Beyond the least-cost path: evaluating corridor redundancy using a graph-theoretic approach

The impact of the landscape matrix on patterns of animal movement and population dynamics has been widely recognized by ecologists. However, few tools are available to model the matrix’s influence on the length, relative quality, and redundancy of dispersal routes connecting habitat patches. Many GIS software packages can use land use/land cover maps to identify the route of least resistance between two points—the least-cost path. The limitation of this type of analysis is that only a single path is identified, even though alternative paths with comparable costs might exist. In this paper, we implemented two graph theory methods that extend the least-cost path approach: the Conditional Minimum Transit Cost (CMTC) tool and the Multiple Shortest Paths (MSPs) tool. Both methods enable the visualization of multiple dispersal routes that, together, are assumed to form a corridor. We show that corridors containing alternative dispersal routes emerge when favorable habitat is randomly distributed in space. As clusters of favorable habitat start forming, corridors become less redundant and dispersal bottlenecks become visible. Our approach is illustrated using data from a real landscape in the Brazilian Atlantic forest. We explored the effect of small, localized disturbance on dispersal routes linking conservation units. Simulated habitat destruction caused the appearance of alternative dispersal routes, or caused existing corridors to become narrower. These changes were observed even in the absence of significant differences in the length or cost of least-cost paths. Last, we discuss applications to animal movement studies and conservation initiatives.     

Improved connectivity analysis using multiple low-cost paths to evaluate habitat for the endangered San Martin titi monkey (Plecturocebus oenanthe) in north-central Peru

Context

Graph-theoretic evaluations of habitat connectivity often rely upon least-cost path analyses to evaluate connectedness of habitat patches, based on an underlying cost surface. We present two improvements upon these methods.

Objectives

As a case study to test these methods, we evaluated habitat connectivity for the endangered San Martin titi monkey (Plecturocebus oenanthe) in north-central Peru, to prioritize habitat patches for conservation.

Methods

First, rather than using a single least-cost path between habitat patches, we analyzed multigraphs made up of multiple low-cost paths. This allows us to differentiate between patches connected through a single narrow corridor, and patches connected by a wide swath of traversable land. We evaluate potential movement pathways by iteratively removing paths and recomputing connectivity metrics. Second, instead of performing a sensitivity analysis by varying costs uniformly across the landscape, we generated landscapes with spatially varying costs.

Results

This approach produced a more informative assessment of connectivity than standard graph analyses. Of the 4340 habitat patches considered across the landscape, we identified the most important 100, those frequently ranked highly through repeated network modifications, for multiple metrics and cost surfaces.

Conclusions

These methods represent a novel approach for assessing connectivity, better accounting for spatial configurations of habitat patches and uncertainty in cost surfaces. The ability to identify habitat patches with more possible routes to other patches is of interest for resiliency planning and prioritization in the face of continued habitat loss and climate change. These methods should be broadly applicable to conservation planning for other wildlife species.

     

Animal behavior, cost-based corridor models, and real corridors

Corridors are popular conservation tools because they are thought to allow animals to safely move between habitat fragments, thereby maintaining landscape connectivity. Nonetheless, few studies show that mammals actually use corridors as predicted. Further, the assumptions underlying corridor models are rarely validated with field data. We categorized corridor use as a behavior, to identify animal-defined corridors, using movement data from fishers (Martes pennanti) tracked near Albany, New York, USA. We then used least-cost path analysis and circuit theory to predict fisher corridors and validated the performance of all three corridor models with data from camera traps. Six of eight fishers tracked used corridors to connect the forest patches that constitute their home ranges, however the locations of these corridors were not well predicted by the two cost-based models, which together identified only 5 of the 23 used corridors. Further, camera trap data suggest the cost-based corridor models performed poorly, often detecting fewer fishers and mammals than nearby habitat cores, whereas camera traps within animal-defined corridors recorded more passes made by fishers, carnivores, and all other non-target mammal groups. Our results suggest that (1) fishers use corridors to connect disjunct habitat fragments, (2) animal movement data can be used to identify corridors at local scales, (3) camera traps are useful tools for testing corridor model predictions, and (4) that corridor models can be improved by incorporating animal behavior data. Given the conservation importance and monetary costs of corridors, improving and validating corridor model predictions is vital.     

