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1.
Currently, the habitat of many species is fragmented, resulting in small local populations with individuals occasionally dispersing between the remaining habitat patches. In a solitary bee metapopulation, extinction probability was related to both local bee population sizes and pollen resources measured as host plant population size. Patch size, on the other hand, had no additional predictive power. The turnover rate of local bee populations in 63 habitat patches over 4 years was high, with 72 extinction events and 31 colonization events, but the pollen plant population was stable with no extinctions or colonizations. Both pollen resources and bee populations had strong and independent effects on extinction probability, but connectivity was not of importance. Colonizations occurred more frequently within larger host plant populations. For metapopulation survival of the bee, large pollen plant populations are essential, independent of current bee population size.  相似文献   

2.
Because spatial connectivity is critical to dispersal success and persistence of species in highly fragmented landscapes, the way that we envision and measure connectivity is consequential for biodiversity conservation. Connectivity metrics used for predictive modeling of spatial turnover and patch occupancy for metapopulations, such as with Incidence Function Models (IFM), incorporate distances to and sizes of possible source populations. Here, our focus is on whether habitat quality of source patches also is considered in these connectivity metrics. We propose that effective areas (weighted by habitat quality) of source patches should be better surrogates for population size and dispersal potential compared to unadjusted patch areas. Our review of a representative sample of the literature revealed that only 12.5% of studies incorporated habitat quality of source patches into IFM-type connectivity metrics. Quality of source patches generally was not taken into account in studies even if habitat quality of focal patches was included in analyses. We provide an empirical example for a metapopulation of a rare wetland species, the round-tailed muskrat (Neofiber alleni), demonstrating that a connectivity metric based on effective areas of source patches better predicts patch colonization and occupancy than a metric that used simple patch areas. The ongoing integration of landscape ecology and metapopulation dynamics could be hastened by incorporating habitat quality of source patches into spatial connectivity metrics applied to species conservation in fragmented landscapes.  相似文献   

3.
4.
This study aimed at comparing six patch connectivity measures by fitting them to field data. We used occupancy data for eight beetle and two pseudoscorpion species from 281 hollow oaks in southeast Sweden. Species occupancy was modelled in relation to tree characteristics and one measure of patch connectivity at a time. For each connectivity measure we searched for the spatial scale that generated the best fit to field data. Connectivity measures that only include occupied patches provided better model fits than those that include all patches. When occupancy data are absent for surrounding habitat patches, information that reflects occurrence probabilities can be included in the connectivity measure. However, in this study incorporation of such information resulted in only a slight improvement of model fit. A frequently used connectivity measure based on the negative exponential function was relatively poor in explaining species’ occurrence; for eight species out of nine a buffer measure was better. A better fit was obtained when the negative exponential function was modified to take into account that habitat patches may “compete” for the immigrants. The spatial scale with the best fit tended to be larger when we used connectivity measures in which dispersal sources are identified with lower precision. Thus, the outcomes from different multiple‐scale studies are not directly comparable if the density of dispersal sources is not measured in the same way. Overall we conclude that buffer measures are useful, as they give good predictions and are easy to understand and use. If a biologically more realistic measure is needed, one that up‐weights the closest patches should be used. Finally, the possibility that habitat patches may compete with each other for immigrants should be considered when selecting a connectivity measure.  相似文献   

5.
We report cases of long‐distance dispersal in Odonata, some of which were directly observed by identifying single individuals of riverine species in unsuitable habitat, mostly desert, far distant from reproduction habitats. The shortest possible linear distances of the observation points to reproduction habitats were measured. Furthermore, established populations of riverine species were recorded in artificial lakes in central and southern Namibia far distant from the next regular reproduction sites. Our records demonstrate that single individuals of riverine species were probably covering distances of several hundred kilometres over arid landscape without any intervening possible reproduction habitat. Although it is likely that only small numbers of individuals of the river populations may disperse long distances, relatively recent colonizations of artificial habitats suggest that a few, or even single, dispersing individuals may lead to large‐scale‐range expansions.  相似文献   

