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1.
Individual dispersal,landscape connectivity and ecological networks   总被引:1,自引:0,他引:1  
Connectivity is classically considered an emergent property of landscapes encapsulating individuals' flows across space. However, its operational use requires a precise understanding of why and how organisms disperse. Such movements, and hence landscape connectivity, will obviously vary according to both organism properties and landscape features. We review whether landscape connectivity estimates could gain in both precision and generality by incorporating three fundamental outcomes of dispersal theory. Firstly, dispersal is a multi‐causal process; its restriction to an ‘escape reaction’ to environmental unsuitability is an oversimplification, as dispersing individuals can leave excellent quality habitat patches or stay in poor‐quality habitats according to the relative costs and benefits of dispersal and philopatry. Secondly, species, populations and individuals do not always react similarly to those cues that trigger dispersal, which sometimes results in contrasting dispersal strategies. Finally, dispersal is a major component of fitness and is thus under strong selective pressures, which could generate rapid adaptations of dispersal strategies. Such evolutionary responses will entail spatiotemporal variation in landscape connectivity. We thus strongly recommend the use of genetic tools to: (i) assess gene flow intensity and direction among populations in a given landscape; and (ii) accurately estimate landscape features impacting gene flow, and hence landscape connectivity. Such approaches will provide the basic data for planning corridors or stepping stones aiming at (re)connecting local populations of a given species in a given landscape. This strategy is clearly species‐ and landscape‐specific. But we suggest that the ecological network in a given landscape could be designed by stacking up such linkages designed for several species living in different ecosystems. This procedure relies on the use of umbrella species that are representative of other species living in the same ecosystem.  相似文献   

2.
Carnivores are often particularly sensitive to landscape fragmentation. Ecological corridors may help to connect local populations, ensuring gene flow and retaining viable meta-populations. We aimed to establish habitat suitability models for two large carnivores in Poland, the grey wolf Canis lupus Linnaeus, 1758 and the Eurasian lynx Lynx lynx Linnaeus, 1758, based on ecological niche factor analysis (ENFA). Secondly, we calculated least cost paths (LCPs) based on cost values obtained from ENFA. Thirdly, we determined structures that might act as barriers, thus diminishing the value of the corridor unless appropriate conservation measures are taken. We compared some of the results with actual dispersal data of four lynx in eastern Poland. Results indicate that both species are highly marginalised. Less habitat that is currently available in Poland is suitable for lynx than for wolves. We determined a total of 76 LCPs. Comparison of these theoretical corridors with actual dispersal routes suggests that the traits of calculated LCPs are mostly within the range of those of real routes. We highlight a variety of features that might act as barriers, such as major roads (including planned highways), urbanized areas, and large un-forested areas. We give suggestions where concerted conservation efforts (eg wildlife passages) might be particularly well-directed.  相似文献   

3.
Habitat destruction and fragmentation are known to strongly affect dispersal by altering the quality of the environment between populations. As a consequence, lower landscape connectivity is expected to enhance extinction risks through a decrease in gene flow and the resulting negative effects of genetic drift, accumulation of deleterious mutations and inbreeding depression. Such phenomena are particularly harmful for amphibian species, characterized by disjunct breeding habitats. The dispersal behaviour of amphibians being poorly understood, it is crucial to develop new tools, allowing us to determine the influence of landscape connectivity on the persistence of populations. In this study, we developed a new landscape genetics approach that aims at identifying land-uses affecting genetic differentiation, without a priori assumptions about associated ecological costs. We surveyed genetic variation at seven microsatellite loci for 19 Alpine newt (Mesotriton alpestris) populations in western Switzerland. Using strips of varying widths that define a dispersal corridor between pairs of populations, we were able to identify land-uses that act as dispersal barriers (i.e. urban areas) and corridors (i.e. forests). Our results suggest that habitat destruction and landscape fragmentation might in the near future affect common species such as M. alpestris. In addition, by identifying relevant landscape variables influencing population structure without unrealistic assumptions about dispersal, our method offers a simple and flexible tool of investigation as an alternative to least-cost models and other approaches.  相似文献   