Connecting mountains and desert valleys for black bears in northern Mexico

Context

Black bear connectivity studies are scarce in the southern distribution where the species is endangered. The identification of corridors is a strategy to promote conservation in human-modified landscapes.

Objectives

Assess and validate long-distance corridors in the southern black bear distribution using resistance models, occurrence records, and radio-telemetry of an individual that dispersed between the Sierras Madres of Mexico.

Methods

We acquired black bear occurrence records from several sources and telemetry records from one dispersal individual in northern Mexico. We generated ensemble habitat suitability models and resistance landscape surfaces to generate cumulative resistant kernel and least-cost paths to identify connectivity core areas and corridors of importance through Natural Protected Areas. Finally, we assessed long-distance corridors.

Results

We developed three habitat suitability models for black bears southern range; one matches the current distribution of the species. When including radio-tracking records, the landscape resistance is reduced to arid sites with low habitat suitability. We used least resistance connectivity surfaces to merge subpopulations within each Sierra Madre. The long-distance corridor models indicate narrow routes that require individuals with plastic behavioral dispersal capacity. Almost 20% of the connectivity core areas are within Natural Protected Areas. These are the first large-scale corridors using resistance layers in the southern black bear distribution.

Conclusions

Corridors can be functional for a range of temperate and dry habitat species. Landscape connectivity models should include the monitoring of dispersal individuals to identify the plasticity of organisms and the tangible barriers for them.

     

The sensitivity of least-cost habitat graphs to relative cost surface values

Maintaining and restoring connectivity among high-quality habitat patches is recognized as an important goal for the conservation of animal populations. To provide an efficient measure of potential connectivity pathways in heterogeneous landscapes, least-cost route analysis has been combined with graph-theoretical techniques. In this study we use spatially explicit least-cost habitat graphs to examine how matrix quality and spatial configuration influence assessments of habitat connectivity. We generated artificial landscapes comprised of three landcover types ranked consistently from low to high quality: inhospitable matrix, hospitable matrix, and habitat. We controlled the area and degree of fragmentation of each landcover in a factorial experiment for a total of 20 combinations replicated 100 times. In each landscape we compared eight sets of relative landcover qualities (cost values of 1 for habitat, between 1.5 and 150 for hospitable matrix, and 3–10,000 for inhospitable matrix). We found that the spatial location of least-cost routes was sensitive to differences in relative cost values assigned to landcover types and that the degree of sensitivity depended on the spatial structure of the landscape. Highest sensitivity was found in landscapes with fragmented habitat and between 20 and 50% hospitable matrix; sensitivity decreased as habitat fragmentation decreased and the amount of hospitable matrix increased. As a means of coping with this sensitivity, we propose identifying multiple low-cost routes between pairs of habitat patches that collectively delineate probable movement zones. These probable movement zones account for uncertainty in least-cost routes and may be more robust to variation in landcover cost values.     

Not all forests are alike: the role of commercial forest in the conservation of landscape connectivity for the giant panda

Context

The downlisting of giant panda (Ailuropoda melanoleuca) from Endangered to Vulnerable in IUCN Red List confirms the effectiveness of current conservation practices. However, future survival of giant panda is still in jeopardy due to habitat fragmentation and climate change. Maintaining movement corridors between habitat patches in the newly established Giant Panda National Park (GPNP) is the key for the long-term sustainability of the species.

Objectives

We evaluated the impacts of conversion from natural forest to plantation on giant panda habitat connectivity, which is permitted within collective forests and encouraged by the policies for the economic benefits of local communities. We modeled distribution of giant panda habitat in Minshan Mountains which harbors its largest population, and delineated movement corridors between core habitat patches under management scenarios of different forest conversion proportions.