6.
Ecosystems are linked by the movement of organisms across habitat boundaries and the arrangement of habitat patches can affect species abundance and composition. In tropical seascapes many coral reef fishes settle in adjacent habitats and undergo ontogenetic habitat shifts to coral reefs as they grow. Few studies have attempted to measure at what distances from nursery habitats these fish migrations (connectivity) cease to exist and how the abundance, biomass and proportion of nursery species change on coral reefs along distance gradients away from nursery areas. The present study examines seascape spatial arrangement, including distances between habitats, and its consequences on connectivity within a tropical seascape in Mozambique using a seascape ecology approach. Fish and habitat surveys were undertaken in 2016/2017 and a thematic habitat map was created in ArcGIS, where cover and distances between habitat patches were calculated. Distance to mangroves and seagrasses were significant predictors for abundance and biomass of most nursery species. The proportions of nursery species were highest in the south of the archipelago, where mangroves were present and decreased with distance to nurseries (mangroves and seagrasses). Some nursery species were absent on reef sites farthest from nursery habitats, at 80 km from mangroves and at 12 km from seagrass habitats. The proportion of nursery/non-nursery snapper and parrotfish species, as well as abundance and biomass of seagrass nursery species abruptly declined at 8 km from seagrass habitats, indicating a threshold distance at which migrations may cease. Additionally, reefs isolated by large stretches of sand and deep water had very low abundances of several nursery species despite being within moderate distances from nursery habitats. This highlights the importance of considering the matrix (sand and deep water) as barriers for fish migration.  相似文献   

7.
A major conclusion of studying metapopulation biology is that species conservation should favor regional rather than local population persistence. Regional persistence is tightly linked to size, spatial configuration and quality of habitat patches. Hence it is important for the management of endangered species that priority patches can be identified. We developed a predictive model of patch occupancy by capercaillie, a threatened grouse species, based on a single snapshot of data. We used logistic regression to predict patch occupancy as a function of patch size, isolation, connectivity, relative altitude, and biogeographical area. The probability of a patch being occupied increased with patch size and increasing altitude, and decreased with increasing distance to the next occupied patch. Patch size was the most important predictor although occupied patches varied considerably in size. Our model only uses data on the number, size and spatial configuration of habitat patches. It is a useful tool to designate priority areas for conservation, i.e. large core patches with high resilience in habitat quality, smaller island‐patches that still have high probability of being inhabited or becoming recolonised, and patches functioning as “stepping stones”. If capercaillie is to be preserved, habitat suitability needs to be maintained in a functional network of patches that account for size and inter‐patch distance thresholds as found in this study. We suggest that similar area‐isolation relationships are valid for almost any region within the distribution range of capercaillie. The thresholds for occupancy are however likely to depend on characteristics of the respective landscape. The outcome of our study emphasises the need for future investigations that explore the relationship between patch occupancy, matrix quality and its resistance to dispersing individuals.  相似文献   

8.
The Pampa grassland of Argentina is one of the most highly threatened biomes in the world. A high proportion of the original grassland cover has been transformed into land for agriculture or degraded. In the southern part of the region, fragmented semi‐natural grasslands over exposed rock still persist and connectivity between them is assumed to be crucial for maintaining viable populations. We quantified overall connectivity of grassland patches in a sector of the Southern Pampa region, and investigated the degree to which landscape connectivity explains entomophilous plant species assemblages in a subset of patches. We characterized each of the 301 patches in the landscape by their degree of intra‐patch and inter‐patch connectivity based on graph theory, and considering threshold dispersal distances from 100 to 1000 m. We surveyed entomophilous plant species in 39 grassland patches and classified the species in three categories (annual herbs, perennial herbs and shrubs) considering their different growth form and longevity. The influence of connectivity variables on entomophilous plant species assemblages variation was explored using Canonical Correspondence Analysis. Although grassland patches were poorly connected at all threshold distances, some of them were found to be critical for global connectivity. Connectivity significantly explained total, annual‐biennial and shrub assemblages for all threshold dispersal distances (6–13% of total variation). Variation in annual species assemblages was associated with intra‐patch and inter‐patch connectivity at short distance (100 m), while variation in shrub species assemblages was explained by intra‐patch and inter‐patch connectivity for distances between 100 m and 1000 m. This study evidenced the low connectivity of the study system, allowed the identification of critical areas for conservation, and provided valuable information to develop management strategies in increasingly human‐dominated landscapes.  相似文献   