4.
Permeability of the landscape matrix between amphibian breeding sites   总被引:1,自引:0,他引:1  
For organisms that reproduce in discrete habitat patches, land cover between patches (known as the matrix) is important for dispersal among breeding sites. Models of patchy populations often incorporate information on the permeability of the matrix to dispersal, sometimes based on expert opinion. I estimated the relative resistance to gene flow of land cover types and barriers using FST calculated from microsatellite markers in two amphibians, within an 800‐km2 area in northern Switzerland. The species included a frog (Rana temporaria: 996 individuals, 48 populations, seven markers) and a newt (Triturus alpestris: 816 individuals, 41 populations, seven markers). Open fields and urban areas were more resistant to gene flow than forested land; roads and highways also reduced permeability. Results were similar for the two species. However, differences in resistance among matrix elements were relatively low: gene flow through urban areas was reduced by only 24–42% relative to forest; a divided highway reduced gene flow by 11–40% and was 7–8 times more resistant than a secondary road. These data offer an empirically based alternative to expert opinion for setting relative resistance values in landscape models.  相似文献   

5.
Abstract Many natural populations in areas of continuous habitat exhibit some form of local genetic structure. Anthropogenic habitat fragmentation can also strongly influence the dynamics of gene flow between populations. We used eight microsatellite markers to investigate the population genetic structure of an abundant forest species, the Australian bush rat (Rattus fuscipes), in the subtropical forests of south‐east Queensland. Five sites were sampled, allowing pairwise comparisons within continuous habitat and across clearings. Weak, but significant population differentiation and a significant pattern of isolation by distance was detected over the small scale (<10 km) of this study. Fine‐scale analysis at a single site (<1 km) showed a significant correlation between individual female genetic distance and geographical distance, but no similar pattern among male individuals. There was no evidence of increased population differentiation across clearings relative to comparisons within continuous forest. This was attributed to dispersal within corridors of remnant and revegetated habitat between the forested areas. We concluded that an inherently restricted dispersal ability, female philopatry and natural habitat heterogeneity play an important part in the development of genetic structure among populations of R. fuscipes. It is important to understand the relationship between landscape features and the pattern of gene flow among continuous populations, as this allows us to predict the impact of fragmentation on natural populations.  相似文献   

6.
Background: There is limited understanding about bird dispersal behaviour and seedling distribution of endangered tree species in patchy environments, although these processes are important for plant species persistence.

Aims: We tested how patch features affected bird behaviour and seed dispersal, and thus seedling distribution of the endangered Chinese yew tree (Taxus chinensis).

Methods: In the present study, we combined field data of bird dispersal behaviour and GIS-based information to elucidate the influence of spatial features of habitat patches on bird dispersal behaviour, and the resulting effects on the seedling distribution of the endangered Chinese yew in two patchy habitats.

Results: Our results showed that the only seed source patch could attract eight bird species for dispersal at the two sites. Post-foraging movements of bird dispersers was strongly related to both topography and the relative locations of habitat patches. Yew seedlings aggregated only at the seed source and bamboo recruitment patches, which was affect by both the spatial distribution of recruitment patches and patch use by dispersers.

Conclusions: Our results emphasise that bamboo patches in both patchy environments provide the necessary conditions for germination of yew seeds, and the post-foraging behaviour of dispersers determines seed deposited in these patches. Our study highlights the importance of the dispersal behaviour of frugivorous birds in the successful regeneration and colonisation of yew populations in patchy habitats.  相似文献   

7.
1. A diverse array of patterns has been reported regarding the spatial extent of population genetic structure and effective dispersal in freshwater macroinvertebrates. In river systems, the movements of many taxa can be restricted to varying degrees by the natural stream channel hierarchy. 2. In this study, we sampled populations of the non‐biting freshwater midge Echinocladius martini in the Paluma bioregion of tropical northeast Queensland to investigate fine scale patterns of within‐ and among‐stream dispersal and gene flow within a purported historical refuge. We amplified a 639‐bp fragment of mitochondrial cytochrome c oxidase subunit I and analysed genetic structure using pairwise ΦST, hierarchical amova , Mantel tests and a parsimony network. Genetic variation was partitioned among stream sections, using Streamtree , to investigate the effect of potential instream dispersal barriers. 3. The data revealed strong natal site fidelity and significant differentiation among neighbouring, geographically proximate streams. We found evidence for only episodic adult flight among sites on separate stream reaches. Overall, however, our data suggested that both larval and adult dispersal was largely limited to within a stream channel. 4. This may arise from a combination of the high density of riparian vegetation physically restricting dispersal and from the joint effects of habitat stability and large population sizes. Together these latter may make it more likely that upstream populations will persist, even in the absence of regular compensatory upstream flight, and will thus reduce the adaptive value of dispersal among streams. Taken together, these data suggest that dispersal of E. martini is highly restricted, to the scale of only a few kilometres, and hence occurs predominantly within the natal stream.  相似文献   