Methods

We applied an integrated species distribution model based on inhomogeneous Poisson point process to combine presence-only data and site occupancy data, and least-cost models to identify potential movement corridors between core habitat patches.

Results

We found that current distribution of plantation has not damaged connectivity between core habitat patches of giant panda. However, it could be severely degraded if mass conversion occurred. Since the GPNP incorporates all the core habitats identified from our model, controlling natural forest conversion inside GPNP would maintain the movement corridors for giant panda.

Conclusions

We recommend no expansion of plantations inside the GPNP, and improving collective forest management for expansion of ecological forest in adjoining habitat patches.

     

Characterising landscape connectivity for conservation planning using a dispersal guild approach

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.

     

Least-cost path length versus accumulated-cost as connectivity measures

Least-cost modelling has become a popular method for measuring connectivity. By representing the landscape as a cost-surface, least-cost paths can be calculated that represent the route of maximum efficiency between two locations as a function of the distance travelled and the costs traversed. Both the length and the accumulated-cost of a least-cost path have been used as measures of connectivity between pairs of locations. However, we are concerned that in some situations the length of a least-cost path may provide a misleading measure of connectivity as it only accounts for the distance travelled while ignoring the costs traversed, and results in a measure that may be little better than Euclidean distance. Through simulations using fractal landscapes we demonstrate that least-cost path length is often highly correlated with Euclidean distance. This indicates that least-cost path length provides a poor measure of connectivity in many situations, as it does not capture sufficient information about the ecological costs to movement represented by the cost-surface. We recommend that in most situations the accumulated-cost of a least-cost path provides a more appropriate measure of connectivity between locations as it accounts for both the distance travelled and costs traversed, and that the generation of vector least-cost paths should be reserved for visualisation purposes.     

Examining speed versus selection in connectivity models using elk migration as an example

Context

Landscape resistance is vital to connectivity modeling and frequently derived from resource selection functions (RSFs). RSFs estimate relative probability of use and tend to focus on understanding habitat preferences during slow, routine animal movements (e.g., foraging). Dispersal and migration, however, can produce rarer, faster movements, in which case models of movement speed rather than resource selection may be more realistic for identifying habitats that facilitate connectivity.

Objective

To compare two connectivity modeling approaches applied to resistance estimated from models of movement rate and resource selection.

Methods

Using movement data from migrating elk, we evaluated continuous time Markov chain (CTMC) and movement-based RSF models (i.e., step selection functions [SSFs]). We applied circuit theory and shortest random path (SRP) algorithms to CTMC, SSF and null (i.e., flat) resistance surfaces to predict corridors between elk seasonal ranges. We evaluated prediction accuracy by comparing model predictions to empirical elk movements.

Results

All connectivity models predicted elk movements well, but models applied to CTMC resistance were more accurate than models applied to SSF and null resistance. Circuit theory models were more accurate on average than SRP models.

Conclusions

CTMC can be more realistic than SSFs for estimating resistance for fast movements, though SSFs may demonstrate some predictive ability when animals also move slowly through corridors (e.g., stopover use during migration). High null model accuracy suggests seasonal range data may also be critical for predicting direct migration routes. For animals that migrate or disperse across large landscapes, we recommend incorporating CTMC into the connectivity modeling toolkit.

     

A comparison of metrics predicting landscape connectivity for a highly interactive species along an urban gradient in Colorado, USA

Many organisms persist in fragmented habitat where movement between patches is essential for long-term demographic and genetic stability. In the absence of direct observation of movement, connectivity or isolation metrics are useful to characterize potential patch-level connectivity. However, multiple metrics exist at varying levels of complexity, and empirical data on species distribution are rarely used to compare performance of metrics. We compared 12 connectivity metrics of varying degrees of complexity to determine which metric best predicts the distribution of prairie dog colonies along an urban gradient of 385 isolated habitat patches in Denver, Colorado, USA. We found that a modified version of the incidence function model including area-weighting of patches and a cost-weighted distance surface best predicted occupancy, where we assumed roads were fairly impermeable to movement, and low-lying drainages provided dispersal corridors. We also found this result to be robust to a range of cost weight parameters. Our results suggest that metrics should incorporate both patch area and the composition of the surrounding matrix. These results provide guidance for improved landscape habitat modeling in fragmented landscapes and can help identify target habitat for conservation and management of prairie dogs in urban systems.     