9.
Anthropogenic changes in land use and the extirpation of apex predators have facilitated explosive growth of mesopredator populations. Consequently, many species have been subjected to extensive control throughout portions of their range due to their integral role as generalist predators and reservoirs of zoonotic disease. Yet, few studies have monitored the effects of landscape composition or configuration on the demographic or behavioral response of mesopredators to population manipulation. During 2007 we removed 382 raccoons (Procyon lotor) from 30 forest patches throughout a fragmented agricultural ecosystem to test hypotheses regarding the effects of habitat isolation on population recovery and role of range expansion and dispersal in patch colonization of mesopredators in heterogeneous landscapes. Patches were allowed to recolonize naturally and demographic restructuring of patches was monitored from 2008–2010 using mark-recapture. An additional 25 control patches were monitored as a baseline measure of demography. After 3 years only 40% of experimental patches had returned to pre-removal densities. This stagnant recovery was driven by low colonization rates of females, resulting in little to no within-patch recruitment. Colonizing raccoons were predominantly young males, suggesting that dispersal, rather than range expansion, was the primary mechanism driving population recovery. Contrary to our prediction, neither landscape connectivity nor measured local habitat attributes influenced colonization rates, likely due to the high dispersal capability of raccoons and limited role of range expansion in patch colonization. Although culling is commonly used to control local populations of many mesopredators, we demonstrate that such practices create severe disruptions in population demography that may be counterproductive to disease management in fragmented landscapes due to an influx of dispersing males into depopulated areas. However, given the slow repopulation rates observed in our study, localized depopulation may be effective at reducing negative ecological impacts of mesopredators in fragmented landscapes at limited spatial and temporal scales.  相似文献   

10.
Ongoing habitat loss and fragmentation is considered a threat to biodiversity as it can create small, isolated populations that are at increased risk of extinction. Tree‐dependent species are predicted to be highly sensitive to forest and woodland loss and fragmentation, but few studies have tested the influence of different types of landscape matrix on gene flow and population structure of arboreal species. Here, we examine the effects of landscape matrix on population structure of the sugar glider (Petaurus breviceps) in a fragmented landscape in southeastern South Australia. We collected 250 individuals across 12 native Eucalyptus forest remnants surrounded by cleared agricultural land or exotic Pinus radiata plantations and a large continuous eucalypt forest. Fifteen microsatellite loci were genotyped and analyzed to infer levels of population differentiation and dispersal. Genetic differentiation among most forest patches was evident. We found evidence for female philopatry and restricted dispersal distances for females relative to males, suggesting there is male‐biased dispersal. Among the environmental variables, spatial variables including geographic location, minimum distance to neighboring patch, and degree of isolation were the most important in explaining genetic variation. The permeability of a cleared agricultural matrix to dispersing gliders was significantly higher than that of a pine matrix, with the gliders dispersing shorter distances across the latter. Our results added to previous findings for other species of restricted dispersal and connectivity due to habitat fragmentation in the same region, providing valuable information for the development of strategies to improve the connectivity of populations in the future.  相似文献   

11.
Lack of landscape connectivity and habitat loss is major threats to biodiversity and ecosystem integrity in nature reserves aimed at conservation. In this study, we used structural pattern and functional connectivity metrics to analyze the spatial patterns and landscape connectivity of habitat patches for the Shangyong sub-reserve of the Xishuangbanna Nature Reserve from 1970, 1990, and 2000. On the basis of vegetation and land cover data, we applied the equivalent connected area ECA(PC) indicator to analyze the changes in forest connectivity. Four distance thresholds (2, 4, 8, 12 km) were considered to compare the patch importance of connectivity by dECA values. The results showed the declining trends of landscape connectivity measured by ECA(PC) index from 1970 to 2000. The importance of connectivity in each forest patch varied with the increment of dispersal distances at the patch level, and some important habitat patches, which exhibit a potential to enhance landscape connectivity, should be given more attention. The least-cost pathways based on network structure were displayed under four dispersal distances in three periods. The results showed that the number of paths among the fragments of forest patches exhibited radical increases for larger dispersal distances. Further correlation analyses of AWF, ECA (IIC), and ECA (PC) showed the weakest and least-frequent correlations with the structural pattern indices, while H presented more significant correlations with the PD fragmentation metric. Furthermore, Kendall's rank correlations between the forest patch area and functional connectivity indicators showed that dECA (PC) and dAWF indicators should provided the area-based prioritization of habitat patches. Moreover, the low-rank correlations showed that dF and dLCP can be considered as effective and appropriate indicators for the evaluation of habitat features and network patterns.  相似文献   

