Comparison and development of new graph-based landscape connectivity indices: towards the priorization of habitat patches and corridors for conservation

The loss of connectivity of natural areas is a major threat for wildlife dispersal and survival and for the conservation of biodiversity in general. Thus, there is an increasing interest in considering connectivity in landscape planning and habitat conservation. In this context, graph structures have been shown to be a powerful and effective way of both representing the landscape pattern as a network and performing complex analysis regarding landscape connectivity. Many indices have been used for connectivity analyses so far but comparatively very little efforts have been made to understand their behaviour and sensitivity to spatial changes, which seriously undermines their adequate interpretation and usefulness. We systematically compare a set of ten graph-based connectivity indices, evaluating their reaction to different types of change that can occur in the landscape (habitat patches loss, corridors loss, etc.) and their effectiveness for identifying which landscape elements are more critical for habitat conservation. Many of the available indices were found to present serious limitations that make them inadequate as a basis for conservation planning. We present a new index (IIC) that achieves all the properties of an ideal index according to our analysis. We suggest that the connectivity problem should be considered within the wider concept of habitat availability, which considers a habitat patch itself as a space where connectivity exists, integrating habitat amount and connectivity between habitat patches in a single measure.     

Quantifying landscape connectivity through the use of connectivity response curves

Habitat connectivity is an essential component of biodiversity conservation. Simulated landscapes were manipulated to quantify the impact of changes to the amount, fragmentation and dispersion of habitat on a widely applied landscape connectivity metric, the probability of connectivity index. Index results for different landscape scenarios were plotted against the dispersal distances used for their calculation to create connectivity response curves for each scenario. Understanding index response to controlled changes in landscape structure at a range of spatial scales can be used to give context to comparison of alternative landscape management scenarios. Increased amounts of habitat, decreased fragmentation and decreased inter-patch distances resulted in increased connectivity index values. Connectivity response curves demonstrated increases in assessed connectivity for scenarios with continuous corridors or “stepping stone” connectors. The sensitivity of connectivity response curves to controlled changes in landscape structure indicate that this approach is able to detect and distinguish between different types of landscape changes, but that delineation of habitat and method of quantifying dispersal probability incorporate assumptions that must be recognized when interpreting results to guide landscape management. Representing landscape connectivity in this manner allows for the impacts of alternative landscape management strategies to be compared visually through comparative plots, or statistically through the parameters that describe connectivity response curves.     

Evaluation of landscape connectivity at community level using satellite-derived NDVI

Landscape connectivity, defined as the degree to which the landscape facilitates or impedes movement among resource patches, has been considered to be a key issue for biodiversity conservation. However, the use of landscape connectivity measurements has been strongly criticised due to uncertainties in the methods used and the lack of validation. Moreover, measurements are typically restricted to the population level, whereas management is generally carried out at the community level. Here, we used satellite imagery and network metrics to predict the landscape connectivity at community level for semi-natural herbaceous patches in an urban area near Paris (France). We tested different measurement methods, both taking into account and ignoring the spatial heterogeneity of matrix resistance estimated by the normalised difference vegetation index (NDVI), and quantifying the link strength between patches with the shortest path and flow metrics. We assessed the fit of these connectivity predictions with empirical data on plant communities embedded in an urban matrix. Our results indicate that the best fit with the empirical data is obtained when the connectivity is estimated with the flow metric and takes into account the matrix heterogeneity. Overall, our study helps to estimate the landscape connectivity of urban areas and makes recommendations for ways in which we might optimise landscape planning with respect to conservation of urban biodiversity.     