8.
1. Using a portion of the mitochondrial cytochrome oxidase I gene, we evaluated the population genetic structure of a geographically rare black fly (Metacnephia coloradensis) that is a habitat specialist in outlet streams of large, productive, alpine lakes in Colorado, U.S.A. Given its rarity and life history traits that restrict dispersal, we hypothesised that genetic structure would show a signature of allopatric fragmentation associated with climatic warming since Pleistocene glaciations. 2. We tested for genetic isolation by distance (IBD) and applied nested clade analysis (NCA) to ask whether current genetic structure is primarily a consequence of historic fragmentation or if there is evidence of ongoing gene flow. 3. Only four populations were located despite a thorough search of potential sites, and they demonstrated a significant degree of genetic structure (FST = 0.17). However, there was some evidence of IBD in a plot of genetic versus geographic distance, and NCA further supported IBD and restricted ongoing gene flow in clades at all nested levels. Compared with a more widespread alpine black fly (Prosimulium neomacropyga) in the same region, M. coloradensis demonstrated significantly less population genetic structure. 4. Although these results counterintuitively implicate limited ongoing gene flow driving current population structure, significant IBD may be a signature of historic gene flow, especially if migration–drift equilibrium has not yet been reached since a late‐Pleistocene fragmentation event. Extraordinarily dense local populations probably allowed M. coloradensis to maintain large effective population sizes and minimise genetic drift rates. 5. Despite large local populations, M. coloradensis is vulnerable to continued rapid environmental change because of its limited geographic distribution and high habitat specificity.  相似文献   

9.
Habitat destruction threatens biodiversity by reducing the amount of available resources and connectivity among geographic areas. For organisms living in fragmented habitats, population persistence may depend on dispersal, which maintains gene flow among fragments and can prevent inbreeding within them. It is centrally important to understand patterns of dispersal for bees living in fragmented areas given the importance of pollination systems and recently documented declines in bee populations. We used population and landscape genetic techniques to characterize patterns of dispersal over a large fragmented area in southern Costa Rica for the orchid bee species Euglossa championi. First, we estimated levels of genetic differentiation among forest fragments as ?PT, an analog to the traditional summary statistic FST, as well as two statistics that may more adequately represent levels of differentiation, GST and Dest. Second, we used a Bayesian approach to determine the number and composition of genetic groups in our sample. Third we investigated how genetic differentiation changes with distance. Fourth, we determined the extent to which deforested areas restrict dispersal. Finally, we estimated the extent to which there were temporal differences in allele frequencies within the same forest fragments. Within years we found low levels of differentiation even over 80 km, and no effect of land use type on level of genetic differentiation. However, we found significant genetic differentiation between years. Taken together our results suggest that there are high levels of gene flow over this geographic area, and that individuals show low site fidelity over time.  相似文献   

10.
Landscape features of anthropogenic or natural origin can influence organisms' dispersal patterns and the connectivity of populations. Understanding these relationships is of broad interest in ecology and evolutionary biology and provides key insights for habitat conservation planning at the landscape scale. This knowledge is germane to restoration efforts for the New England cottontail (Sylvilagus transitionalis), an early successional habitat specialist of conservation concern. We evaluated local population structure and measures of genetic diversity of a geographically isolated population of cottontails in the northeastern United States. We also conducted a multiscale landscape genetic analysis, in which we assessed genetic discontinuities relative to the landscape and developed several resistance models to test hypotheses about landscape features that promote or inhibit cottontail dispersal within and across the local populations. Bayesian clustering identified four genetically distinct populations, with very little migration among them, and additional substructure within one of those populations. These populations had private alleles, low genetic diversity, critically low effective population sizes (3.2–36.7), and evidence of recent genetic bottlenecks. Major highways and a river were found to limit cottontail dispersal and to separate populations. The habitat along roadsides, railroad beds, and utility corridors, on the other hand, was found to facilitate cottontail movement among patches. The relative importance of dispersal barriers and facilitators on gene flow varied among populations in relation to landscape composition, demonstrating the complexity and context dependency of factors influencing gene flow and highlighting the importance of replication and scale in landscape genetic studies. Our findings provide information for the design of restoration landscapes for the New England cottontail and also highlight the dual influence of roads, as both barriers and facilitators of dispersal for an early successional habitat specialist in a fragmented landscape.  相似文献   

