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1.
2.

Context

A challenge devising revegetation strategies in fragmented landscapes is conserving for the widest spectrum of biodiversity. Habitat network reconstruction should improve landscape capacity to maintain species populations. However, the location of revegetation often fails to account for species occurrence and dispersal processes operating across spatial scales.

Objectives

Our objective was to integrate metapopulation theory with estimates of landscape capacity and dispersal pathways to highlight connectivity gaps. Maintenance of populations could thereby be facilitated through reconnecting habitat networks across regional and broader scales, with assumed benefit for the dispersal needs of less sensitive species.

Methods

Predicted occupancy and metapopulation capacity were calculated for a generic focal species derived from fragmentation-sensitive woodland birds, mammals and reptiles. A metapopulation connectivity analysis predicted regional dispersal links to identify likely routes through which individuals may move to contribute to the viability of the population. We used the revegetation programmes of the Brigalow–Nandewar Biolinks project, eastern New South Wales, Australia, to demonstrate our approach.

Results

Landscape capacity of the current landscape varied across the region. Low-value links between populations provided greatest opportunities for revegetation and improved landscape capacity. Where regional connectivity did not indicate a pathway between populations, broader scale connectivity provided guidance for revegetation.

Conclusions

The metapopulation-based model, coupled with a habitat dispersal network analysis, provided a platform to inform revegetation locations and better support biodiversity. Our approach has application for directing on-ground action to support viable populations, assess the impact of revegetation schemes or monitor the progress of staged implementations.
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3.

Context

Corridors are usually delineated as areas of minimum cumulative resistance to movement through a resistance surface and characterized by their effective distance (accumulated resistance along the least-cost path). The results of these assessments depend on resistance values, which are typically derived from the inverse of habitat suitability models or from presence data of individuals within their home ranges, rather than from data on dispersal or exploratory movements.

Objective

Evaluate the extent to which corridor delineation and effective distance estimates may vary depending on whether home range locations or dispersal data are used to characterize species habitat selection and landscape resistance to movement.

Methods

We analyzed a large telemetry dataset (GPS collars) for the endangered Iberian lynx. We modeled corridors and effective distances three ways: (1) considering only GPS locations within home ranges, (2) considering only locations in dispersal or exploratory movements outside home ranges, and (3) considering all locations together.

Results

Delineated least-cost corridors followed similar trajectories and sometimes overlapped in the three models. The estimated effective distances were 42 % lower in the dispersal-based model than in the model based solely on home range use.

Conclusions

Models derived exclusively from locations within home ranges may provide lower connectivity estimates than models derived from dispersal locations, affecting estimates of resistance to move between habitat areas, even when the most likely movement routes are similar. Although dispersal data are costly to gather, they potentially provide more realistic assessments of the actual isolation of populations in heterogeneous landscapes.
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4.

Context

Land use changes have modified the extent and structure of native vegetation, resulting in fragmentation of native species habitat. Connectivity is increasingly seen as a requirement for effective conservation in these landscapes, but the question remains: ‘connectivity for which species?’.

Objective

The aim of this study was to develop and then apply a rapid, expert-based, dispersal guild approach where species are grouped on similar fine-scale dispersal behaviour (such as between scattered trees) and habitat characteristics.

Methods

Dispersal guilds were identified using clustering techniques to compare dispersal and habitat parameters elicited from experts. We modelled least-cost paths and corridors between patches and individual movement probabilities within these corridors for each of the dispersal guilds using Circuitscape. We demonstrate our approach with a case study in the Tasmanian Northern Midlands, Australia.

Results

The dispersal guild approach grouped the 12 species into five dispersal guilds. The connectivity modelling of those five guilds found that broadly dispersing species in this landscape, such as medium-sized carnivorous mammals, were unaffected by fragmentation while from the perspective of the three dispersal guilds made up of smaller mammals, the landscape appeared highly fragmented.

Conclusions

Our approach yields biologically defensible outputs that are broadly applicable, particularly for conservation planning where data and resources are limited. It is a useful first step in multi-species conservation planning which aims to identify those species most in need of conservation efforts.
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5.

Context

Landscape and habitat filters are major drivers of biodiversity of small habitat islands by influencing dispersal and extinction events in plant metapopulations.