Two measures of landscape-graph connectivity: assessment across gradients in area and configuration

Landscape connectivity is critical to species persistence in the face of habitat loss and fragmentation. Graph theory is a well-defined method for quantifying connectivity that has tremendous potential for ecology, but its application has been limited to a small number of conservation scenarios, each with a fixed proportion of habitat. Because it is important to distinguish changes in habitat configuration from changes in habitat area in assessing the potential impacts of fragmentation, we investigated two metrics that measure these different influences on connectivity. The first metric, graph diameter, has been advocated as a useful measure of habitat configuration. We propose a second area-based metric that combines information on the amount of connected habitat and the amount of habitat in the largest patch. We calculated each metric across gradients in habitat area and configuration using multifractal neutral landscapes. The results identify critical connectivity thresholds as a function of the level of fragmentation and a parallel is drawn between the behavior of graph theory metrics and those of percolation theory. The combination of the two metrics provides a means for targeting sites most at risk of suffering low potential connectivity as a result of habitat fragmentation.     

Habitat connectivity for pollinator beetles using surface metrics

Measuring habitat connectivity in complex landscapes is a major focus of landscape ecology and conservation research. Most studies use a binary landscape or patch mosaic model for describing spatial heterogeneity and understanding pattern-process relationships. While the value of landscape gradient approaches proposed by McGarigal and Cushman are recognized, applications of these newly proposed three dimensional surface metrics remain under-used. We created a gradient map of habitat quality from several GIS layers and applied three dimensional surface metrics to measure connectivity between 67 locations in Indiana, USA surveyed for one group of ecosystem service providers, flower longicorn beetles (Cerambycidae: Lepturinae). The three dimensional surface metrics applied to the landscape gradient model showed great potential to explain the differences of lepturine assemblages among the 2,211 studied landscapes (between site pairs). Surface kurtosis and its interaction with geographic distance were among the most important metrics. This approach provided unique information about the landscape through four configuration metrics. There were some uniform trends of the responses of many species to some of surface metrics, however some species responded differently to other metrics. We suggest that three dimensional surface metrics applied to a habitat surface map created with insight into species requirements is a valuable approach to understanding the spatial dynamics of species, guilds, and ecosystem services.     

The relative impact of forest patch and landscape attributes on black howler monkey populations in the fragmented Lacandona rainforest, Mexico

Land-use change is forcing many animal populations to inhabit forest patches in which different processes can threaten their survival. Some threatening processes are mainly related to forest patch characteristics, but others depend principally on the landscape spatial context. Thus, the impact of both patch and landscape spatial attributes needs to be assessed to have a better understanding of the habitat spatial attributes that constraint the maintenance of populations in fragmented landscapes. Here, we evaluated the relative effect of three patch-scale (i.e., patch size, shape, and isolation) and five landscape-scale metrics (i.e., forest cover, fragmentation, edge density, mean inter-patch isolation distance, and matrix permeability) on population composition and structure of black howler monkeys (Alouatta pigra) in the Lacandona rainforest, Mexico. We measured the landscape-scale metrics at two spatial scales: within 100 and 500 ha landscapes. Our findings revealed that howler monkeys were more strongly affected by local-scale metrics. Smaller and more isolated forest patches showed a lower number of individuals but at higher densities. Population density also tended to be positively associated to matrices with higher proportion of secondary forests and arboreal crops (i.e. with greater permeability), most probably because these matrices can offer supplementary foods. The immature-to-female ratio also increased with matrix permeability, shape complexity, and edge density; habitat characteristics that can increase landscape connectivity and sources availability. The prevention of habitat loss and isolation, and the increment of matrix permeability are therefore needed for the conservation of this endangered Neotropical mammal.     

Connectivity measures: a review

One of the central problems in contemporary ecology and conservation biology is the drastic change of landscapes induced by anthropogenic activities, resulting in habitat loss and fragmentation. For many wild living species, local extinctions of fragmented populations are common and recolonization is critical for regional survival. Successful recolonization depends on the availability of dispersing individuals and the degree of landscape connectivity. The obvious implications of landscape connectivity for conservation biology have led to a proliferation of connectivity measures. However, general relationships between landscape connectivity and landscape structure are lacking, and so are the relationships between different connectivity metrics. Consequently, there is a need to develop landscape metrics that more accurately characterize the landscape with an emphasis on the underlying processes. Here we review various definitions of landscape connectivity, explain their mathematical connotations, and make some unifying conclusions and suggestions for future research.     

