuclear DNA to empirically infer geographic partitioning of genetic variation and to identify evolutionarily distinct lineages for conservation action. The distribution of Philippine tarsier genetic diversity is neither congruent with expectations based on biogeographical patterns documented in other Philippine vertebrates, nor does it agree with the most recent Philippine tarsier taxonomic arrangement. We identify three principal evolutionary lineages that do not correspond to the currently recognized subspecies, highlight the discovery of a novel cryptic and range-restricted subcenter of genetic variation in an unanticipated part of the archipelago, and identify additional geographically structured genetic variation that should be the focus of future studies and conservation action. Conservation of this flagship species necessitates establishment of protected areas and targeted conservation programs within the range of each genetically distinct variant of the Philippine tarsier.  相似文献   

8.
The Philippine archipelago is recognized as a biodiversity hotspot because of its high levels of endemism and numerous threatened species. Avian lineages in the Philippines feature morphologically distinct allopatric taxa, which have been variably treated either as species or subspecies depending on species concepts and recognition criteria. To understand how alternative species limits would alter diversity metrics and patterns of endemism in the Philippines, we selected 19 focal lineages of birds, each containing multiple described taxa within the Mindanao Island Group. We delimited species in an integrative, lineage-based framework using three operational criteria: species must (1) form well-supported, geographically circumscribed clades, (2) be monophyletic with significant genetic differentiation identified by a coalescent model, and (3) feature fixed differences in phenotypic characters. Our criteria identified 40 species from the original 19 focal lineages, a 50–74% increase over recent comprehensive taxonomic treatments. Genetic criteria in isolation identified an additional 10 populations that could be cryptic species in need of further study. We identified fine-scale endemism within the Mindanao Island Group, with multiple unrecognized avian endemics restricted to Samar/Leyte, Bohol Island, and the Zamboanga Peninsula. Genetic and phenotypic information support the hypothesis that polytypic bird species in the Philippines tend to be composed of evolutionarily distinct, range-restricted, allopatric replacements rather than widespread and variable “superspecies”. We conclude that lack of species recognition has resulted in underestimates of species diversity and overlooked fine-scale endemism in the Philippines. Recognizing this diversity would alter conservation priorities, shifting efforts to protect microendemics on smaller islands and finer scale endemic areas within larger islands.  相似文献   

9.

Aim

We used an eco-phylogenetic approach to investigate the diversity and assembly patterns of tropical dry forests (TDFs) in Central India. We aimed at informing conservation and restoration practices in these anthropogenically disturbed forests by identifying potential habitats of conservation significance and elements of regional biodiversity most vulnerable to human impact and climate change.

Location

Tropical dry forests of Madhya Pradesh, Central India.

Methods

We analysed the species richness, stem density, basal area and phylogenetic structure (standardized effect size of MNTD, MPD, PD and community evolutionary distinctiveness cED) of 117 tree species assemblages distributed across a ~230 to ~940 m elevational gradient. We examined how these community measures and taxonomic (Sørensen) and phylogenetic (UniFrac) beta diversity varied with elevation, precipitation, temperature and climatic stress.

Results

Species richness, phylogenetic diversity, stem density and basal area were positively correlated with elevation, with high-elevation plots exhibiting cooler temperatures, higher precipitation and lower stress. High-elevation assemblages also trended towards greater phylogenetic dispersion, which diminished at lower elevations and in drier, more stressful plots. Phylogenetic turnover was observed across the elevation gradient, and species evolutionary distinctiveness increased at lower elevations and under harsher abiotic conditions.

Main Conclusions

Harsher abiotic conditions at low elevations may act as a selective filter on plant lineages, leading to phylogenetically clustered low-diversity assemblages. These assemblages contained more evolutionarily distinct species that may contribute disproportionately to biodiversity. Conversely, milder abiotic conditions at high elevations may serve as refuges for drought-sensitive species, resulting in more diverse assemblages. Conservation practices that prioritize both high- and low-elevation habitats could promote the persistence of evolutionarily distinct species and areas of high biodiversity within the Central Indian landscape. Establishing connectivity between these habitats may provide a range of climatic conditions for species to retreat to or persist within as climates change.  相似文献   