Integrating GIS and homing experiments to study avian movement costs

Forest cover reduction may affect movements of forest animals, but resistance to animal movements in and out of forests remains unknown despite its importance for modelling. We tested whether ovenbird (Seiurus aurocapilla), a forest-interior songbird, responds similarly to the amount of forest cover while moving locally (~2 km) and over entire landscapes (~25 km). We compared spatially-explicit simulations to field data to address the issue of resistance to movement in open areas. We caught, banded and translocated 143 territorial males 0.8–27 km away from their territory early in the breeding season. Seventy-eight percent and 50% of translocated males returned (homed) within 10 days following “local” and “landscape” translocations respectively. Independent of translocation distance, homing times increased with decreasing forest in the landscape. With a Geographic Information System (GIS), we simulated “least-cost” paths that homing ovenbirds would ideally take, when resistance to movement in open areas ranged 1–1000 times the resistance to movement in forest. The length, the cumulative cost, and variability of simulated least-cost movement paths increased with increasing resistance in open areas. With landscape translocations, least-cost path length explained homing time better than Euclidean distance, and based on an information-theoretic approach, resistance to movement was estimated to be 27 times greater in open areas than in forests (95% confidence interval: 16–45). However, least-cost path length did not perform better than Euclidean distance with local translocations, and the cumulative cost of least-cost paths was not associated to homing time in either translocation scale. We conclude that resistance to animal movements in open areas can be addressed by a combination of GIS modelling and translocation experiments, and is between one and two orders of magnitude greater than resistance to movements in forests, in the case of ovenbirds.     

Habitat suitability modelling to correlate gene flow with landscape connectivity

Landscape connectivity is important in designing corridor and reserve networks. Combining genetic distances among individuals with least-cost path (LCP) modelling helps to correlate indirect measures of gene flow with landscape connectivity. Applicability of LCP modelling, however, is reduced if knowledge on dispersal pathways or routes is lacking. Therefore, we integrated habitat suitability modelling into LCP analysis to avoid the subjectivity common in LCP analyses lacking knowledge on dispersal pathways or routes. We used presence-only data and ecological niche factor analysis to model habitat suitability for the spiny rat, Niviventer coninga, in a fragmented landscape of western Taiwan. We adapted the resultant habitat suitability map for incorporation into LCP analyses. Slightly increased Mantel correlations indicated that a class-weighted suitability map better explained genetic distances among individuals than did geographical distances. The integration of habitat suitability modelling into LCP analysis can thus generate information on distribution of suitable habitats, on potential routes of dispersal, for placement of corridors, and evaluate landscape connectivity. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Circuit theory emphasizes the importance of edge-crossing decisions in dispersal-scale movements of a forest passerine

Measuring landscape connectivity in ways that reflect an animal’s propensity or reluctance to move across a given landscape is key for planning effective conservation strategies. Resistance distance, based on circuit theory, is one such measure relevant for modeling how broad-scale animal movements over long time periods may lead to gene flow across the landscape. Despite the success of circuit theory in landscape genetic studies, its applicability to model finer-scale processes such as the movement patterns of individual animals within their breeding grounds (e.g., while prospecting for territories) has yet to be tested. Here, we applied both circuit models and least-cost models to understand the relationship between landscape connectivity and return time of Ovenbirds (Seiurus aurocapilla) that had been translocated at least 20 km from their home territory near Québec City, Canada. Using an iterative optimization process, we derived resistance values for three cover types (forest, edge, and open) that resulted in resistance distance values that best explained Ovenbird return times. We also identified the cover-type resistance values that yielded length of least-cost path estimates that best explained return times of the translocated birds. The circuit theory and least-cost path methods were equally supported by the data despite being based on different sets of resistance values. The optimal resistance values for calculating resistance distance indicated that for Ovenbirds, traversing a given distance of edge habitat presented a substantially greater resistance than that of open areas. On the other hand, optimized resistances of edge and open were very similar for calculating length of least-cost path. The circuit theory approach suggested that for an Ovenbird moving through fragmented habitat, the number of forest-open transitions (i.e., edge-crossings) that an individual must make is critical to understanding return times after translocation. The least-cost path approach, on the other hand, suggested that the birds strongly avoid all open areas, regardless of size. Circuit theory offers an important new approach for understanding landscapes from the perspective of individuals moving within their breeding range, at finer spatial scales and shorter time scales than have been previously considered.     