Objectives

We assessed the effects of landscape and habitat filters on the species richness, abundance and trait composition of grassland specialist and generalist plants in small habitat islands. We studied traits related to functional spatial connectivity (dispersal ability by wind and animals) and temporal connectivity (clonality and seed bank persistence) using model selection.

Methods

We sampled herbaceous plants, landscape (local and regional isolation) and habitat filters (inclination, woody encroachment and disturbance) in 82 grassland islands in Hungary.

Results

Isolation decreased the abundance of good disperser specialist plants due to the lack of directional vectors transferring seeds between suitable habitat patches. Clonality was an effective strategy, but persistent seed bank did not support the survival of specialist plants in isolated habitats. Generalist plants were unaffected by landscape filters due to their wide habitat breadth and high propagule availability. Clonal specialist plants could cope with increasing woody encroachment due to their high resistance against environmental changes; however, they could not cope with intensive disturbance. Steep slopes providing environmental heterogeneity had an overall positive effect on species richness.

Conclusions

Specialist plants were influenced by the interplay of landscape filters influencing their abundance and habitat filters affecting species richness. Landscape filtering by isolation influenced the abundance of specialist plants by regulating seed dispersal. Habitat filters sorted species that could establish and persist at a site by influencing microsite availability and quality.
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6.

Context

Methods quantifying habitat patch importance for maintaining habitat network connectivity have been emphasized in helping to prioritize conservation actions. Functional connectivity is accepted as depending on landscape resistance, and several measures of functional inter-patch distance have been designed. However, how the inter-patch distance, i.e., based on least-cost path or multiple paths, influences the identification of key habitat patches has not been explored.

Objectives

We compared the prioritization of habitat patches according to least-cost distance (LCD) and resistance distance (RD), using common binary and probabilistic connectivity metrics.

Methods

Our comparison was based on a generic functional group of forest mammals with different dispersal distances, and was applied to two landscapes differing in their spatial extent and fragmentation level.

Results

We found that habitat patch prioritization did not depend on distance type when considering the role of patch as contributing to dispersal fluxes. However, the role of patch as a connector facilitating dispersal might be overestimated by LCD-based indices compared with RD for short- and medium-distance dispersal. In particular, when prioritization was based on dispersal probability, the consideration of alternatives routes identified the connectors that probably provided functional connectivity for species in the long term. However, the use of LCD might help identify landscape areas that need critical restoration to improve individual dispersal.

Conclusions

Our results provide new insights about the way that inter-patch distance is viewed changes the evaluation of functional connectivity. Accordingly, prioritization methods should be carefully selected according to assumptions about population functioning and conservation aims.
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7.

Context

In modern agricultural landscapes, fragmentation of partial habitats is a significant filter for multi-habitat users, reducing local taxonomic and functional diversity. There is compelling evidence that small species are more susceptible than large species. The impact of habitat fragmentation on intraspecific body-size distribution, however, is yet unexplored.

Objectives

We tested habitat fragmentation, a major driver of pollinator loss, for its impact on intraspecific body-size distributions of solitary wild-bee species. Subsequently, we tested individual body size for its impact on pollination services.

Methods

We sampled 1272 individuals of the four most common Andrena wild bee species in 22 newly established flowering fields (0.21–0.41 ha) in Hessen, Central Germany, over two consecutive years. Study sites were located in a ca. 80 ha landscape context of increasing habitat fragmentation. We analysed the pollen loads of the most abundant species.

Results

Body size within local populations of the two medium-sized bees increased with fragmentation, suggesting intraspecific selection for higher dispersal capacity. Pollen analysis carried out for the most common species revealed that larger individuals visited a significantly smaller plant spectrum. Habitat fragmentation may thus alter pollination services without necessarily affecting species richness or composition.

Conclusions

Systematic body-size variation at the population level thus explains the considerable variability between simple community measures and ecosystem functioning. Filtering processes at the individual level require increased understanding for targeting pollination services under current and future land-use change.
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8.

Context

Environmental processes and dispersal are primary determinants of metacommunity dynamics. The relative importance of these effects may vary between species of different abundance classes, given variation in life history traits. Under high disturbance conditions, rare species may be more easily eliminated from their optimal habitats and their distribution may therefore be more heavily dependent upon dispersal from nearby habitat patches than common species.

Objectives

We tested if metacommunity dynamics vary between abundance classes in a high disturbance environment.