Interactive effects of area and connectivity on the diversity of tachinid parasitoids in highly fragmented landscapes

Although many empirical and theoretical studies have elucidated the effects of habitat fragmentation on the third trophic level, little attention has been paid to the impacts of this driver on more generalist groups of non-hymenopteran parasitoids. Here, we used the highly-diverse group of tachinid flies as an alternative model to test the effects of landscape fragmentation on insect parasitoids. Our aims were: (i) to evaluate the relative importance of habitat area and connectivity losses and their potential interaction on tachinid diversity, (ii) to test whether the effects of habitat fragmentation changes seasonally, and (iii) to further assess the effect of habitat diversity on tachinid diversity and whether different parasitoid-host associations modify the species richness response to fragmentation. In 2012 a pan-trap sampling was conducted in 18 semi-natural grasslands embedded in intensive agricultural landscapes along statistically orthogonal gradients of habitat area, connectivity and habitat diversity. We found an interaction between habitat area and connectivity indicating that tachinid abundance and species richness were more negatively affected by habitat loss in landscapes with low rather than with relatively large habitat connectivity. Although tachinid communities exhibited large within-year species turnover, we found that the effects of landscape fragmentation did not change seasonally. We found that habitat diversity and host association did not affect tachinid species diversity. Our results have important implications for biodiversity conservation as any attempts to mitigate the negative effects of habitat loss need to take the general level of habitat connectivity in the landscape into account.     

Multiscale effects of habitat and surrounding matrices on waterbird diversity in the Yangtze River Floodplain

Context

With the expansion in urbanization, understanding how biodiversity responds to the altered landscape becomes a major concern. Most studies focus on habitat effects on biodiversity, yet much less attention has been paid to surrounding landscape matrices and their joint effects.

Objective

We investigated how habitat and landscape matrices affect waterbird diversity across scales in the Yangtze River Floodplain, a typical area with high biodiversity and severe human-wildlife conflict.

Methods

The compositional and structural features of the landscape were calculated at fine and coarse scales. The ordinary least squares regression model was adopted, following a test showing no significant spatial autocorrelation in the spatial lag and spatial error models, to estimate the relationship between landscape metrics and waterbird diversity.

Results

Well-connected grassland and shrub surrounded by isolated and regular-shaped developed area maintained higher waterbird diversity at fine scales. Regular-shaped developed area and cropland, irregular-shaped forest, and aggregated distribution of wetland and shrub positively affected waterbird diversity at coarse scales.

Conclusions

Habitat and landscape matrices jointly affected waterbird diversity. Regular-shaped developed area facilitated higher waterbird diversity and showed the most pronounced effect at coarse scales. The conservation efforts should not only focus on habitat quality and capacity, but also habitat connectivity and complexity when formulating development plans. We suggest planners minimize the expansion of the developed area into critical habitats and leave buffers to maintain habitat connectivity and shape complexity to reduce the disturbance to birds. Our findings provide important insights and practical measures to protect biodiversity in human-dominated landscapes.

     

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.     

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.     

Can we face different types of storms under the same umbrella? Efficiency and consistency of connectivity umbrellas across different patchy landscape patterns

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.