12.
Connectivity of habitat patches is crucial for wildlife dispersal and survival, and identifying patches with high importance for maintaining connectivity can aid effective wildlife management. Knowledge of the habitat distribution of the Tibetan antelope in the Altun Mountain National Nature Reserve, which is essential for connectivity analysis, remains limited. We analyzed potential habitat distribution and priority patches using GIS-based habitat suitability modeling with three weighting factors and evaluated the connectivity of habitat patches under four dispersal distance scenarios. Patches with high habitat suitability covered 25.39 % of the total area, and these patches were selected for connectivity analysis as resource patches. Connectivity analysis indicated that, although the overall probability of connectivity (PC) showed an upward trend with increasing dispersal distance, the importance of each patch varied considerably under different dispersal distance scenarios. Transfer analysis of patch numbers between different importance levels revealed that the number of patches becoming less important was higher than the number of patches becoming more important when dispersal distance increased. In addition, nine patches covering 38.49 % of the suitable habitat area were identified as priority patches, in particular the patches near the Kardun and the Karchuka inspection stations. We also found that the habitat distribution of the Tibetan antelope obtained from the suitability model matched the population distribution determined by a field survey. Correlation analysis between patch area and the percentage of PC index value loss (dPC) revealed that the larger patches in this region were more likely to be important for maintaining connectivity.  相似文献   

13.
The South African grassland biome is one of the most threatened biomes in South Africa. Approximately 45% of the grassland biome area is transformed, degraded or severely invaded by alien plants and the remaining natural areas are highly fragmented. In this fragmented landscape, the connectivity between habitat patches is very important to maintain viable populations. In this study we aimed to quantify connectivity of the grassland biome in Mpumalanga using graph theory in order to identify conservation priorities and to direct conservation efforts. Graph theory‐based connectivity indices have the ability to combine spatially explicit habitat data with species specific dispersal data and can quantify structural and functional connectivity over large landscapes. We used these indices to quantify the overall connectivity of the study area, to determine the influence of abandoned croplands on overall connectivity, and to identify the habitat patches and vegetation types most in need of maintaining overall connectivity. Natural areas were identified using 2008 land cover data for Mpumalanga. Connectivity within the grassland biome of Mpumalanga was analysed for grassland species with dispersal distances ranging from 50 to 1000 m. The grassland habitat patches were mostly well connected, with 99.6% of the total habitat area connected in a single component at a threshold distance of 1000 m. The inclusion of abandoned croplands resulted in a 33% increase in connectivity at a threshold distance of 500 m. The habitat patches most important for maintaining overall connectivity were the large patches of continuous habitat in the upper and lower centres of the study area and the most important vegetation types were the Wakkerstroom Montane Grassland and the Eastern Temperate Freshwater Wetlands. These results can be used to inform management decisions and reserve design to improve and maintain connectivity in this biome.  相似文献   

14.
Habitat loss can alter animal movements and disrupt animal seed dispersal mutualisms; however, its effects on spatial patterns of seed dispersal are not well understood. To explore the effects of habitat loss on seed dispersal distances and seed dispersion (aggregation), we created a spatially explicit, individual‐based model of an animal dispersing seeds (SEADS—Spatially Explicit Animal Dispersal of Seeds) in a theoretical landscape of 0%–90% habitat loss based on three animal traits: movement distance, gut retention time, and time between movements. Our model design had three objectives: to determine the effects of (1) animal traits and (2) habitat loss on seed dispersal distances and dispersion and (3) determine how animal traits could mitigate the negative effects of habitat loss on these variables. SEADS results revealed a complex interaction involving all animal traits and habitat loss on dispersal distances and dispersion, driven by a novel underlying mechanism of fragment entrapment. Unexpectedly, intermediate habitat loss could increase dispersal distances and dispersion relative to low and high habitat loss for some combinations of animal traits. At intermediate habitat loss, movement between patches was common, and increased dispersal distances and dispersion compared to continuous habitats because animals did not stop in spaces between fragments. However, movement between patches was reduced at higher habitat loss as animals became trapped in fragments, often near the parent plant, and dispersed seeds in aggregated patterns. As movement distance increased, low time between movements and high gut retention time combinations permitted more movement to adjacent patches than other combinations of animal traits. Because habitat loss affects movement in a nonlinear fashion under some conditions, future empirical tests would benefit from comparisons across landscapes with more than two levels of fragmentation.  相似文献   

