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.     

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.     

Dispersal traits determine plant response to habitat connectivity in an urban landscape

Identification of trait syndromes that make species vulnerable to habitat fragmentation is essential in predicting biodiversity change. Plants are considered particularly vulnerable if their capacities for persistence in and for dispersal among local habitats are low. Here we investigated the influence of easily measured functional traits on the presence of 45 plant species in an urban landscape in north-west Germany where patches were separated by distances consistent with regular plant dispersal range. To describe the spatial configuration of patches we calculated species-specific patch connectivities. Then we assessed plant connectivity responses in distribution models calculated from connectivities and environmental predictors. Twenty (45%) of the analysed species showed a positive connectivity response after accounting for species-specific habitat requirements. These species differed from non-responsive species by functional traits associated with dispersal, including reduced seed numbers and higher terminal velocities relative to non-responsive species. Persistence traits played however no role which we attribute to the environmental conditions of urban habitats and their spatiotemporal characteristics. Our study underlines that even ruderal plants experience dispersal limitation and demonstrates that easily measured functional traits may be used as indicators of fragmentation vulnerability in urban systems allowing generalizations to larger species sets.     

Landscape connectivity and animal behavior: functional grain as a key determinant for dispersal

Landscape connectivity can be viewed from two perspectives that could be considered as extremes of a gradient: functional connectivity (refers to how the behavior of a dispersing organism is affected by landscape structure and elements) and structural connectivity (depends on the spatial configuration of habitat patches in the landscape like vicinity or presence of barriers). Here we argue that dispersal behavior changes with landscape configuration stressing the evolutionary dimension that has often been ignored in landscape ecology. Our working hypothesis is that the functional grain of resource patches in the landscape is a crucial factor shaping individual movements, and therefore influencing landscape connectivity. Such changes are likely to occur on the short-term (some generations). We review empirical studies comparing dispersal behavior in landscapes differing in their fragmentation level, i.e., with variable resource grain. We show that behavioral variation affecting each of the three stages of the dispersal process (emigration, displacement or transfer in the matrix, and immigration) is indeed likely to occur according to selective pressures resulting from changes in the grain of the landscape (mortality or deferred costs). Accordingly, landscape connectivity results from the interaction between the dispersal behavior of individuals and the grain of each particular landscape. The existence of this interaction requires that connectivity estimates (being based on individual-based models, least cost distance algorithms, and structural connectivity metrics or even Euclidian distance) should be carefully evaluated for their applicability with respect to the required level of precision in species-specific and landscape information.     

A multi-scale assessment of population connectivity in African lions (<Emphasis Type="Italic">Panthera leo)</Emphasis> in response to landscape change

Context

Habitat loss and fragmentation are among the major drivers of population declines and extinction, particularly in large carnivores. Connectivity models provide practical tools for assessing fragmentation effects and developing mitigation or conservation responses. To be useful to conservation practitioners, connectivity models need to incorporate multiple scales and include realistic scenarios based on potential changes to habitat and anthropogenic pressures. This will help to prioritize conservation efforts in a changing landscape.

Objectives

The goal of our paper was to evaluate differences in population connectivity for lions (Panthera leo) across the Kavango-Zambezi Trans-frontier Conservation Area (KAZA) under different landscape change scenarios and a range of dispersal distances.

Methods

We used an empirically optimized resistance surface, based on analysis of movement pathways of dispersing lions in southern Africa to calculate resistant kernel connectivity. We assessed changes in connectivity across nine landscape change scenarios, under each of which we explored the behavior of lions with eight different dispersal abilities.

Results

Our results demonstrate that reductions in the extent of the protected area network and/or fencing protected areas will result in large declines in the extent of population connectivity, across all modeled dispersal abilities. Creation of corridors or erection of fences strategically placed to funnel dispersers between protected areas increased overall connectivity of the population.

Conclusions

Our results strongly suggest that the most effective means of maintaining long-term population connectivity of lions in the KAZA region involves retaining the current protected area network, augmented with protected corridors or strategic fencing to direct dispersing individuals towards suitable habitat and away from potential conflict areas.

     

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.     