     

A scale-sensitive connectivity analysis to identify ecological networks and conservation value in river networks

Existing methods for connectivity analysis still encounter difficulties in explaining functional relationships between network structure and ecological patterns over larger territories or complex structures like dendritic river networks. We propose a method that addresses the problem of scale and resolution in the connectivity analysis of dendritic network structures, illustrated here for the re-colonization of the French Loire river basin by the European otter. The ecological niche factor approach is applied to infer favourable habitat in the river network based on large scale data of land use and hydro-morphology of river segments for the entire river basin. These analyses identified the stressors to the riparian zone of channel straightening, urbanisation and forest fragmentation as the principal factors explaining otter occurrence. Using this estimate of habitat favourability, we used the Integral Index of Connectivity to quantify habitat availability and connectivity in the dendritic river network. When we calculate the integral index of connectivity over different spatial extents by constraining network distances, the scale-sensitivity of the network’s connectivity emerges. Accounting for high mobility by entering larger network distances in the analysis identifies conservation networks and priorities mainly in downstream parts of the river basin, whereas with smaller network distances, more restricted high quality areas in central and upstream parts are highlighted. The presented approach performed better than distribution modelling approaches in explaining species occurrence over the river network and confirms the crucial aspect of connectivity in otter re-colonization.     

How should we measure landscape connectivity?

The methods for measuring landscape connectivity have never been compared or tested for their responses to habitat fragmentation. We simulated movement, mortality and boundary reactions across a wide range of landscape structures to analyze the response of landscape connectivity measures to habitat fragmentation. Landscape connectivity was measured as either dispersal success or search time, based on immigration into all habitat patches in the landscape. Both measures indicated higher connectivity in more fragmented landscapes, a potential for problematic conclusions for conservation plans. We introduce cell immigration as a new measure for landscape connectivity. Cell immigration is the rate of immigration into equal-sized habitat cells in the landscape. It includes both within- and between-patch movement, and shows a negative response to habitat fragmentation. This complies with intuition and existing theoretical work. This method for measuring connectivity is highly robust to reductions in sample size (i.e., number of habitat cells included in the estimate), and we hypothesize that it therefore should be amenable to use in empirical studies. The connectivity measures were weakly correlated to each other and are therefore generally not comparable. We also tested immigration into a single patch as an index of connectivity by comparing it to cell immigration over the landscape. This is essentially a comparison between patch-scale and landscape-scale measurement, and revealed some potential for patch immigration to predict connectivity at the landscape scale. However, this relationship depends on the size of the single patch, the dispersal characteristics of the species, and the amount of habitat in the landscape. We conclude that the response of connectivity measures to habitat fragmentation should be understood before deriving conclusions for conservation management.     

The importance of functional connectivity in the conservation of a ground-dwelling mammal in an urban Australian landscape

The distribution of the northern brown bandicoot (Isoodon macrourus), a medium-sized ground-dwelling marsupial, was examined in habitat fragments within the urban landscape of the city of Brisbane, Australia. From surveys conducted in 68 fragments, bandicoots were found to be present in 33 (49%) despite widespread habitat loss and fragmentation. Logistic regression analysis revealed that of 13 measured independent variables, functional connectivity was the only factor that significantly predicted the presence of bandicoots within fragments, with connectivity positively correlated with the likelihood of occupation. Functional connectivity was equated to the likelihood of bandicoot immigration into the focal fragment from the nearest occupied fragment, based on the estimated resistance to movement offered by the intervening matrix. Within Brisbane, riparian habitat fragments typically have a relatively high level of functional connectivity, as thin strips of vegetation fringing waterways serve as corridors between larger riparian areas and facilitate the movement of bandicoots between patches. Analyses based on the Akaike Information Criterion revealed that the optimal model based on landscape context variables was convincingly better supported by the data than the optimal model produced from fragment characteristics. However, it is important to examine both internal attributes of habitat fragments and external features of the surrounding landscape when modelling the distribution of ground-dwelling fauna in urban environments, or other landscapes with a highly variable matrix. As urban centres throughout the world expand, it is crucial that the ecology of local wildlife be considered to ensure functional connection is maintained between habitat patches, especially for the conservation of species that are highly susceptible to fragmentation.     