15.
1. Three independent methods were used to investigate population structure in the butterfly Plebejus argus . First, migration and dispersal ability were measured by mark–release–recapture in seven adjacent habitat patches, and by release of butterflies in unoccupied habitat. Secondly, colonization of newly created habitat was observed over 7 years. Finally, genetic differentiation of local populations within a metapopulation was investigated. Sampled local populations included parts of the mark–release–recapture study area.
2. Plebejus argus is relatively sedentary: the maximum movement detected was 395 m, and only 2% of individuals moved further than 100 m between recaptures on different days. None the less, adjacent local populations in the mark–release–recapture study area were linked by occasional migration, with ≈ 1.4% of individuals moving between patches separated by 13–200 m.
3. Despite low mobility, observed colonizations occurred rapidly over distances of 1 km. Because P. argus occurs at high population densities, 1.4% migration can generate enough migrants to colonize newly suitable habitat quickly at this spatial scale.
4. Mark–release–recapture data were used to predict that there would be limited genetic differentiation through drift between local populations at this spatial scale. The prediction was supported by allele frequency data for the same local populations.
5. Genetic differentiation often indicates higher levels of migration than are revealed by the movements of marked individuals. This study shows that when experimental releases and extensive marking are undertaken in areas that are large relative to most movements, indirect measures of gene flow and direct measures of dispersal can concur.
6. Evidence from the three different approaches was complementary, indicating that P. argus occurs as metapopulations within the study area.  相似文献   

16.
In landscape ecology, substantial theoretical progress has been made in understanding how critical threshold levels of habitat loss may result in sudden changes in landscape connectivity to animal movement. Empirical evidence for such thresholds in real systems, however, remains scarce. Streambed landscapes provide a strong testing ground for studying critical thresholds because organisms are faced with substantial environmental heterogeneity while attempting to overcome the physical force of water. In this study, I report on the results from a series of experiments investigating the influence of habitat abundance and current velocity on the movement dynamics of two stream herbivores (caddisfly larva Agapetus boulderensis and snail Physa sp.) that differ substantially in how they perceive landscape structure. Specifically, I ask whether critical thresholds to herbivore movement exist in streambed landscapes. By exploiting the pattern recognition capabilities of artificial neural networks, I found that the rate, sinuosity and directionality of movement by Agapetus and Physa varied nonlinearly according to the abundance of habitat patches, current velocity and habitat-current interaction. Both the study organisms exhibited threshold responses to habitat abundance, yet the location and slope of these thresholds differed between species and with respect to different current velocities. These results suggest that a critical threshold in functional connectivity (i.e. the connection of habitat patches by dispersal) is not an inherent property of the landscape, but in fact emerges from the interplay of species' interactions with landscape structure. Moreover, current velocity interacted with habitat abundance to elicit strong upstream-oriented movement for both the species. This suggests that dispersing individuals may be polarized in the upstream direction and therefore functional connectivity is not equal in all directions. Such results highlight the need for future research addressing the sources of variability of critical threshold effects in ecological phenomena.  相似文献   

17.
道路对林地景观连接度的影响——以巩义市为例   总被引:1,自引:0,他引:1  
梁国付  许立民  丁圣彦 《生态学报》2014,34(16):4775-4784
基于景观连接度原理,借用景观连接度指数,在地理信息系统支持下,探讨了巩义市山区林地景观在不同距离阈值下连接度的变化,定量分析了道路对林地景观连接度的影响。结果显示,随着景观距离阈值的增大,无论是否有道路,林地景观整体可能连通性指数值都表现为逐渐增大;对林地景观连接度起"非常高"和"高"作用的林地斑块数量都比较少,但占林地总面积比例较大,面积大的林地斑块在提高景观连接度中起的作用较大;道路的分割使得林地斑块重要值降低,就单一斑块而言,随着景观距离阈值的增大,分割成的小斑块的重要值降低程度在逐渐减小。  相似文献   