11.
Dispersal is crucial for gene flow and often determines the long‐term stability of meta‐populations, particularly in rare species with specialized life cycles. Such species are often foci of conservation efforts because they suffer disproportionally from degradation and fragmentation of their habitat. However, detailed knowledge of effective gene flow through dispersal is often missing, so that conservation strategies have to be based on mark–recapture observations that are suspected to be poor predictors of long‐distance dispersal. These constraints have been especially severe in the study of butterfly populations, where microsatellite markers have been difficult to develop. We used eight microsatellite markers to analyse genetic population structure of the Large Blue butterfly Maculinea arion in Sweden. During recent decades, this species has become an icon of insect conservation after massive decline throughout Europe and extinction in Britain followed by reintroduction of a seed population from the Swedish island of Öland. We find that populations are highly structured genetically, but that gene flow occurs over distances 15 times longer than the maximum distance recorded from mark–recapture studies, which can only be explained by maximum dispersal distances at least twice as large as previously accepted. However, we also find evidence that gaps between sites with suitable habitat exceeding ~20 km induce genetic erosion that can be detected from bottleneck analyses. Although further work is needed, our results suggest that M. arion can maintain fully functional metapopulations when they consist of optimal habitat patches that are no further apart than ~10 km.  相似文献   

12.
Theory predicts that dispersal throughout metapopulations has a variety of consequences for the abundance and distribution of species. Immigration is predicted to increase abundance and habitat patch occupancy, but gene flow can have both positive and negative demographic consequences. Here, we address the eco‐evolutionary effects of dispersal in a wild metapopulation of the stick insect Timema cristinae, which exhibits variable degrees of local adaptation throughout a heterogeneous habitat patch network of two host‐plant species. To disentangle the ecological and evolutionary contributions of dispersal to habitat patch occupancy and abundance, we contrasted the effects of connectivity to populations inhabiting conspecific host plants and those inhabiting the alternate host plant. Both types of connectivity should increase patch occupancy and abundance through increased immigration and sharing of beneficial alleles through gene flow. However, connectivity to populations inhabiting the alternate host‐plant species may uniquely cause maladaptive gene flow that counters the positive demographic effects of immigration. Supporting these predictions, we find the relationship between patch occupancy and alternate‐host connectivity to be significantly smaller in slope than the relationship between patch occupancy and conspecific‐host connectivity. Our findings illustrate the ecological and evolutionary roles of dispersal in driving the distribution and abundance of species.  相似文献   

13.
Man-made habitat fragmentation is a major concern in river ecology and is expected to have particularly detrimental effects on aquatic species with limited dispersal abilities, like the bullhead (Cottus gobio). We used ten microsatellite markers to investigate small-scale patterns of gene flow, current dispersal and neutral genetic diversity in a morphologically diverse river where fragmented and unfragmented sections could be compared. We found high genetic differentiation between sampling sites with a maximum F ST of 0.32 between sites separated by only 35 km. A significant increase of genetic differentiation with geographical distance was observed in the continuous river section as well as in the full dataset which included headwater populations isolated by anthropogenic barriers. Several lines of evidence are consistent with the hypothesis that such barriers completely block upstream movement while downstream dispersal may be little affected. In the unfragmented habitat, dispersal rates were also higher in the direction of water flow than against it. The resulting asymmetry in gene flow likely contributes to the decrease of genetic variation observed from the lower reaches towards the headwaters, which is particularly pronounced in physically isolated populations. Our findings suggest that headwater populations, due to their isolation and low genetic variation, may be particularly vulnerable to extinction.  相似文献   