Methods

Standardized butterfly sampling was conducted in the urban parks of Hong Kong. To estimate the strength of environmental processes, we measured an array of environmental variables for all sampled parks. Spatial predictors were generated to estimate the effect of dispersal.

Results

For shaping common species compositions, we found environmental processes (and specifically environmental variables including floral density and surrounding woody plant cover) slightly more important than spatial processes. For rare species, only spatial processes were significant while environmental processes were insignificant. Our result contrasts previous studies in natural metacommunities, which have shown that both common and rare species compositions are shaped by environmental processes and similar variables.

Conclusions

Our results demonstrate that high disturbance conditions may inhibit rare species establishment and persistence in urban landscapes. Local habitat management may not be sufficient in conserving rare species in urban environments—spatial context and configuration should be considered in planning for biodiversity. We also highlight the utility of community deconstruction analysis in providing insights into rare species metacommunity dynamics.
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9.

Context

Seed dispersal is recognized as having profound effects on the distribution, dynamics and structure of plant populations and communities. However, knowledge of how landscape structure shapes carnivore-mediated seed dispersal patterns is still scarce, thereby limiting our understanding of large-scale plant population processes.

Objectives

We aim to determine how the amount and spatial configuration of forest cover impacted the relative abundance of carnivorous mammals, and how these effects cascaded through the seed dispersal kernels they generated.

Methods

Camera traps activated by animal movement were used for carnivore sampling. Colour-coded seed mimics embedded in common figs were used to know the exact origin of the dispersed seed mimics later found in carnivore scats. We applied this procedure in two sites differing in landscape structure.

Results

We did not find between-site differences in the relative abundance of the principal carnivore species contributing to seed dispersal patterns, Martes foina. Mean dispersal distance and the probability of long dispersal events were higher in the site with spatially continuous and abundant forest cover, compared to the site with spatially aggregated and scarcer forest cover. Seed deposition closely matched the spatial patterning of forest cover in both study sites, suggesting behaviour-based mechanisms underpinning seed dispersal patterns generated by individual frugivore species.

Conclusions

Our results provide the first empirical evidence of the impact of landscape structure on carnivore-mediated seed dispersal kernels. They also indicate that seed dispersal kernels generated strongly depend on the effect that landscape structure exerts on carnivore populations, particularly on habitat-use preferences.
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10.

Context

Animals selectively use landscapes to meet their energetic needs, and trade-offs in habitat use may depend on availability and environmental conditions. For example, habitat selection at high temperatures may favor thermal cover at the cost of reduced foraging efficiency under consistently warm conditions.

Objective

Our objective was to examine habitat selection and space use in distinct populations of moose (Alces alces). Hypothesizing that endotherm fitness is constrained by heat dissipation efficiency, we predicted that southerly populations would exhibit greater selection for thermal cover and reduced selection for foraging habitat.

Methods

We estimated individual step selection functions with shrinkage for 134 adult female moose in Minnesota, USA, and 64 in Ontario, Canada, to assess habitat selection with variation in temperature, time of day, and habitat availability. We averaged model coefficients within each site to quantify selection strength for habitats differing in forage availability and thermal cover.

Results

Moose in Ontario favored deciduous and mixedwood forest, indicating selection for foraging habitat across both diel and temperature. Habitat selection patterns of moose in Minnesota were more dynamic and indicated time- and temperature-dependent trade-offs between use of foraging habitat and thermal cover.

Conclusions

We detected a scale-dependent functional response in habitat selection driven by the trade-off between selection for foraging habitat and thermal cover. Landscape composition and internal state interact to produce complex patterns of space use, and animals exposed to increasingly high temperatures may mitigate fitness losses from reduced foraging efficiency by increasing selection for foraging habitat in sub-prime foraging landscapes.
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11.

Context

Landscape-scale population dynamics are driven in part by movement within and dispersal among habitat patches. Predicting these processes requires information about how movement behavior varies among land cover types.

Objectives

We investigated how butterfly movement in a heterogeneous landscape varies within and between habitat and matrix land cover types, and the implications of these differences for within-patch residence times and among-patch connectivity.

