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1.
Functional diversity changes during tropical forest succession   总被引:1,自引:0,他引:1  
Functional diversity (FD) ‘those components of biodiversity that influence how an ecosystem operates or functions’ is a promising tool to assess the effect of biodiversity loss on ecosystem functioning. FD has received ample theoretical attention, but empirical studies are limited. We evaluate changes in species richness and FD during tropical secondary forest succession after shifting cultivation in Mexico. We also test whether species richness is a good predictor of FD. FD was calculated based on a combination of nine functional traits, and based on two individual traits important for primary production (specific leaf area) and carbon sequestration (wood density). Stand basal area was a good predictor of successional changes in diversity and FD, in contrast to fallow age. Incidence-based FD indices increased logarithmically with stand basal area, but FD weighted by species’ importance values lacked pattern with succession. Species richness and diversity are strong predictors of FD when all traits were considered; linear relationships indicate that all species are equally functionally complementary, suggesting there is little functional redundancy. In contrast, when FD was calculated for individual traits and weighted for abundances, species richness may underestimate FD.Selection of functional trait(s) critically determines FD, with large consequences for studies relating biodiversity to ecosystem functioning. Careful consideration of the traits required to capture the ecosystem process of interest is thus essential.  相似文献   

2.
Vegetation complexity is an important predictor of animal species diversity. Specifically, taller vegetation should provide more potential ecological niches and thus harbor communities with higher species richness and functional diversity (FD). Resource use behavior is an especially important functional trait because it links species to their resource base with direct relevance to niche partitioning. However, it is unclear how exactly the diversity of resource use behavior changes with vegetation complexity. To address this question, we studied avian FD in relation to vegetation complexity along a continental‐scale vegetation gradient. We quantified foraging behavior of passerine birds in terms of foraging method and substrate use at 21 sites (63 transects) spanning 3,000 km of woodlands and forests in Australia. We also quantified vegetation structure on 630 sampling points at the same sites. Additionally, we measured morphological traits for all 111 observed species in museum collections. We calculated individual‐based, abundance‐weighted FD in morphology and foraging behavior and related it to species richness and vegetation complexity (indexed by canopy height) using structural equation modeling, rarefaction analyses, and distance‐based metrics. FD of morphology and foraging methods was best predicted by species richness. However, FD of substrate use was best predicted by canopy height (ranging 10–30 m), but only when substrates were categorized with fine resolution (17 categories), not when categorized coarsely (8 categories). These results suggest that, first, FD might increase with vegetation complexity independently of species richness, but whether it does so depends on the studied functional trait. Second, patterns found might be shaped by how finely we categorize functional traits. More complex vegetation provided larger "ecological space" with more resources, allowing the coexistence of more species with disproportionately more diverse foraging substrate use. We suggest that the latter pattern was driven by nonrandom accumulation of functionally distinct species with increasing canopy height.  相似文献   

3.
Functional diversity (FD) is a key facet of biodiversity used to address central questions in ecology. Despite recent methodological advances, FD remains a complex concept and no consensus has been reached either on how to quantify it, or on how it influences ecological processes. Here we define FD as the distribution of trait values within a community. When and how to account for intraspecific trait variability (ITV) when measuring FD remains one of the main current debates. It remains however unclear to what extent accounting for population‐level ITV would modify FD quantification and associated conclusions. In this paper, we address two critical questions: (1) How sensitive are different components of FD to the inclusion of population‐level ITV? (2) Does the omission of ITV obscure the understanding of ecological patterns? Using a mixture of empirical data and simulation experiments, we conducted a sensitivity analysis of four commonly used FD indices (community weighted mean traits, functional richness, Rao's quadratic entropy, Petchey and Gaston's FD index) and their relationships with environmental gradients and species richness, by varying both the extent (plasticity or not) and the structure (contingency to environmental gradient due to local adaptation) of population‐level ITV. Our results suggest that ITV may strongly alter the quantification of FD and the detection of ecological patterns. Our analysis highlights that 1) species trait values distributions within communities are crucial to the sensitivity to ITV, 2) ITV structure plays a major role in this sensitivity and 3) different indices are not evenly sensitive to ITV, the single‐trait FD from Petchey and Gaston being the most sensitive among the four metrics tested. We conclude that the effects of intraspecific variability in trait values should be more systematically tested before drawing central conclusions on FD, and suggest the use of simulation studies for such sensitivity analyses.  相似文献   