14.
The increasing fragmentation of natural habitats may strongly affect patterns of dispersal and gene flow among populations, and thus alter evolutionary dynamics. We examined genetic variation at twelve microsatellite loci in the Agile frog (Rana dalmatina) from 22 breeding ponds in the Iberian Peninsula, the southwest limit of its range, where populations of this species are severely fragmented and are of conservation concern. We investigated genetic diversity, structure and gene flow within and among populations. Diversity as observed heterozygosities ranged from 0.257 to 0.586. The mean number of alleles was 3.6. Just one population showed a significant F IS value. Four populations show evidence of recent bottlenecks. Strong pattern of structure was observed due to isolation by distance and to landscape structure. The average degree of genetic differentiation among populations was F ST = 0.185. Three operational conservation units with metapopulation structure were identified. Additionally, there are some other isolated populations. The results reinforce the view that amphibian populations are highly structured even in small geographic areas. The knowledge of genetic structure pattern and gene flow is fundamental information for developing programmes for the preservation of R. dalmatina at the limits of its geographic distribution.  相似文献   

15.
Abstract: Natural and anthropogenic landscape features, such as rivers, mountain ranges, and roads can alter animal dispersal paths and movement patterns. Consequently landscape, through its effects on dispersal, may influence many ecological processes, including disease transmission, invasion dynamics, and gene flow. To investigate influences of landscape features on dispersal patterns of a large mammal, we captured and radiomarked 363 juvenile male white-tailed deer (Odocoileus virginianus), including 212 confirmed dispersers, in 2 topographically dissimilar study areas in Pennsylvania, USA. Dispersal azimuths were uniformly distributed in the western study area (WSA), where there was irregular, hilly topography. Mean dispersal azimuths paralleled ridge direction in the eastern study area, where long parallel ridges were aligned northeast-southwest. Major roads in both areas and a large river in the WSA were semipermeable barriers to dispersal of juvenile males; dispersal paths were less likely to intersect these linear features. Dispersal movements were direct and brief, typically lasting <12 hours. For all dispersers, we found no evidence for preference or avoidance of establishing adult, postdispersal ranges in proximity to roads; however, deer that encountered roads near the terminus of their dispersal path were more likely to stop on the near side. Further, for deer that established postdispersal home ranges near major roads, these features influenced range placement such that locations were typically clustered on one side of the road. The influence of roads, rivers, and mountains on dispersal paths and postdispersal locations of white-tailed deer suggest that landscape-specific features should be considered in conservation and management of this and possibly other species of large mammals.  相似文献   

16.
Measures of gene flow in the Columbian ground squirrel   总被引:2,自引:0,他引:2  
F. Stephen Dobson 《Oecologia》1994,100(1-2):190-195
From analyses of published data and a review of the literature, I studied indirect and direct measures of gene flow among populations of Columbian ground squirrels, Spermophilus columbianus. New analyses were used to examine an allozyme data set (seven polymorphic loci) that had been collected by Zammuto and Millar (1985a) from six populations of ground squirrels that were spread over 183 km. G-tests indicated significant variation in allele frequencies among populations, but F-statistics revealed relatively little population differentiation (average F ST=0.026). F ST values were used to estimate rates of gene flow indirectly and indicated fairly high rates of gene flow (average N e m=13.5). Recorded dispersal distances of individual ground squirrels were fairly short (most<4 km, maximum recorded distance was 8.5 km), and the minimum distance between populations used to create the allozyme data set was about 25 km. Thus, direct dispersal among the populations in the allozyme data set was highly unlikely. Small genetically effective populations may have experienced high rates of migration over short distances (about 43% of adults in local populations were immigrants), however, resulting in homogeneous allele frequencies over the geographic range. This explanation provides an alternative to invoking gene flow in the recent past to explain discrepancies between dispersal distances in the field and homogenization of allele frequencies over large ranges, Mammalian species that have virtually complete dispersal of subadult males from the natal area might be expected to exhibit relatively high rates of gene flow, regardless of actual dispersal distances. Genetically effective populations may be much smaller than more extensive ecological populations and experience higher rates of gene flow.  相似文献   