Modeling scale-dependent landscape pattern,dispersal, and connectivity from the perspective of the organism

Understanding the impacts of habitat fragmentation on dispersal is an important issue in landscape and conservation ecology. Here I examine the effects of fine- to broad-scale patterns in landscape structure on dispersal success of organisms with differing life-history traits. An individual-based model was used to simulate dispersal of amphibian-like species whose movements were driven by land cover and moisture conditions. To systematically control spatial pattern, a landscape model was created by merging simulated land cover maps with synthetic topographic surfaces. Landscapes varied in topographic roughness and spatial contagion in agriculture and urban land cover. Simulations included three different species types that varied in their maximum potential dispersal distances by 1-, 2-, or 4-fold. Two sets of simulations addressed effects of varying aspects of landscape structure on dispersal success. In the first set of simulations, which incorporated variable distances between breeding patches, dispersal success was lowest for all species types when anthropogenic cover was patchily distributed. In the second set, with interpatch distances held constant as landscape composition varied, dispersal success decreased as anthropogenic cover became spatially contagious. Both sets revealed strong main effects of species characteristics, interpatch distances and landscape composition on dispersal success; furthermore, scale-dependent patterns in land cover and moisture gradients had a stronger effect on longer- than shorter-ranging species types. Taken together, these simulations suggest that heuristic conservation strategies could potentially be developed based on important but limited life history information.     

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.     

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.

     

Dispersal success on fractal landscapes: a consequence of lacunarity thresholds

Habitat fragmentation is expected to disrupt dispersal, and thus we explored how patch metrics of landscape structure, such as percolation thresholds used to define landscape connectivity, corresponded with dispersal success on neutral landscapes. We simulated dispersal as either a purely random process (random direction and random step lengths) or as an area-limited random walk (random direction, but movement limited to an adjacent cell at each dispersal step) and quantified dispersal success for 1000 individuals on random and fractal landscape maps across a range of habitat abundance and fragmentation. Dispersal success increased with the number of cells a disperser could search (m), but poor dispersers (m<5) searching via area-limited dispersal on fractal landscapes were more successful at locating suitable habitat than random dispersers on either random or fractal landscapes. Dispersal success was enhanced on fractal landscapes relative to random ones because of the greater spatial contagion of habitat. Dispersal success decreased proportionate to habitat loss for poor dispersers (m=1) on random landscapes, but exhibited an abrupt threshold at low levels of habitat abundance (p<0.1) for area-limited dispersers (m<10) on fractal landscapes. Conventional metrics of patch structure, including percolation, did not exhibit threshold behavior in the region of the dispersal threshold. A lacunarity analysis of the gap structure of landscape patterns, however, revealed a strong threshold in the variability of gap sizes at low levels of habitat abundance (p<0.1) in fractal landscapes, the same region in which abrupt declines in dispersal success were observed. The interpatch distances or gaps across which dispersers must move in search of suitable habitat should influence dispersal success, and our results suggest that there is a critical gap-size structure to fractal landscapes that interferes with the ability of dispersers to locate suitable habitat when habitat is rare. We suggest that the gap structure of landscapes is a more important determinant of dispersal than patch structure, although both are ultimately required to predict the ecological consequences of habitat fragmentation.     

Evaluating functional connectivity with matrix behavior uncertainty for an endangered butterfly

Understanding animal responses to landscape elements helps forecast population reactions to changing landscape conditions. The challenge is that some behaviors are poorly known and difficult to estimate. We assessed how uncertainty in behavioral responses to dense woods, an avoided landscape structure, impacts functional connectivity among reproductive habitat patches for Fender’s blue butterfly, an endangered prairie species of western Oregon, USA. We designed a factorial simulation experiment using a spatially explicit individual-based model to project functional connectivity for female butterflies across current and alternative landscapes. We varied the probability of dense woods entry and turning angle standard deviation for movements within the dense woods over a range of biologically reasonable and observed values. Butterflies in the current landscape (46 % dense woods) and one with prairie encroached by forest (60 % dense woods) showed reductions in functional connectivity estimates consistent with the expectations of habitat fragmentation. Although dense woods entrance uncertainty impacted functional connectivity projections, uncertainty in the dense woods turning angle standard deviation had comparatively little impact on connectivity estimates. Reduction and reconfiguration of the current dense woods to 27 % cover (restored landscape) appeared to facilitate a corridor behavior in dispersing individuals, likely providing a functional connectivity estimate comparable to the historic landscape (<5 % dense woods). Our simulations suggest that additional study of butterfly movement within the dense woods is unnecessary and that a partial reduction in dense woods would be sufficient to achieve historic levels of functional connectivity for Fender’s blue across the study landscape.     

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.     