Multi-scaled habitat considerations for conserving urban biodiversity: native reptiles and small mammals in Brisbane,Australia

The rapid expansion of the world’s urban population is a major driver of contemporary landscape change and ecosystem modification. Urbanisation destroys, degrades and fragments native ecosystems, replacing them with a heterogeneous matrix of urban development, parks, roads, and isolated remnant fragments of varying size and quality. This presents a major challenge for biodiversity conservation within urban areas. To make spatially explicit decisions about urban biodiversity conservation actions, urban planners and managers need to be able to separate the relative influence of landscape composition and configuration from patch and local (site)-scale variables for a range of fauna species. We address this problem using a hierarchical landscape approach for native, terrestrial reptiles and small mammals living in a fragmented semi-urban landscape of Brisbane, Australia. Generalised linear modelling and hierarchical partitioning analysis were applied to quantify the relative influence of landscape composition and configuration, patch size and shape, and local habitat composition and structure on the species’ richness of mammal and reptile assemblages. Landscape structure (composition and configuration) and local-scale habitat structure variables were found to be most important for influencing reptile and mammal assemblages, although the relative importance of specific variables differed between reptile and mammal assemblages. These findings highlight the importance of considering landscape composition and configuration in addition to local habitat elements when planning and/or managing for the conservation of native, terrestrial fauna diversity in urban landscapes.     

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.     

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.     

Spatial planning of a climate adaptation zone for wetland ecosystems

Here we present a spatial planning approach for the implementation of adaptation measures to climate change in conservation planning for ecological networks. We analyse the wetland ecosystems of the Dutch National Ecological Network for locations where the effectiveness of the network might be weakened because of climate change. We first identify potential dispersal bottlenecks where connectivity might be insufficient to facilitate range expansions. We then identify habitat patches that might have a too low carrying capacity for populations to cope with additional population fluctuations caused by weather extremes. Finally, we describe the spatial planning steps that were followed to determine the best locations for adaptation measures. An essential part of our adaptation strategy is to concentrate adaptation measures in a ‘climate adaptation zone’. Concentrating adaptation measures is a cost-effective planning strategy, rendering the largest benefit per area unit. Measures are taken where abiotic conditions are optimal and measures to enhance the spatial cohesion of the network are taken close to existing areas, thus creating the highest possible connectivity with the lowest area demands. Another benefit of a climate adaptation zone is that it provides a spatial protection zone where activities that will have a negative impact on ecosystem functioning might be avoided or mitigated. The following adaptation measures are proposed within the climate adaptation zone: (1) link habitat networks to enable species to disperse from present to future suitable climate zones, (2) enlarge the carrying capacity by either enlarging the size of natural areas or by improving habitat quality to shorten population recovery after disturbances, (3) increase the heterogeneity of natural areas, preferably by stimulating natural landscape-forming processes, to avoid large synchronised extinctions after extreme weather events. The presented approach can be generalised to develop climate adaptation zones for other ecosystem types inside or outside Europe, where habitat fragmentation is a limiting factor in biodiversity responses to climate change.     

Characterizing connectivity relationships in freshwaters using patch-based graphs

Spatial graphs in landscape ecology and conservation have emerged recently as a powerful methodology to model patterns in the topology and connectivity of habitat patches (structural connectivity) and the movement of genes, individuals or populations among these patches (potential functional connectivity). Most spatial graph’s applications to date have been in the terrestrial realm, whereas the use of spatially explicit graph-based methods in the freshwater sciences has lagged far behind. Although at first patch-based spatial graphs were not considered suitable for representing the branching network of riverine landscapes, here we argue that the application of graphs can be a useful tool for quantifying habitat connectivity of freshwater ecosystems. In this review we provide an overview of the potential of patch-based spatial graphs in freshwater ecology and conservation, and present a conceptual framework for the topological analysis of stream networks (i.e., riverscape graphs) from a hierarchical patch-based context. By highlighting the potential application of graph theory in freshwater sciences we hope to illustrate the generality of spatial network analyses in landscape ecology and conservation.