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.

     

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.     

Assessing landscape functional connectivity in a forest carnivore using path selection functions

Context

Understanding connectivity patterns in relation to habitat fragmentation is essential to landscape management. However, connectivity is often judged from expert opinion or species occurrence patterns, with very few studies considering the actual movements of individuals. Path selection functions provide a promising tool to infer functional connectivity from animal movement data, but its practical application remains scanty.

Objectives

We aimed to describe functional connectivity patterns in a forest carnivore using path-level analysis, and to explore how connectivity is affected by land cover patterns and road networks.

Methods

We radiotracked 22 common genets in a mixed forest-agricultural landscape of southern Portugal. We developed path selection functions discriminating between observed and random paths in relation to landscape variables. These functions were used together with land cover information to map conductance surfaces.

Results

Genets moved preferentially within forest patches and close to riparian habitats. Functional connectivity declined with increasing road density, but increased with the proximity of culverts, viaducts and bridges. Functional connectivity was favoured by large forest patches, and by the presence of riparian areas providing corridors within open agricultural land. Roads reduced connectivity by dissecting forest patches, but had less effect on riparian corridors due to the presence of crossing structures.

Conclusions

Genet movements were jointly affected by the spatial distribution of suitable habitats, and the presence of a road network dissecting such habitats and creating obstacles in areas otherwise permeable to animal movement. Overall, the study showed the value of path-level analysis to assess functional connectivity patterns in human-modified landscapes.

     

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.     

Characterizing the importance of habitat patches and corridors in maintaining the landscape connectivity of a Pholidoptera transsylvanica (Orthoptera) metapopulation

Since the fragmentation of natural habitats is one of the most serious problems for many endangered species, it is highly interesting to study the properties of fragmented landscapes. As a basic property, landscape connectivity and its effects on various ecological processes are frequently in focus. First, we discuss the relevance of some graph properties in quantifying connectivity. Then, we propose a method how to quantify the relative importance of habitat patches and corridors in maintaining landscape connectivity. Our combined index explicitly considers pure topological properties and topographical measures, like the quality of both patches (local population size) and corridors (permeability). Finally, for illustration, we analyze the landscape graph of the endangered, brachypterous bush-cricket Pholidoptera transsylvanica. The landscape contains 11 patches and 13 corridors and is situated on the Aggtelek Karst, NE-Hungary. We characterize the importance of each node and link of the graph by local and global network indices. We show how different measures of connectivity may suggest different conservation preferences. We conclude, accordingly to our present index, by identifying one specific habitat patch and one specific corridor being in the most critical positions in maintaining connectivity.This revised version was published online in May 2005 with corrections to the Cover Date.     

Movement behavior in response to landscape structure: the role of functional grain

Landscape structure can influence the fine-scale movement behavior of dispersing animals, which ultimately may influence ecological patterns and processes at broader scales. Functional grain refers to the finest scale at which an organism responds to spatial heterogeneity among patches and extends to the limits of its perceptual range. To determine the functional grain of a model insect, red flour beetle (Tribolium castaneum), we examined its movement behavior in response to experimental flour landscapes. Landscape structure was varied by manipulating habitat abundance (0%, 10%, 30%, and 100%) and grain size of patches (fine-2 × 2 cm, intermediate-5 × 5 cm, and coarse-10 × 10 cm) in 50 × 50 cm landscapes. Pathway metrics indicated that beetles used a similar proportion of all landscape types. Several pathway metrics indicated a graded response from the fine to the coarse grain landscape. Lacunarity analysis of beetle pathways indicated a non-linear change in space use between the fine and intermediate landscapes and the coarse-grained landscape. Beetles moved more slowly and tortuously (with many turns), and remained longer in both the overall landscape and individual patches, in fine-grained compared to coarse-grained landscapes. Our research demonstrates how detailed examination of movement pathways and measures of lacunarity can be useful in determining functional grain. Spatially explicit, organism-centered studies focusing on behavioral responses to different habitat configurations can serve as an important first step to identify behavioral rules of movement that may ultimately lead to more accurate predictions of space use in landscapes.     

Genetic analysis of landscape connectivity in tree populations

Genetic connectivity in plant populations is determined by gene movement within and among populations. When populations become genetically isolated, they are at risk of loss of genetic diversity that is critical to the long-term survival of populations. Anthropogenic landscape change and habitat fragmentation have become so pervasive that they may threaten the genetic connectivity of many plant species. The theoretical consequences of such changes are generally understood, but it is not immediately apparent how concerned we should be for real organisms, distributed across real landscapes. Our goals here are to describe how one can study gene movement of both pollen and seeds in the context of changing landscapes and to explain what we have learned so far. In the first part, we will cover methods of describing pollen movement and then review evidence for the impact of fragmentation in terms of both the level of pollen flow into populations and the genetic diversity of the resulting progeny. In the second part, we will describe methods for contemporary seed movement, and describe findings about gene flow and genetic diversity resulting from seed movement. Evidence for pollen flow suggests high connectivity, but it appears that seed dispersal into fragments may create genetic bottlenecks due to limited seed sources. Future work should address the interaction of pollen and seed flow and attention needs to be paid to both gene flow and the diversity of the incoming gene pool. Moreover, if future work is to model the impact of changing landscapes on propagule movement, with all of its ensuing consequences for genetic connectivity and demographic processes, we will need an effective integration of population genetics and landscape ecology.     