4.
Question: Indices of functional diversity have been seen as the key for integrating information on species richness with measures that focus on those components of community composition related to ecosystem functioning. For comparing species richness among habitats on an equal‐effort basis, so‐called sample‐based rarefaction curves may be used. Given a study area that is sampled for species presence and absence in N plots, sample‐based rarefaction generates the expected number of accumulated species as the number of sampled plots increases from 1 to N. Accordingly, the question for this study is: can we construct a ‘functional rarefaction curve’ that summarizes the expected functional dissimilarity between species when n plots are drawn at random from a larger pool of N plots? Methods: In this paper, we propose a parametric measure of functional diversity that is obtained by combining sample‐based rarefaction techniques that are usually applied to species richness with Rao's quadratic diversity. For a given set of N presence/absence plots, the resulting measure summarizes the expected functional dissimilarity at an increasingly larger cumulative number of plots n (nN). Results and Conclusions: Due to its parametric nature, the proposed measure is progressively more sensitive to rare species with increasing plot number, thus rendering this measure adequate for comparing the functional diversity of species assemblages that have been sampled with variable effort.  相似文献   

5.
Different components of biodiversity may vary independently of each other along environmental gradients giving insights into the mechanisms that regulate species coexistence. In particular, the functional diversity (FD) or the presence of rare or endemic species in natural assemblages do not necessarily increase with species diversity. We studied if different components of plant species diversity (species richness, Simpson diversity, evenness) varied similarly to FD (measured as a generalization of the Simpson index) and rarity along grazing intensity and climatic gradients. Plots under different sheep grazing regimes (high and low intensity, abandonment) were surveyed in five locations along a climatic gradient in north-eastern Spain, from semi-arid lowland to moist upland locations. Variation in species diversity, functional diversity and rarity followed different patterns. Species diversity was lowest in water-stressed environments (arid locations and southern aspects) and increased with grazing more makedly in humid locations. The FD was comparable between the most species-poor and species-rich locations and decreased with grazing in the moistest location, i.e. where species diversity markedly increased. The FD did not show a strong correlation with species richness nor with the Simpson index and less specious communities could show the highest functional diversity. The rarest species in the region were more frequently found in the abandoned areas, which held the lowest species diversity. Consequently, the mechanisms that enhance the diversity of species do not necessarily support a functional differentiation among those species or the maintenance of rare species in a region. We hypothesize that the degree of dependence of functional diversity on species diversity might be mostly related to the amplitude of the species' traits pool and on how species partition the niche space available.  相似文献   

6.
Functional trait composition of plant communities has been proposed as a helpful key for understanding the mechanisms of biodiversity effects on ecosystem functioning. In this study, we applied a step‐wise modeling procedure to test the relative effects of taxonomic diversity, functional identity, and functional diversity on macrophytes community productivity along water depth gradient. We sampled 42 plots and 1513 individual plants and measured 16 functional traits and abundance of 17 macrophyte species. Results showed that there was a significant decrease in taxonomic diversity, functional identity (i.e., stem dry mass content, leaf [C] and leaf [N]), and functional diversity (i.e., floating leaf, mean Julian flowering date and rooting depth) with increasing water depth. For the multiple‐trait functional diversity (FD) indices, functional richness decreased, while functional divergence increased with water depth gradient. Macrophyte community productivity was strongly determined by functional trait composition within community, but not significantly affected by taxonomic diversity. Community‐weighted means (CWM) showed a two times higher explanatory power relative to FD indices in determining variations in community productivity. For nine of sixteen traits, CWM and FD showed significant correlations with community productivity, although the strength and direction of those relations depended on selected trait. Furthermore, functional composition in a community affected productivity through either additive or opposite effects of CWM and FD, depending on the particular traits being considered. Our results suggested both mechanisms of mass ratio and niche complementarity can operate simultaneously on variations in community productivity, and considering both CWM and FD would lead to a more profound understanding of traits–productivity relationships.  相似文献   