Influence of separating home range and dispersal movements on characterizing corridors and effective distances

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.

     

Thresholds in seascape connectivity: influence of mobility, habitat distribution, and current strength on fish movement

Assessing connectivity of the marine environment is a fundamental challenge for marine conservation and planning, yet conceptual development in habitat connectivity has been based on terrestrial examples rather than marine ecosystems. Here, we explore differences in marine environments that could affect localized movement of marine organisms and demonstrate the importance of incorporating them into seascape models. We link a fish-based cost surface model to simulated seascapes to test hypotheses about the effects of fish mobility, water current strength, and their interactions on functional connectivity of a seascape. Our models predict that sedentary fish should be more sensitive to habitat change than more mobile fish. Furthermore, highly mobile fish should be more sensitive to water currents than habitat change. In our models, the cost of swimming against a current (of any strength) exceeded its benefits, resulting in overall decreases in connectivity with increasing current strengths. We further hypothesized that thresholds in functional connectivity will be affected by both fish mobility and water current strength. Connectivity thresholds in the models occurred when 10–50 % of benthic habitat was favourable; below these thresholds there was a rapid increase in path cost. Thresholds were influenced by the interaction of relative habitat costs (simulated fish mobility) and habitat fragmentation: thresholds for less mobile fish (higher relative cost) were reached at lower habitat abundance when habitat was fragmented, while thresholds for mobile fish were less affected by fragmentation. Our approach suggests mobility and water current are useful indicators of connectivity in marine environments and should be incorporated in seascape models.     

A method for building corridors in spatial conservation prioritization

We introduce a novel approach to building corridors in spatial conservation prioritization. The underlying working principle is the use of a penalty structure in an iterative algorithm used for producing a spatial priority ranking. The penalty term aims to prevent loss or degradation of structural connections, or, equivalently, to promote to a higher rank landscape elements that are required to keep networks connected. The proposed method shows several convenient properties: (1) it does not require a priori specification of habitat patches, end points or related thresholds, (2) it does not rely on resistance coefficients for different habitats, (3) it does not require species targets, and (4) the cost of additional connectivity via corridors can be quantified in terms of habitat quality lost across species. Corridor strength and width parameters control the trade-off between increased structural connectivity via corridors and other considerations relevant to conservation planning. Habitat suitability or dispersal suitability layers used in the analysis can be species specific, thus allowing analysis both in terms of structural and functional connectivity. The proposed method can also be used for targeting habitat restoration, by identifying areas of low habitat quality included in corridors. These methods have been implemented in the Zonation software, and can be applied to large scale and high resolution spatial prioritization, effectively integrating corridor design and spatial conservation prioritization. Since the method operates on novel principles and combines with a large number of features already operational in Zonation, we expect it to be of utility in spatial conservation planning.     