10.
The ratio of species extinctions to introductions has been comparable for many insular assemblages, suggesting that introductions could have ‘compensated’ for extinctions. However, the capacity for introduced species to replace ecological roles and evolutionary history lost following extinction is unclear. We investigated changes in bird functional and phylogenetic diversity in the wake of extinctions and introductions across a sample of 32 islands worldwide. We found that extinct and introduced species have comparable functional and phylogenetic alpha diversity. However, this was distributed at different positions in functional space and in the phylogeny, indicating a ‘false compensation’. Introduced and extinct species did not have equivalent functional roles nor belong to similar lineages. This makes it unlikely that novel island biotas composed of introduced taxa will be able to maintain ecological roles and represent the evolutionary histories of pre‐disturbance assemblages and highlights the importance of evaluating changes in alpha and beta diversity concurrently.  相似文献   

11.
There is an increasing interest in measuring loss of phylogenetic diversity and evolutionary distinctiveness which together depict the evolutionary history of conservation interest. Those losses are assessed through the evolutionary relationships between species and species threat status or extinction probabilities. Yet, available information is not always sufficient to quantify the threat status of species that are then classified as data deficient. Data‐deficient species are a crucial issue as they cause incomplete assessments of the loss of phylogenetic diversity and evolutionary distinctiveness. We aimed to explore the potential bias caused by data‐deficient species in estimating four widely used indices: HEDGE, EDGE, PDloss, and Expected PDloss. Second, we tested four different widely applicable and multitaxa imputation methods and their potential to minimize the bias for those four indices. Two methods are based on a best‐ vs. worst‐case extinction scenarios, one is based on the frequency distribution of threat status within a taxonomic group and one is based on correlates of extinction risks. We showed that data‐deficient species led to important bias in predictions of evolutionary history loss (especially high underestimation when they were removed). This issue was particularly important when data‐deficient species tended to be clustered in the tree of life. The imputation method based on correlates of extinction risks, especially geographic range size, had the best performance and enabled us to improve risk assessments. Solving threat status of DD species can fundamentally change our understanding of loss of phylogenetic diversity. We found that this loss could be substantially higher than previously found in amphibians, squamate reptiles, and carnivores. We also identified species that are of high priority for the conservation of evolutionary distinctiveness.  相似文献   

12.
The geographical distribution of species richness and species range size of African anthropoid primates (catarrhines) is investigated and related to patterns of habitat and dietary niche breadth. Catarrhine species richness is concentrated in the equatorial regions of central and west Africa; areas that are also characterised by low average species range sizes and increased ecological specificity. Species richness declines with increasing latitude north and south of the equator, while average species range size, habitat and dietary breadth increase. Relationships between species richness, species range size and niche breadth remain once latitudinal and longitudinal effects have been removed. Among areas of lowest species richness, however, there is increased variation in terms of average species range size and niche breadth, and two trends are identified. While most such areas are occupied by a few wide-ranging generalists, others are occupied by range-restricted specialist species. That conservation efforts increasingly focus on regions of high species richness may be appropriate if these regions are also characterised by species that are more restricted in both their range size and their ecological versatility, although special consideration may be required for some areas of low species richness.  相似文献   

13.
Amidst a worldwide decline in amphibian populations, those species endemic to islands remain an important focus for conservation efforts. The Sooglossidae are a family of frog species endemic to the Seychelles islands that are believed to have evolved in isolation for approximately 75 million years. Formerly thought to inhabit just two Seychelles islands (Mahé and Silhouette), a third population was discovered on Praslin in 2009. Phylogenetic analysis based on 438 bp of mitochondrial 16S rRNA suggests that the Praslin population is most closely related to Sooglossus sechellensis from Silhouette, and identifies these as two separate clades which together sit distinct from the population on Mahé. An average of 4.06% uncorrected pairwise sequence divergence between the Praslin and Silhouette populations suggests substantial evolutionary divergence rather than recent introduction. Discriminant function analysis also revealed differences in morphology in frogs from Praslin and Mahé. DNA sequences of two Praslin specimens group more closely with the Mahé population, indicating some shared haplotypes that suggest recent secondary contact. Tests for a genetic signature of recent population expansion on either island were not significant. Our results suggest substantial evolutionary divergence between the three populations of S. sechellensis, most likely following isolation due to changes in sea level in the Indian Ocean. Whilst further genetic sampling and ecological studies are needed, our initial phylogenetic analyses suggest that the sooglossid population on Praslin should be managed as an evolutionarily significant unit to retain the uniqueness of its genetic diversity and its evolutionary trajectory within this ancient family of amphibians.  相似文献   