7.
高寒草地植物物种多样性与功能多样性的关系   总被引:5,自引:0,他引:5  
物种多样性与功能多样性的关系是生态学当前研究的热点问题之一,不同区域典型生态系统物种多样性和功能多样性的关系研究有利于生物多样性保护理论的全面发展。以青藏高原地区的主要草地生态系统—高寒草甸和高寒草原为研究对象,采用4个物种多样性指数(Patrick丰富度指数、Shannon-Weiner多样性指数、Pielou均匀度指数和Simpson优势度指数)和9个功能多样性指数(FAD功能性状距离指数、MFAD功能性状平均距离指数、基于样地的FDp和基于群落的FDc功能树状图指数、FRic功能体积指数、FEve功能均匀度指数、Rao功能离散度常二次熵指数、FDiv功能离散指数、FDis功能分散指数),分析了高寒草地植物物种多样性、功能多样性关系及其与初级生产力的关系,以期阐明3个科学问题:不同草地类型的高寒草地生态系统植物物种多样性和功能多样性有何差异?高寒草地生态系统的植物物种多样性和功能多样性有何关系?高寒草地生态系统物种多样性、功能多样性对生态系统功能的影响有何异同?研究结果表明:(1)与高寒草原相比,高寒草甸具有更高的物种多样性、功能丰富度和功能离散度;(2)高寒草甸中,Patrick丰富度与功能丰富度指数(FAD、MFAD、FDp、FDc)和功能离散度指数(FDiv)的具有较强的相关性,最优拟合方程分别为幂函数和二次多项式函数;(3)高寒草原中,Patrick丰富度与功能丰富度指数(FAD、MFAD、FDp、FDc、FRic)、Shannon指数和Simpson指数与FEve指数的相关性较强,最优拟合方程为二次多项式函数,Pielou指数与FEve指数的相关性较强,最优拟合方程为指数函数;(4)高寒草甸的初级生产力分别与物种丰富度指数Patrick、功能离散指数FDiv具有较强的相关性;而高寒草原的初级生产力与4个物种多样性指数间均具有较强的相关性,与功能离散指数FDiv具有较强的相关性,最佳拟合方程均为二次多项式函数。研究的总体结论为:物种多样性、功能多样性、二者之间的关系以及二者与生态系统服务功能(以初级生产力为例)之间的关系在高寒草甸和高寒草原群落中表现迥异,因此在研究青藏高原高寒草地的生态功能时,不能仅仅测度传统的物种多样性,还应测度与物种多样性、生态功能密切相关的功能多样性。  相似文献   

8.
Habitat discontinuity is one of the main causes of diversity reduction in lotic ecosystems. We tested the predictions of the Serial Discontinuity Concept (SDC) caused by small reservoirs on the functional diversity (FD) of Chironomidae assemblages in Neotropical Savanna streams. We obtained taxonomic information from segments upstream and downstream of small reservoirs. In addition, abiotic variables, such as stream segment width, flow velocity, dissolved oxygen, pH, conductivity, water temperature, and organic matter were measured. We analyzed the Chironomidae assemblage FD using the functional richness (FRic), functional dispersion (FDis), trait relevance, as well as the species richness metrics. We used a non-parametric paired tests to compare differences in the FD indices, species richness, and the abiotic variables between the upstream and downstream segments. The results suggest that there were reductions in the FRic, FDis, species richness, and organic matter percentage below reservoirs. Moreover, depth, width, and dissolved oxygen increased in the downstream segments. The discontinuity length presented a negative influence in the FD indexes toward downstream segments as proposed by the SDC theory. We concluded that the discontinuity length affected the dispersal ability of some Chironomidae taxa, causing richness reduction and dissimilarity in the assemblages’ functional traits toward downstream stretches.  相似文献   

9.
Three metrics of species diversity – species richness, the Shannon index and the Simpson index – are still widely used in ecology, despite decades of valid critiques leveled against them. Developing a robust diversity metric has been challenging because, unlike many variables ecologists measure, the diversity of a community often cannot be estimated in an unbiased way based on a random sample from that community. Over the past decade, ecologists have begun to incorporate two important tools for estimating diversity: coverage and Hill diversity. Coverage is a method for equalizing samples that is, on theoretical grounds, preferable to other commonly used methods such as equal-effort sampling, or rarefying datasets to equal sample size. Hill diversity comprises a spectrum of diversity metrics and is based on three key insights. First, species richness and variants of the Shannon and Simpson indices are all special cases of one general equation. Second, richness, Shannon and Simpson can be expressed on the same scale and in units of species. Third, there is no way to eliminate the effect of relative abundance from estimates of any of these diversity metrics, including species richness. Rather, a researcher must choose the relative sensitivity of the metric towards rare and common species, a concept which we describe as ‘leverage.' In this paper we explain coverage and Hill diversity, provide guidelines for how to use them together to measure species diversity, and demonstrate their use with examples from our own data. We show why researchers will obtain more robust results when they estimate the Hill diversity of equal-coverage samples, rather than using other methods such as equal-effort sampling or traditional sample rarefaction.  相似文献   