Are hedgerows effective corridors between fragments of woodland habitat? An evidence-based approach

Anthropogenic modification of the countryside has resulted in much of the landscape consisting of fragments of once continuous habitat. Increasing habitat connectivity at the landscape-scale has a vital role to play in the conservation of species restricted to such remnant patches, especially as species may attempt to track zones of habitat that satisfy their niche requirements as the climate changes. Conservation policies and management strategies frequently advocate corridor creation as one approach to restore connectivity and to facilitate species movements through the landscape. Here we examine the utility of hedgerows as corridors between woodland habitat patches using rigorous systematic review methodology. Systematic searching yielded 26 studies which satisfied the review inclusion criteria. The empirical evidence currently available is insufficient to evaluate the effectiveness of hedgerow corridors as a conservation tool to promote the population viability of woodland fauna. However, the studies did provide anecdotal evidence of positive local population effects and indicated that some species use hedgerows as movement conduits. More replicated and controlled field investigations or long-term monitoring are required in order to allow practitioners and policy makers to make better informed decisions about hedgerow corridor creation and preservation. The benefits of such corridors in regard to increasing habitat connectivity remain equivocal, and the role of corridors in mitigating the effects of climate change at the landscape-scale is even less well understood.     

Prioritization of habitat patches for landscape connectivity conservation differs between least-cost and resistance distances

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.

     

Independent effects of connectivity predict homing success by northern flying squirrel in a forest mosaic

Landscape composition and configuration, often termed as habitat loss and fragmentation, are predicted to reduce species population viability, partly due to the restriction of movement in the landscape. Unfortunately, measuring the effects of habitat loss and fragmentation on functional connectivity is challenging because these variables are confounded, and often the motivation for movement by target species is unknown. Our objective was to determine the independent effects of landscape connectivity from the perspective of a mature forest specialist—the northern flying squirrel (Glaucomys sabrinus). To standardize movement motivation, we translocated 119 squirrels, at varying distances (0.18–3.8 km) from their home range across landscapes representing gradients in both habitat loss and fragmentation. We measured the physical connectedness of mature forest using an index of connectivity (landscape coincidence probability). Patches were considered connected if they were within the mean gliding distance of a flying squirrel. Homing success increased in landscapes with a higher connectivity index. However, homing time was not strongly predicted by habitat amount, connectivity index, or mean nearest neighbour and was best explained as a simple function of sex and distance translocated. Our study shows support for the independent effects of landscape configuration on animal movement at a spatial scale that encompasses several home ranges. We conclude that connectivity of mature forest should be considered for the conservation of some mature forest specialists, even in forest mosaics where the distinction between habitat and movement corridors are less distinct.     

Connectivity providers for semi-aquatic vertebrates: the case of the endangered otter in Italy

Modeling habitat connectivity for conservation of semi-aquatic vertebrates is a particularly challenging task, due to the fine-scale and linear distribution of riverine habitats and to the capacity of species to move both on freshwater and terrestrial realms. We showed how the integrated analytical framework provided by the habitat availability (reachability) metrics and their fractions can be used to effectively evaluate the distinctive roles and contributions of both habitat patches (aquatic and riparian) and linkage areas (permeable land matrix) to the connectivity and functioning of a complex system composed of multiple river catchments. Analysis focused on the Eurasian otter (Lutra lutra L.), one of the most endangered mammals in Italy. We developed a network connectivity model based on suitable otter habitats and multiple least-cost paths between catchments. A graph analytical approach was used to identify critical nodes and links for the potential expansion and long-term viability of the species in the region. Our results showed that few basins concentrate most of the importance for sustaining the overall habitat connectivity, due to the extension of suitable habitats they contain, their strong connections with other basins, and their importance as stepping stones that uphold ecological fluxes between otherwise weakly connected habitats. The potential contribution of each basin to enhance the dispersal and expansion of otters in the area strongly depended on the key functional paths (sequences of links and nodes) among the catchments. We identified vacant basins that could be colonized by otters in the near future, and connecting areas in the intermediate matrix that might be preferentially used to conduct and promote dispersal movements and gene flow in the area. The novel approach here adopted could be easily extended to other semiaquatic species and catchment systems, offering a management strategy to preserve the hydrographic network as an integrated system, as well as a joint evaluation of the role of both the river courses and the matrix in between in a single landscape model.