14.
Protected areas (PAs) are pivotal tools for biodiversity conservation on the Earth. Europe has had an extensive protection system since Natura 2000 areas were created in parallel with traditional parks and reserves. However, the extent to which this system covers not only taxonomic diversity but also other biodiversity facets, such as evolutionary history and functional diversity, has never been evaluated. Using high-resolution distribution data of all European tetrapods together with dated molecular phylogenies and detailed trait information, we first tested whether the existing European protection system effectively covers all species and in particular, those with the highest evolutionary or functional distinctiveness. We then tested the ability of PAs to protect the entire tetrapod phylogenetic and functional trees of life by mapping species'' target achievements along the internal branches of these two trees. We found that the current system is adequately representative in terms of the evolutionary history of amphibians while it fails for the rest. However, the most functionally distinct species were better represented than they would be under random conservation efforts. These results imply better protection of the tetrapod functional tree of life, which could help to ensure long-term functioning of the ecosystem, potentially at the expense of conserving evolutionary history.  相似文献   

15.
Although selectively logged tropical forests have high bird species richness, it is known that their species composition is substantially changed when compared with intact forests. Thus, we need to improve the understanding on how functional trait diversity of birds is affected in this habitat type in order to support the development of more effective conservation actions to maintain functional roles and community stability. Here, we evaluate traits responses to variations in forest vegetation integrity and how the pattern of niche occupancy is affected by this increase in species richness. We then evaluated the effects of vegetation integrity in the Atlantic rainforest on range of trait space occupied, niche packing, and trait composition in local bird communities. We also evaluate the mechanisms driving niche expansion and packing using null models. Our results show that trait composition changes in communities: (1) lower vegetation integrity increases foraging in understory and consumption of grains and ectothermic vertebrates by birds; (2) higher vegetation integrity drives higher and wider beaks and increase foraging for invertebrates in canopy. We also found that lower vegetation integrity not only is associated with the increase of species richness, but also with both expansion and packing of niche space occupied by the community. However, only niche packing had predominantly smaller values than expected by chance, indicating a strong effect of environmental filters on niche occupancy density. Although bird assemblages in more intact vegetations have lower species richness, they have greater functional distance between bird species suggesting greater stability, with a low probability of local extinctions due to a lower intensity of interspecific competition. This demonstrates that isolated assessments of species richness are potentially illusory and can lead to unsuccessful conservation measures, such as proposing selective logging in primary forests based on the supposed benefit of increased bird species richness in vegetations less intact. Furthermore, the functional composition tends to change with changes in vegetation integrity degree, thus altering the functional role provided by communities. Consequently, forests with high vegetation integrity status should be maintained, despite the lower species richness.  相似文献   

16.
Species of the Heteropogon-Themeda clade are ecologically important grasses distributed across the tropics, including widespread species, such as the pantropical Heteropogon contortus and Themeda triandra, and range-restricted species such as Heteropogon ritchiei and Themeda anathera. Here, we examine habitat preferences of the grassland/savanna and wetland species by describing bioclimatic niche characteristics, characterizing functional traits, and investigating the evolution of functional traits of 31 species in the Heteropogon-Themeda clade in relation to precipitation and temperature. The climatic limits of the clade are linked to mean annual precipitation and seasonality that also distinguish seven wetland species from 24 grassland/savanna species. Tests of niche equivalency highlighted the unique bioclimatic niche of the wetland species. However, climatic factors do not fully explain species geographic range, and other factors are likely to contribute to their distribution ranges. Trait analyses demonstrated that the wetland and grassland/savanna species were separated by culm height, leaf length, leaf area, awn length, and awn types. Phylogenetic analyses showed that the wetland species had tall stature with long and large leaves and lack of hygroscopic awns, which suggest selective pressures in the shift between savanna/grassland and wetland. The two most widespread species, H. contortus and T. triandra, have significantly different bioclimatic niches, but we also found that climatic niche alone does not explain the current geographic distributions of H. contortus and T. triandra. Our study provides a new understanding of the biogeography and evolutionary history of an ecologically important clade of C4 tropical grasses.  相似文献   