10.
Cacao agroforestry have been considered as biodiversity‐friendly farming practices by maintaining habitats for a high diversity of species in tropical landscapes. However, little information is available to evaluate whether this agrosystem can maintain functional diversity, given that agricultural changes can affect the functional components, but not the taxonomic one (e.g., species richness). Thus, considering functional traits improve the understanding of the agricultural impacts on biodiversity. Here, we measured functional diversity (functional richness‐FD, functional evenness‐FEve, and functional divergence‐Rao) and taxonomic diversity (species richness and Simpson index) to evaluate changes of bird diversity in cacao agroforestry in comparison with nearby mature forests (old‐growth forests) in the Brazilian Atlantic Forest. We used data from two landscapes with constraining areas of mature forest (49% Una and 4.8% Ilhéus) and cacao agroforestry cover (6% and 82%, respectively). To remove any bias of species richness and to evaluate assembly processes (functional overdispersion or clustering), all functional indices were adjusted using null models. Our analyses considered the entire community, as well as separately for forest specialists, habitat generalists, and birds that contribute to seed dispersal (frugivores/granivores) or invertebrate removal (insectivores). Our findings showed that small cacao agroforestry in the forested landscape sustains functional diversity (FD and FEve) as diverse as nearby forests when considering the entire community, forest specialist, and habitat generalists. However, we observed declines for frugivores/granivores and insectivores (FD and Rao). These responses of bird communities differed from those observed by taxonomic diversity, suggesting that even species‐rich communities in agroforestry may capture lower functional diversity. Furthermore, communities in both landscapes showed either functional clustering or neutral processes as the main driver of functional assembly. Functional clustering may indicate that local conditions and resources were changed or lost, while neutral assemblies may reveal high functional redundancy at the landscape scale. In Ilhéus, the neutral assembly predominance suggests an effect of functional homogenization between habitats. Thus, the conservation value of cacao agroforestry to harbor species‐rich communities and ecosystem functions relies on smallholder production with reduced farm management in a forested landscape. Finally, we emphasize that seed dispersers and insectivores should be the priority conservation targets in cacao systems.  相似文献   

11.
The effects of species loss on ecosystems depend on the community’s functional diversity (FD). However, how FD responds to environmental changes is poorly understood. This applies particularly to higher trophic levels, which regulate many ecosystem processes and are strongly affected by human-induced environmental changes. We analyzed how functional richness (FRic), evenness (FEve), and divergence (FDiv) of important generalist predators—epigeic spiders—are affected by changes in woody plant species richness, plant phylogenetic diversity, and stand age in highly diverse subtropical forests in China. FEve and FDiv of spiders increased with plant richness and stand age. FRic remained on a constant level despite decreasing spider species richness with increasing plant species richness. Plant phylogenetic diversity had no consistent effect on spider FD. The results contrast with the negative effect of diversity on spider species richness and suggest that functional redundancy among spiders decreased with increasing plant richness through non-random species loss. Moreover, increasing functional dissimilarity within spider assemblages with increasing plant richness indicates that the abundance distribution of predators in functional trait space affects ecological functions independent of predator species richness or the available trait space. While plant diversity is generally hypothesized to positively affect predators, our results only support this hypothesis for FD—and here particularly for trait distributions within the overall functional trait space—and not for patterns in species richness. Understanding the way predator assemblages affect ecosystem functions in such highly diverse, natural ecosystems thus requires explicit consideration of FD and its relationship with species richness.  相似文献   