Movements of cactus bugs: Patch transfers,matrix resistance,and edge permeability

Individual movement is a key process affecting the distribution of animals in heterogeneous landscapes. For specialist species in patchy habitat, a central issue is how dispersal distances are related to landscape structure. We compared dispersal distances for cactus bugs (Chelinidea vittiger) on two naturally fragmented landscapes (≤ 4% suitable habitat) with different matrix structures (i.e., vegetation height of nonsuitable habitat between suitable patches). Using mark-release-recapture studies, we determined that most transfers between cactus patches occurred during the mating season. Dispersal distances were reduced by > 50% on the landscape that had reduced structural connectivity due to relatively high matrix structure and low patch density. An experiment with detailed movement pathways demonstrated that greater matrix structure decreased mean step lengths, reduced directionality, and thus decreased net displacement by > 60%. However, habitat edges between two matrix elements that differed substantially in resistance to movement were completely permeable. Therefore, the difference in distributions of dispersal distances between the two landscapes mainly reflected the average resistance of matrix habitat and not the level of matrix heterogeneity per se. Our study highlights the merits of combining estimates of dispersal distances with insights on mechanisms from detailed movement pathways, and emphasizes the difficulty of treating dispersal distances of species as fixed traits independent of landscape structure.     

Genetic isolation by distance and landscape connectivity in the American marten (Martes americana)

Empirical studies of landscape connectivity are limited by the difficulty of directly measuring animal movement. ‘Indirect’ approaches involving genetic analyses provide a complementary tool to ‘direct’ methods such as capture–recapture or radio-tracking. Here the effect of landscape on dispersal was investigated in a forest-dwelling species, the American marten (Martes americana) using the genetic model of isolation by distance (IBD). This model assumes isotropic dispersal in a homogeneous environment and is characterized by increasing genetic differentiation among individuals separated by increasing geographic distances. The effect of landscape features on this genetic pattern was used to test for a departure from spatially homogeneous dispersal. This study was conducted on two populations in homogeneous vs. heterogeneous habitat in a harvested boreal forest in Ontario (Canada). A pattern of IBD was evidenced in the homogeneous landscape whereas no such pattern was found in the near-by harvested forest. To test whether landscape structure may be accountable for this difference, we used effective distances that take into account the effect of landscape features on marten movement instead of Euclidean distances in the model of isolation by distance. Effective distances computed using least-cost modeling were better correlated to genetic distances in both landscapes, thereby showing that the interaction between landscape features and dispersal in Martes americana may be detected through individual-based analyses of spatial genetic structure. However, the simplifying assumptions of genetic models and the low proportions in genetic differentiation explained by these models may limit their utility in quantifying the effect of landscape structure.     

Can landscape indices predict ecological processes consistently?

The ecological interpretation of landscape patterns is one of the major objectives in landscape ecology. Both landscape patterns and ecological processes need to be quantified before statistical relationships between these variables can be examined. Landscape indices provide quantitative information about landscape pattern. Response variables or process rates quantify the outcome of ecological processes (e.g., dispersal success for landscape connectivity or Morisita's index for the spatial distribution of individuals). While the principal potential of this approach has been demonstrated in several studies, the robustness of the statistical relationships against variations in landscape structure or against variations of the ecological process itself has never been explicitly investigated. This paper investigates the consistency of correlations between a set of landscape indices (calculated with Fragstats) and three response variables from a simulated dispersal process across heterogeneous landscapes (cell immigration, dispersal success and search time) against variation in three experimental treatments (control variables): habitat amount, habitat fragmentation and dispersal behavior. I found strong correlations between some landscape indices and all three response variables. However, 68% of the statistical relationships were highly inconsistent and sometimes ambiguous for different landscape structures and for differences in dispersal behavior. Correlations between one landscape index and one response variable could range from highly positive to highly negative when derived from different spatial patterns. I furthermore compared correlation coefficients obtained from artificially generated (neutral) landscape models with those obtained from Landsat TM images. Both landscape representations produced equally strong and weak statistical relationships between landscape indices and response variables. This result supports the use of neutral landscape models in theoretical analyses of pattern-process relationships.     

Targeting and evaluating biodiversity conservation action within fragmented landscapes: an approach based on generic focal species and least-cost networks

The focus of biodiversity conservation is shifting to larger spatial scales in response to habitat fragmentation and the need to integrate multiple landscape objectives. Conservation strategies increasingly incorporate measures to combat fragmentation such as ecological networks. These are often based on assessment of landscape structure but such approaches fail to capitalise on the potential offered by more ecologically robust assessments of landscape function and connectivity. In this paper, we describe a modelling approach to identifying functional habitat networks and demonstrate its application to a fragmented landscape where policy initiatives seek to improve conditions for woodland biodiversity including increasing woodland cover. Functional habitat networks were defined by identifying suitable habitat and by modelling connectivity using least-cost approaches to account for matrix permeability. Generic focal species (GFS) profiles were developed, in consultation with stakeholders, to represent species with high and moderate sensitivity to fragmentation. We demonstrated how this form of analysis can be used to aid the spatial targeting of conservation actions. This ‘targeted’ action scenario was tested for effectiveness against comparable scenarios, which were based on random and clumped actions within the same landscape. We tested effectiveness using structural metrics, network-based metrics and a published functional connectivity indicator. Targeting actions within networks resulted in the highest mean woodland area and highest connectivity indicator value. Our approach provides an assessment of landscape function by recognising the importance of the landscape matrix. It provides a framework for the targeting and evaluation of alternative conservation options, offering a pragmatic, ecologically-robust solution to a current need in applied landscape ecology.     