Methods

We empirically measured movement behavior in the Baltimore checkerspot butterfly (Euphydryas phaeton) in three land cover classes that broadly constitute habitat and two classes that constitute matrix. We also measured habitat preference at boundaries. We predicted patch residence times and interpatch dispersal using movement parameters estimated separately for each habitat and matrix land cover subclass (5 categories), or for combined habitat and combined matrix land cover classes (2 categories). We evaluated the effects of including edge behavior on all metrics.

Results

Overall, movement was slower within habitat land cover types, and faster in matrix cover types. Butterflies at forest edges were biased to remain in open areas, and connectivity and patch residence times were most affected by behavior at structural edges. Differences in movement between matrix subclasses had a greater effect on predictions about connectivity than differences between habitat subclasses. Differences in movement among habitat subclasses had a greater effect on residence times.

Conclusions

Our findings highlight the importance of careful classification of movement and land cover in heterogeneous landscapes, and reveal how subtle differences in behavioral responses to land cover can affect landscape-scale outcomes.
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12.

Context

Native vegetation is often used as a proxy for habitat to estimate habitat availability in landscapes. This approach may lead to incorrect estimates of the impacts of habitat loss and fragmentation on species, which have not been thoroughly quantified so far.

Objectives

We quantified to what extent the loss of native vegetation reflect actual habitat loss by native species in landscapes. We tested the hypothesis that habitat availability declines at greater rates than native vegetation and thus is overestimated when it is quantified on the basis of native vegetation.

Methods

Using simulations, we quantified how the loss of native vegetation in artificial and real landscapes affects habitat availability for species with different habitat requirements. We contrasted a generalist species, which uses all native vegetation, with 10 habitat-specialist species classified into three categories (interior, patchy and riparian species).

Results

Habitat availability generally declined at greater rates than native vegetation for all specialist species. This pattern was apparent for different specialist species in a broad range of landscape types. Interior species always lost habitat availability more rapidly than the generalist species. Most riparian species lost habitat availability more rapidly than the generalist species. Responses of patchy species were more complex, depending on their dispersal abilities and landscape structure.

Conclusions

Habitat availability is likely to be overestimated when native vegetation is used as proxy for habitat, because habitat availability will generally decline at greater rates than native vegetation. Therefore, a species-centered approach should be adopted when estimating habitat availability in landscapes.
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13.

Context

Habitat loss and habitat fragmentation negatively affect amphibian populations. Roads impact amphibian species through barrier effects and traffic mortality. The landscape variable ‘accessible habitat’ considers the combined effects of habitat loss and roads on populations.

Objectives

The aim was to test whether accessible habitat was a better predictor of amphibian species richness than separate measures of road effects and habitat loss. I assessed how accessible habitat and local habitat variables determine species richness and community composition.

Methods

Frog and tadpole surveys were conducted at 52 wetlands in a peri-urban area of eastern Australia. Accessible habitat was delineated using a highway. Regressions were used to examine relationships between species richness and eleven landscape and local habitat variables. Redundancy analysis was used to examine relationships between community composition and accessible habitat and local habitat variables.

Results

Best-ranked models of species richness included both landscape and local habitat variables. There were positive relationships between species richness and accessible habitat and distance to the highway, and uncertain relationships with proportion cover of native vegetation and road density. There were negative relationships between species richness and concreted wetlands and wetland electrical conductivity. Four species were positively associated with accessible habitat, whereas all species were negatively associated with wetland type.

Conclusions

Barrier effects caused by the highway and habitat loss have negatively affected the amphibian community. Local habitat variables had strong relationships with species richness and community composition, highlighting the importance of both availability and quality of habitat for amphibian conservation near major roads.
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14.

Context

The problem of how ecological mechanisms create and interact with patterns across different scales is fundamental not only for understanding ecological processes, but also for interpretations of ecological dynamics and the strategies that organisms adopt to cope with variability and cross-scale influences.

Objectives

Our objective was to determine the consistency of the role of individual habitat patches in pattern-process relationships (focusing on the potential for dispersal within a network of patches in a fragmented landscape) across a range of scales.

Methods

Network analysis was used to assess and compare the potential connectivity and spatial distribution of highland fynbos habitat in and between protected areas of the Western Cape of South Africa. Connectivity of fynbos patches was measured using ten maximum threshold distances, ranging from five to 50 km, based on the known average dispersal distances of fynbos endemic bird species.

Results

Network connectivity increased predictably with scale. More interestingly, however, the relative contributions of individual protected areas to network connectivity showed strong scale dependence.