12.
Land use intensification can greatly reduce species richness and ecosystem functioning. However, species richness determines ecosystem functioning through the diversity and values of traits of species present. Here, we analyze changes in species richness and functional diversity (FD) at varying agricultural land use intensity levels. We test hypotheses of FD responses to land use intensification in plant, bird, and mammal communities using trait data compiled for 1600+ species. To isolate changes in FD from changes in species richness we compare the FD of communities to the null expectations of FD values. In over one-quarter of the bird and mammal communities impacted by agriculture, declines in FD were steeper than predicted by species number. In plant communities, changes in FD were indistinguishable from changes in species richness. Land use intensification can reduce the functional diversity of animal communities beyond changes in species richness alone, potentially imperiling provisioning of ecosystem services.  相似文献   

13.
Brody Sandel 《Ecography》2018,41(5):837-844
Phylogenetic diversity indices are widely used to characterize the structure and diversity of ecological communities. Most indices are based on a metric that is expected to vary with species richness, so they are standardized to remove this richness‐dependence. With this standardization, values of 0 are consistent with random phylogenetic structure, while phylogenetic clustering is associated with either negative or positive values (depending on the index). One common interpretation of phylogenetic clustering is that it indicates some combination of environmental and biological filtering that restricts the species that can be present in a community. Increasingly, studies are comparing phylogenetic indices along environmental gradients to infer differences in the factors structuring communities. This comparison implicitly assumes that index values are comparable among communities with different numbers of species. Using a set of simulations, I show here that this assumption is incorrect. Holding the strength of filtering constant, communities composed of more species show larger absolute index values. This problem is most pronounced when the environmental filter favors a moderate‐sized clade strongly over others and when using the net relatedness index (NRI) to measure clustering. This bias potentially casts doubt on studies studying phylogenetic index patterns along gradients where richness also varies. Fortunately, the arising generality that more stressful environments have lower species richness and stronger clustering is opposite to this bias and therefore robust. I also show that a simple rarefaction can remove the richness‐dependence of these indices, at the expense of increased error.  相似文献   

14.
Many techniques have been developed to estimate species richness and beta diversity. Those techniques, dependent on sampling, require abundance or presence/absence data. Palaeontological data is by nature incomplete, and presence/absence data is often the only type of data that can be used to provide an estimate of ancient biodiversity. We used a simulation approach to investigate the behaviour of commonly used similarity indices, and the reliability of classifications derived from these indices, when working with incomplete data. We drew samples, of varying number and richness, from artificial species lists, which represented original life assemblages, and calculated error rates for classifications of the parent lists and samples. Using these results, we estimated the Minimum Sample Richness (MSR) needed to achieve 95% classification accuracy. Results were compared for classifications derived from several commonly used similarity indexes (Dice, Jaccard, Simpson and Raup–Crick). MSR was similar for the Dice, Jaccard and Simpson indices. MSR for the Raup–Crick index was often much lower, suggesting that it is preferable for classifying patchy data, however the performance of this index was less stable than the other three in the simulations, which required an even lower MSR. MSR can be found for all presence/absence data from the contour graphs and equations as long as the absolute species richness and the beta diversity can be estimated.  相似文献   

15.
Functional diversity (FD), species richness and community composition   总被引:15,自引:0,他引:15  
Functional diversity is an important component of biodiversity, yet in comparison to taxonomic diversity, methods of quantifying functional diversity are less well developed. Here, we propose a means for quantifying functional diversity that may be particularly useful for determining how functional diversity is related to ecosystem functioning. This measure of functional diversity “FD” is defined as the total branch length of a functional dendrogram. Various characteristics of FD make it preferable to other measures of functional diversity, such as the number of functional groups in a community. Simulating species' trait values illustrates how the relative importance of richness and composition for FD depends on the effective dimensionality of the trait space in which species separate. Fewer dimensions increase the importance of community composition and functional redundancy. More dimensions increase the importance of species richness and decreases functional redundancy. Clumping of species in trait space increases the relative importance of community composition. Five natural communities show remarkably similar relationships between FD and species richness.  相似文献   

16.
17.
Two main effects are proposed to explain biodiversity–ecosystem functioning relationships: niche complementarity and selection effects. Both can be functionally defined using the functional diversity (FD) and functional identity (FI) of the community respectively. Herein, we present results from the first tree diversity experiment that separated the effect of selection from that of complementarity by varying community composition in high‐density plots along a gradient of FD, independent of species richness and testing for the effects of FD and community weighted means of traits (a proxy for FI) on stem biomass increment (a proxy for productivity). After 4 years of growth, most mixtures did not differ in productivity from the averages of their respective monocultures, but some did overyield significantly. Those positive diversity effects resulted mostly from selection effects, primarily driven by fast‐growing deciduous species and associated traits. Net diversity effect did not increase with time over 4 years.  相似文献   