Simulating the effects of climate change on population connectivity of American marten (Martes americana) in the northern Rocky Mountains, USA

We utilize empirically derived estimates of landscape resistance to assess current landscape connectivity of American marten (Martes americana) in the northern Rocky Mountains, USA, and project how a warming climate may affect landscape resistance and population connectivity in the future. We evaluate the influences of five potential future temperature scenarios involving different degrees of warming. We use resistant kernel dispersal models to assess population connectivity based on full occupancy of suitable habitat in each of these hypothetical future resistance layers. We use the CDPOP model to simulate gene exchange among individual martens in each of these hypothetical future climates. We evaluate: (1) changes in the extent, connectivity and pattern of marten habitat, (2) changes in allelic richness and expected heterozygosity, and (3) changes in the range of significant positive genetic correlation within the northern Idaho marten population under each future scenario. We found that even moderate warming scenarios resulted in very large reductions in population connectivity. Calculation of genetic correlograms for each scenario indicates that climate driven changes in landscape connectivity results in decreasing range of genetic correlation, indicating more isolated and smaller genetic neighborhoods. These, in turn, resulted in substantial loss of allelic richness and reductions in expected heterozygosity. In the U.S. northern Rocky Mountains, climate change may extensively fragment marten populations to a degree that strongly reduces genetic diversity. Our results demonstrate that for species, such as the American marten, whose population connectivity is highly tied to climatic gradients, expected climate change can result in profound changes in the extent, pattern, connectivity and gene flow of populations.     

Population dynamics and habitat connectivity affecting the spatial spread of populations – a simulation study

In this paper we show how the spatialconfiguration of habitat quality affects the spatial spread of apopulation in a heterogeneous environment. Our main result is thatfor species with limited dispersal ability and a landscape withisolated habitats, stepping stone patches of habitat greatlyincrease the ability of species to disperse. Our results showthat increasing reproductive rate first enables and thenaccelerates spatial spread, whereas increasing the connectivity has aremarkable effect only in case of low reproductive rates. Theimportance of landscape structure varied according to thedemographic characteristics of the population. To show this wepresent a spatially explicit habitat model taking into accountpopulation dynamics and habitat connectivity. The population dynamicsare based on a matrix projection model and are calculated on eachcell of a regular lattice. The parameters of the Leslie matrix dependon habitat suitability as well as density. Dispersal between adjacentcells takes place either unrestricted or with higher probability inthe direction of a higher habitat quality (restricted dispersal).Connectivity is maintained by corridors and stepping stones ofoptimal habitat quality in our fragmented model landscape containinga mosaic of different habitat suitabilities. The cellular automatonmodel serves as a basis for investigating different combinations ofparameter values and spatial arrangements of cells with high and lowquality.This revised version was published online in May 2005 with corrections to the Cover Date.     

Measuring landscape connectivity: On the importance of within-patch connectivity

Context

Many connectivity metrics have been used to measure the connectivity of a landscape and to evaluate the effects of land-use changes and potential mitigation measures. However, there are still gaps in our understanding of how to accurately quantify landscape connectivity.

Objectives

A number of metrics only measure between-patch connectivity, i.e. the connectivity between different habitat patches, which can produce misleading results. This paper demonstrates that the inclusion of within-patch connectivity is important for accurate results.

Methods

The behavior of two metrics is compared: the Connectance Index (CONNECT), which measures only between-patch connectivity, and the effective mesh size (m_eff), which includes both within-patch and between-patch connectivity. The connectivity values of both metrics were calculated on a set of simulated landscapes. Twenty cities were then added to these landscapes to calculate the resulting changes in connectivity.

Results

We found that when using CONNECT counter-intuitive results occurred due to not including within-patch connectivity, such as scenarios where connectivity increased with increasing habitat loss and fragmentation. These counter-intuitive results were resolved when using m_eff. For example, landscapes with low habitat amount may be particularly sensitive to urban development, but this is not reflected by CONNECT.

Conclusions

Applying misleading results from metrics like CONNECT can have detrimental effects on natural ecosystems, because reductions in within-patch connectivity by human activities are neglected. Therefore, this paper provides evidence for the crucial need to consider the balance between within-patch connectivity and between-patch connectivity when calculating the connectivity of landscapes.

     

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.