Conclusions

Conservation approaches that rely on single-scale analyses of connectivity and context (e.g., based on data for a single species with a given dispersal distance) are inadequate to identify key land parcels. Landscape planning, and specifically the assessment of the value of individual areas for dispersal, must therefore be undertaken with a multi-scale approach. Developing a better understanding of scaling dependencies in fragmenting landscapes is of high importance for both ecological theory and conservation planning.
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15.

Context

Methods for measuring restoration success that include functional connectivity between species’ populations are rare in landscape ecology and restoration practices. We developed an approach that analyzes connectivity between populations of target species and their dispersal probabilities to assess restoration success based on easily accessible input data. Applying this method to landscape development scenarios can help optimize restoration planning.

Objectives

We developed an assessment for restoration success and restoration planning based on functional connectivity between species’ populations and spatially explicit scenarios. The method was used in a case study to test its applicability.

Methods

Based on data on available habitat, species’ occurrence and dispersal ranges, connectivity metrics and dispersal probabilities for target species are calculated using the software Conefor Sensinode. The metrics are calculated for scenarios that reflect possible changes in the landscape to provide a basis for future restoration planning. We applied this approach to floodplain meadows along the Upper Rhine for four plant species and three future scenarios.

Results

In the case study, habitats of the target species were poorly connected. Peucedanum officinale and Sanguisorba officinalis were more successful in recolonizing new habitats than Iris spuria and Serratula tinctoria. The scenarios showed that restoration of species-rich grassland was beneficial for dispersal of the target species. As expected in the agriculturally dominated study area, restoration of former arable land significantly increased dispersal probabilities.

Conclusions

In the case study, the developed approach was easily applicable and provided reasonable results. Its implementation will be helpful in decision-making for future restoration planning.
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16.

Context

Landscape changes can be an important modifier of disease. Habitat fragmentation commonly results in reduced connectivity in host populations and increased use of the remaining habitat. For environmentally transmitted parasites, this presents a possible trade-off between transmission potential at the local and global level.

Objectives

We quantify the effects of fragmentation on the transmission of an environmentally transmitted parasite, teasing apart the relative effects of habitat composition and configuration on both host movement behaviour and subsequent infection patterns.

Methods

We use a spatially-explicit epidemiological model to simulate the effects of habitat fragmentation, using, as an example, whipworm (Trichuris sp.) within a red colobus monkey population (Procolobus rufomitratus).

Results

We found that habitat fragmentation did not always lead to a trade-off between population connectivity and concentration of habitat use in host movement behaviour or in final population infection patterns. However, our simulation results suggest the spatial configuration of the remaining habitat became increasingly influential on behavioural and infection outcomes as habitat was removed. Additionally, we found common fragmentation metrics provided little ability to explain variation in propagation of infections.

Conclusions

Our results suggest an interaction between habitat configuration and composition should be considered when assessing disease related impacts of habitat fragmentation on environmentally transmitted parasites, especially in cases where habitat loss is high (≥?30%). We also propose that spatially-explicit simulations that capture a host’s response to fragmentation could aid in the development of novel landscape metrics targeted towards specific host-parasite-landscape systems.
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17.

Context

The umbrella approach applied to landscape connectivity is based on the principle that the conservation or restoration of the dispersal habitats for some species also can facilitate the movement of others. Species traits alone do not seem to be enough to identify good connectivity umbrella species, showing the need to investigate the influence of additional factors on this property.

Objectives

We test whether the potential of a species as a connectivity umbrella can be influenced by landscape composition and configuration.

Methods

We simulated movement routes for eight hypothetical species in artificial patchy landscapes with different levels of fragmentation, habitat amount and matrix permeability. We determined the effectiveness of the connectivity umbrella of the virtual species using pairwise intersections of important habitats for their movements in all landscapes.

Results

The connectivity umbrella performance of all species was affected by the interaction of fragmentation level and habitat amount. In general, species performance increased with decreasing fragmentation and increasing habitat amount. In most landscapes and considering the same dispersal threshold, species able to move more easily through the matrix showed higher umbrella performance than those for which the matrix offered greater resistance.

Conclusions

The connectivity umbrella is not a static feature that depends only on the species traits, but rather a dynamic property that also varies according to the landscape attributes. Therefore, we do not recommend spatial transferability of the connectivity umbrella species identified in a landscape to others that have divergent levels of fragmentation and habitat quantity.
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18.