17.
  1. Habitat fragmentation is one of the main threats to biodiversity. Barriers to dispersal caused by anthropogenic habitat alteration may affect phylogeographic patterns in freshwater mussels. Knowledge of the phylogenetic and phylogeographic patterns of unionoids is vital to inform protection of their biodiversity.
  2. Here, we assessed influences of dams and their environmental effects on the genetic diversity and population connectivity of a broadly distributed freshwater mussel, Nodularia douglasiae, in Poyang Lake Basin.
  3. The results showed high genetic diversity in areas without dams and low genetic diversity in areas with dams. High genetic differentiation and low gene flow were found among the 11 populations. Genetic variation was significantly correlated with dissolved oxygen levels.
  4. The observation of low genetic diversity in populations separated by dams indicated that those populations were subjected to genetic erosion and demographic decline because they are disconnected from other populations with higher diversity. High genetic differentiation and low gene flow among the 11 populations could be correlated with anthropogenic habitat alteration.
  5. These results indicated that anthropogenic habitat alterations have led to the decline in freshwater mussel diversity. Therefore, we recommend maintaining favourable habitat conditions and connectivity of rivers or lakes, and strengthening study of life histories with host-test experiments to identify potential host fish species to strengthen the knowledge base underpinning freshwater mussel conservation.
  相似文献   

18.
Zooplanktonic taxa have a greater number of distinct populations and species than might be predicted based on their large population sizes and open‐ocean habitat, which lacks obvious physical barriers to dispersal and gene flow. To gain insight into the evolutionary mechanisms driving genetic diversification in zooplankton, we developed eight microsatellite markers to examine the population structure of an abundant, globally distributed mesopelagic copepod, Haloptilus longicornis, at 18 sample sites across the Atlantic and Pacific Oceans (= 761). When comparing our microsatellite results with those of a prior study that used a mtDNA marker (mtCOII,= 1059, 43 sample sites), we unexpectedly found evidence for the presence of a cryptic species pair. These species were globally distributed and apparently sympatric, and were separated by relatively weak genetic divergence (reciprocally monophyletic mtCOII lineages 1.6% divergent; microsatellite FST ranging from 0.28 to 0.88 across loci, P < 0.00001). Using both mtDNA and microsatellite data for the most common of the two species (= 669 for microsatellites, = 572 for mtDNA), we also found evidence for allopatric barriers to gene flow within species, with distinct populations separated by continental landmasses and equatorial waters in both the Atlantic and Pacific Ocean basins. Our study shows that oceanic barriers to gene flow can act as a mechanism promoting allopatric diversification in holoplanktonic taxa, despite the high potential dispersal abilities and pelagic habitat for these species.  相似文献   

19.
Assessing patterns of genetic structure and diversity of threatened species has become an essential tool for determining conservation status and designing management strategies. We examine the genetic structure of the Sierra Madre sparrow (Xenospiza baileyi), a species restricted to fragmented patches of subalpine bunchgrass in three small isolated areas of northwestern and central Mexico. Coding and non-coding regions of mtDNA (1,878 bp) from individuals of the only three known populations revealed the existence of a single major lineage, with closely related haplotypes being shared between populations across the range. The sharing of haplotypes between the distant northwest and central populations (~800 km) suggests a recent fragmentation of a formerly contiguous population. Despite a lack of large-scale phylogeographic structure, haplotype frequencies at local scales revealed significant genetic differentiation and high F ST values between all three remaining populations, even between localities separated by less than 12 km. These results suggest restricted gene flow and limited dispersal, likely due to the species’ inability to cross areas of unsuitable habitat. On the basis of genetic interchange and ecological equivalence criteria, we recommend that the species be managed as a single unit, permitting the strengthening of the small population in the northwest with individuals from central Mexico, and/or the translocation of individuals to new areas of suitable habitat.  相似文献   

20.
Climate change, genotypic diversity and gene flow in reef-building corals   总被引:10,自引:0,他引:10  
In the ocean, large‐scale dispersal and replenishment by larvae is a key process underlying biological changes associated with global warming. On tropical reefs, coral bleaching, degradation of habitat and declining adult stocks are also likely to change contemporary patterns of dispersal and gene flow and may lead to range contractions or expansions. On the Great Barrier Reef, where adjacent reefs form a highly interconnected system, we use allozyme surveys of c. 3000 coral colonies to show that populations are genetically diverse, and rates of gene flow for a suite of five species range from modest to high among reefs up to 1200 km apart. In contrast, 700 km further south on Lord Howe Island, genetic diversity is markedly lower and populations are genetically isolated. The virtual absence of long‐distance dispersal of corals to geographically isolated, oceanic reefs renders them extremely vulnerable to global warming, even where local threats are minimal.  相似文献   

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