18.
Connectivity losses lead to a reduction of the amount of habitat resources that can be reached and used by species, and hence to a decline in the ranges and abundance of multiple taxa. Despite the recognized important role of small habitat patches for many species inhabiting fragmented landscapes, their potential contribution as stepping stones for maintaining overall landscape connectivity has received less attention. Using connectivity metrics based on a graph-theoretic approach we (i) quantified the connectivity of grassland patches in a sector of the Pampa region in Argentina, using a range of dispersal distances (from 100 to 10,000 m) representative of the scale of dispersal of different species; (ii) identified the most relevant patches for maintaining overall connectivity; and (iii) studied the importance of small patches (defined for different area thresholds of 5, 20, and 50 ha) as connectivity providers in the landscape. Although grassland patches were in general poorly connected at all distances, some of them were critical for overall connectivity and were found to play different crucial roles in the patch network. The location of small patches in the grassland network allowed them to function as stepping stones, yielding significant connectivity gains for species that move large distances (>5000 m) for the three area thresholds considered. Thus, under the spatial pattern of the studied landscape, species that move long distances would benefit from stepping stones, while less mobile organisms would benefit from, and mostly rely on the largest patches. We recommend that future management activities should (i) aim at preserving the grassland patches with the highest potential as stepping stones to promote landscape-level connectivity; and (ii) pay more attention to the conservation of key small patches, particularly given that usually they are those more vulnerable to land clearing for agriculture.  相似文献   

19.
Landscape geometry determines community response to disturbance   总被引:1,自引:0,他引:1  
Ecological communities are impacted by anthropogenic changes in both habitat geometry (i.e. amount, shape, fragmentation and connectivity of habitat) and disturbance regime. Although the effect of each of these drivers on diversity is well-documented, few studies have considered how habitat geometry and disturbance interact to affect diversity. We used a miniature landscape of moss patches to experimentally manipulate both habitat geometry and disturbance frequency on microarthropod communities. Species richness and abundance in local patches declined linearly with disturbance rate in all experimental landscapes, but the speed of this decline (a measure of ecological resilience) depended on the size and connectivity of the surrounding region. Reductions in region size had little effect on community resilience to disturbance until habitat loss resulted in complete loss of connectivity between patches, suggesting a threshold in community response to habitat loss. Beyond this threshold, repeated disturbance resulted in rapid declines in patch species richness and abundance and substantial changes in community composition. These effects of habitat geometry and disturbance on diversity were scale-dependent. Gamma (regional-scale) diversity was unaffected by habitat geometry, suggesting experimental reductions in alpha (local-scale) diversity were offset by increases in beta diversity. There was no effect of body size, abundance, or trophic position in determining species response to disturbance. Taxonomic grouping had a weak effect, with oribatids less affected by drought. We conclude that, in this system, dispersal from the surrounding metacommunity is vital in allowing recovery of local communities from disturbance. When habitat loss and fragmentation disrupt this process, extinctions result. Studies that examine separately the effects of habitat alterations and disturbance on diversity may therefore underestimate their combined effects.  相似文献   

20.
Habitat fragmentation is a threat to biodiversity because it restricts the ability of animals to move. Maintaining landscape connectivity could promote connections between habitat patches, which is extremely important for the preservation of gene flow and population viability. This paper aims to evaluate the landscape connectivity of forest areas as it relates to the conservation of the Yunnan snub-nosed monkey (Rhinopithecus bieti), an emblematic and endemic endangered primate species. Specifically, this study seeks to model ways to improve connectivity via cropland reforestation scenarios which incorporate graph theory and genetic distances. The connectivity improvement assessment is performed at two nested scales. At the regional scale, the aim is to quantitatively assess the gain in connectivity from different reforestation scenarios, in which croplands are replaced by different kinds of forest habitats. At the local scale, the goal is to prioritize and to locate croplands based on the gain in connectivity that they would provide if they were reforested. The results indicate that the four reforestation scenarios have different impacts on connectivity; the fourth scenario, in which reforestation is accomplished with plant species that provide optimal monkey habitat, yields the greatest increase in connectivity (+24% versus less than +2% for the others). Prioritization of the 1482 cropland patches shows that the 10 best patches increase connectivity from 0.04% to 9.1% as the isolation threshold distance increases. This kind of graph theoretic approach appears to be a useful tool for connectivity assessment and the development of conservation measures for species impacted by human activities.  相似文献   

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