Context

Dispersal is essential for species persistence and landscape genetic studies are valuable tools for identifying potential barriers to dispersal. Macaws have been studied for decades in their natural habitat, but we still have no knowledge of how natural landscape features influence their dispersal.

Objectives

We tested for correlations between landscape resistance models and the current population genetic structure of macaws in continuous rainforest to explore natural barriers to their dispersal.

Methods

We studied scarlet macaws (Ara macao) over a 13,000 km2 area of continuous primary Amazon rainforest in south-eastern Peru. Using remote sensing imagery from the Carnegie Airborne Observatory, we constructed landscape resistance surfaces in CIRCUITSCAPE based on elevation, canopy height and above-ground carbon distribution. We then used individual- and population-level genetic analyses to examine which landscape features influenced gene flow (genetic distance between individuals and populations).

Results

Across the lowland rainforest we found limited population genetic differentiation. However, a population from an intermountain valley of the Andes (Candamo) showed detectable genetic differentiation from two other populations (Tambopata) located 20–60 km away (F ST = 0.008, P = 0.001–0.003). Landscape resistance models revealed that genetic distance between individuals was significantly positively related to elevation.

Conclusions

Our landscape resistance analysis suggests that mountain ridges between Candamo and Tambopata may limit gene flow in scarlet macaws. These results serve as baseline data for continued landscape studies of parrots, and will be useful for understanding the impacts of anthropogenic dispersal barriers in the future.
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19.

Context

Many nearshore species are distributed in habitat patches connected only through larval dispersal. Genetic research has shown some spatial structure of such metapopulations and modeling studies have shed light onto possible patterns of connectivity and barriers. However, little is known about human impact on their spatial structure and patterns of connectivity.

Objectives

We examine the effects of fishing on the spatial and temporal dynamics of metapopulations of sedentary marine species (red sea urchin and red abalone) interconnected by larval dispersal.

Methods

We constructed a metapopulation model to simulate abalone and sea urchin metapopulations experiencing increasing levels of fishing mortality. We performed the modularity analysis on the yearly larval connectivity matrices produced by these simulations, and analyzed the changes of modularity and the formation of modules over time as indicators of spatial structure.

Results

The analysis revealed a strong modular spatial structure for abalone and a weak spatial signature for sea urchin. In abalone, under exploitation, modularity takes step-wise drops on the path to extinction, and modules breakdown into smaller fragments followed by module and later metapopulation collapse. In contrast, sea urchin showed high modularity variation, indicating high- and low-mixing years, but an abrupt collapse of the metapopulation under strong exploitation.

Conclusions

The results identify a disruption in larval connectivity and a pattern of collapse in highly modular nearshore metapopulations. These results highlight the ability of modularity to detect spatial structure in marine metapopulations, which varies among species, and to show early changes in the spatial structure of exploited metapopulations.
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20.

Context

Restoring or establishing corridors between residual forest patches is one of the most adopted strategies for the conservation of animal populations and ecosystem processes in fragmented landscapes.

Objectives

This study aimed to assess whether it is more effective to focus restoration actions on existing corridors or to establish habitats in other strategic areas that can create new dispersal pathways to enhance connectivity.

Methods

We considered a real agroecosystem in northern Italy, based our analyses on graph-theory and habitat availability metrics, and focused on the Hazel Dormouse as the target species. We compared the connectivity increase resulting from (i) the simulated restoration of existing priority corridors, i.e., those with significant presence of forest but in which restoration actions would still result in considerable connectivity gains, or (ii) the simulated plantation of 30 hedgerows along new priority pathways, i.e., those areas with no current forest cover in which habitat creation would be more beneficial for connectivity.

Results

Implementing new priority pathways resulted in substantially larger connectivity gains (+?38%) than when restoration efforts were concentrated in improving already existing corridors (+?11%).

Conclusions

Establishing hedgerows along new pathways allowed enhancing the complementary and functionality of the full set of landscape corridors and proved more efficient than just strengthening the areas where dispersal flows were already concentrated. We demonstrated the importance of analytical procedures able to compare the effectiveness of different management strategies for enhancing connectivity. Our approach may be applied to multiple species sensitive to fragmentation in other heterogeneous landscapes and geographical contexts.
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