18.
Documenting and exploring the patterns of diversity of life on Earth has always been a central theme in biology. Species richness despite being the most commonly used measure of diversity in macroecological studies suffers from not considering the evolutionary and ecological differences among species. Phylogenetic diversity (PD) and functional diversity (FD) have been proposed as alternative measures to overcome this limitation. Although species richness, PD and FD are closely related, their relationships have never been investigated on a global scale. Comparing PD and FD with species richness corroborated the general assumptions of surrogacy of the different diversity measures. However, the analysis of the residual variance suggested that the mismatches between the diversity measures are influenced by environmental conditions. PD increased relative to species richness with increasing mean annual temperature, whereas FD decreased with decreasing seasonality relative to PD. We also show that the tropical areas are characterized by a FD deficit, a phenomenon, that suggests that in tropical areas more species can be packed into the ecological space. We discuss potential mechanisms that could have resulted in the gradient of spatial mismatch observed in the different biodiversity measures and draw parallels to local scale studies. We conclude that the use of multiple diversity measures on a global scale can help to elucidate the relative importance of historical and ecological processes shaping the present gradients in mammalian diversity.  相似文献   

19.
Diversity estimates play a key role in ecological assessments. Species richness and abundance are commonly used to generate complex diversity indices that are dependent on the quality of these estimates. As such, there is a long‐standing interest in the development of monitoring techniques, their ability to adequately assess species diversity, and the implications for generated indices. To determine the ability of substratum community assessment methods to capture species diversity, we evaluated four methods: photo quadrat, point intercept, random subsampling, and full quadrat assessments. Species density, abundance, richness, Shannon diversity, and Simpson diversity were then calculated for each method. We then conducted a method validation at a subset of locations to serve as an indication for how well each method captured the totality of the diversity present. Density, richness, Shannon diversity, and Simpson diversity estimates varied between methods, despite assessments occurring at the same locations, with photo quadrats detecting the lowest estimates and full quadrat assessments the highest. Abundance estimates were consistent among methods. Sample‐based rarefaction and extrapolation curves indicated that differences between Hill numbers (richness, Shannon diversity, and Simpson diversity) were significant in the majority of cases, and coverage‐based rarefaction and extrapolation curves confirmed that these dissimilarities were due to differences between the methods, not the sample completeness. Method validation highlighted the inability of the tested methods to capture the totality of the diversity present, while further supporting the notion of extrapolating abundances. Our results highlight the need for consistency across research methods, the advantages of utilizing multiple diversity indices, and potential concerns and considerations when comparing data from multiple sources.  相似文献   

20.
Aim In times of biodiversity crisis, it is extremely important to understand diversity gradients. In particular, the study of the diversity of ecological functions is a key issue for the management of ecosystem integrity. Here we identify areas of low functional diversity of the native fish fauna in European drainage basins and we determine the relative importance of three underlying mechanisms: environmental filtering, geographic isolation and climatic history. Location The European continent. Methods Based on 14 morphological traits that are closely related to fish function (habitat and dietary niches), three independent functional diversity indices [functional richness (FR), functional evenness (FE), functional divergence (FD)] were calculated for 128 European drainage basins with a total of 230 fish species. The indices were standardized for species richness using null models. The patterns of the standardized indices are described and three potentially underlying mechanisms are tested using variance partitioning and multi‐linear regression models. Results FR and FD were highest in eastern European drainage basins and in Great Britain and lowest in the Mediterranean. FE patterns were less pronounced. All observed patterns were mainly governed by geographic isolation and present environmental conditions. Within the environmental conditions, average annual temperature and precipitation were good predictors for functional diversity. The role of habitat diversity and size was negligible. Main conclusions Geographic isolation coupled with harsh environmental conditions such as extreme temperatures and low precipitation, as in Mediterranean regions, can lead to low FR and FD. This can be explained by extinction that could not be compensated by re‐colonization and high speciation. Due to their high functional redundancy, communities in these areas might better withstand further species extinctions on a small scale. Over the short term, however, their often extremely low FR suggests a less flexible functioning that can hinder their ability to withstand today's rapid environmental and anthropogenic threats.  相似文献   

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