17.
Insular systems are usually characterized by have a high species diversity, endemism, and evolutionary uniqueness. Although ecological and evolutionary factors shaping insular diversity and endemism are relatively well established, there is a little understanding about climatic niche dynamics for many insular adaptive radiations. Here, we evaluate the tempo and mode of climatic niche evolution in an iconic insular radiation of lizards. By using an extensive dataset of phylogenetic and coarse-grain climatic niches, we evaluated phylogenetic niche divergence and niche conservatism across temporal and spatial scales in the Caribbean Anolis lizard radiation. We found several instances of niche shifts during the anole radiation across islands. Many of these niche shifts converged to similar climatic regimes between different islands. Furthermore, we find evidence that single-island endemic species are more limited by low suitability of climatic conditions outside its native islands than oceanic barriers due to the high climatic heterogeneity observed at least between Greater Antillean islands. These results suggest that within-lineage climatic niche conservatism has been prevalent in short time scales and likely played a role driving the exceptional insular endemism observed today.  相似文献   

18.
1. Conservation plans are required to safeguard freshwater biodiversity in the face of increasing threats. Traditionally plans have used surrogates for biodiversity that do not account for the evolutionary process, but genetic data in the form of comparative phylogeography can fulfil this role. 2. Comparative phylogeographic analyses of multiple freshwater fish and decapod crustacean species were carried out with specimens from two model systems, namely the sand dune islands of Fraser and North Stradbroke in eastern Australia. 3. Almost all of the species studied from both islands displayed an intraspecific evolutionary split between sides of the island (east/west on North Stradbroke Island, and north/south on Fraser Island), indicating that each side of each island hosts its own distinct community of populations of freshwater animals. 4. The probable process responsible for both of these divergent communities is different source populations for each side of each island. 5. This study shows that biodiversity will not always follow obvious geography and that significant diversity may exist at small scales within multiple species. These evolutionarily relevant units of biodiversity should be incorporated at the beginning of the conservation and resource management planning process.  相似文献   

19.
《新西兰生态学杂志》2011,34(1):115-136
New Zealand?s offshore and outlying islands have long been a focus of conservation biology as sites of local endemism and as last refuges for many species. During the c. 730 years since New Zealand has been settled by people, mammalian predators have invaded many islands and caused local and global extinctions. New Zealand has led international efforts in island restoration. By the late 1980s, translocations of threatened birds to predator-free islands were well under way to safeguard against extinction. Non-native herbivores and predators, such as goats and cats, had been eradicated from some islands. A significant development in island restoration in the mid-1980s was the eradication of rats from small forested islands. This eradication technology has been refined and currently at least 65 islands, including large and remote Campbell (11 216 ha) and Raoul (2938 ha) Islands, have been successfully cleared of rats. Many of New Zealand?s offshore islands, especially those without predatory mammals, are home to large numbers of breeding seabirds. Seabirds influence ecosystem processes on islands by enhancing soil fertility and through soil disturbance by burrowing. Predators, especially rats, alter ecosystem processes and cause population reductions or extinctions of native animals and plants. Islands have been promoted as touchstones of a primaeval New Zealand, but we are now increasingly aware that most islands have been substantially modified since human settlement of New Zealand. Archaeological and palaeoecological investigations, together with the acknowledgement that many islands have been important mahinga kai (sources of food) for Maori, have all led to a better understanding of how people have modified these islands. Restoration technology may have vaulted ahead of our ability to predict the ecosystem consequences of its application on islands. However, research is now being directed to help make better decisions about restoration and management of islands, decisions that take account of island history and key drivers of island ecosystem functioning.  相似文献   

20.
Comparing niche divergence among closely related taxa can yield important insights into the ecological distinctiveness of genetically similar forms, and identify the processes that are responsible for diversification in such organisms. Here, we apply newly developed techniques for analysing niche divergence to assess how ecologically distinct a group of closely related rattlesnakes (Sistrurus sp.) are and to explore the role that niche divergence may have played in their diversification. We find that all taxa even the most recently evolved subspecies (approximately 100,000 years old) are now ecologically distinct, implying a role for ecology in the diversification process. Statistical analysis based on comparisons with null models show that niche divergence between forms is more common than niche conservation. Finally, there is nonlinear relationship between phylogenetic and niche divergence in this group whereby niche divergence develops more rapidly between recently diverged subspecies than more distantly related forms. Overall, our results argue that ecology may play an important role in the diversification process in these snakes.  相似文献   

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