Static vs dynamic connectivity: how landscape changes affect connectivity predictions in the Iberian Peninsula

Landscape Ecology - Climate and land-use changes affect species ranges and movements. However, these changes are usually overlooked in connectivity studies, and this could have adverse consequences...     

Considering landscape connectivity and gene flow in the Anthropocene using complementary landscape genetics and habitat modelling approaches

Landscape Ecology - A comprehensive understanding of how rapidly changing environments affect species gene flow is critical for mitigating future biodiversity losses. While recent methodological...     

Limiting factors and landscape connectivity: the American marten in the Rocky Mountains

In mobile animals, movement behavior can maximize fitness by optimizing access to critical resources and minimizing risk of predation. We sought to evaluate several hypotheses regarding the effects of landscape structure on American marten foraging path selection in a landscape experiencing forest perforation by patchcut logging. We hypothesized that in the uncut pre-treatment landscape marten would choose foraging paths to maximize access to cover types that support the highest density of prey. In contrast, in the post-treatment landscapes we hypothesized marten would choose paths primarily to avoid crossing openings, and that this would limit their ability to optimally select paths to maximize foraging success. Our limiting factor analysis shows that different resistant models may be supported under changing landscape conditions due to threshold effects, even when a species’ response to landscape variables is constant. Our results support previous work showing forest harvest strongly affects marten movement behavior. The most important result of our study, however, is that the influence of these features changes dramatically depending on the degree to which timber harvest limits available movement paths. Marten choose foraging paths in uncut landscapes to maximize time spent in cover types providing the highest density of prey species. In contrast, following landscape perforation by patchcuts, marten strongly select paths to avoid crossing unforested areas. This strong response to patch cutting reduces their ability to optimize foraging paths to vegetation type. Marten likely avoid non-forested areas in fragmented landscapes to reduce risk of predation and to benefit thermoregulation in winter, but in doing so they may suffer a secondary cost of decreased foraging efficiency.     

Making the connection: combining habitat suitability and landscape connectivity to understand species distribution in an agricultural landscape

Landscape Ecology - The current biodiversity crisis has intensified the need to predict species responses to landscape modification and has renewed attention on the fundamental question of what...     

Linking movement behavior and fine-scale genetic structure to model landscape connectivity for bobcats (Lynx rufus)

Spatial heterogeneity can constrain the movement of individuals and consequently genes across a landscape, influencing demographic and genetic processes. In this study, we linked information on landscape composition, movement behavior, and genetic differentiation to gain a mechanistic understanding of how spatial heterogeneity may influence movement and gene flow of bobcats in the agricultural landscape of Iowa (USA). We analyzed movement paths of 23 animals to parameterize landscape resistance surfaces, applied least cost path analysis to generate measures of effective geographic distance between DNA collection locations of 625 bobcats, and tested the correlation between genetic distance and the different models of geographic distance. We found that bobcats showed a strong preference for forest over any other habitat type, and that incorporating information on habitat composition both along the path and in the surrounding landscape provided the best model of movement. Measures of effective geographic distance were significantly correlated with genetic distance, but not once the effects of Euclidean distance were accounted for. Thus, despite the impact of habitat composition on movement behavior, we did not detect a signature of a landscape effect in genetic structure. Our results are consistent with the issue of limiting factors: the high uniformity of forest fragmentation across southern Iowa, the primary study area, results in a landscape resistance pattern virtually indistinguishable from the isolation-by-distance pattern. The northern portion of the state, however, is predicted to pose a high level of resistance to bobcat movement, which may impede the regional genetic connectivity of populations across the Midwest.     

Sensitivity of resource selection and connectivity models to landscape definition

Context

The definition of the geospatial landscape is the underlying basis for species-habitat models, yet sensitivity of habitat use inference, predicted probability surfaces, and connectivity models to landscape definition has received little attention.

Objectives

We evaluated the sensitivity of resource selection and connectivity models to four landscape definition choices including (1) the type of geospatial layers used, (2) layer source, (3) thematic resolution, and (4) spatial grain.

Methods

We used GPS telemetry data from pumas (Puma concolor) in southern California to create multi-scale path selection function models (PathSFs) across landscapes with 2500 unique landscape definitions. To create the landscape definitions, we identified seven geospatial layers that have been shown to influence puma habitat use. We then varied the number, sources, spatial grain, and thematic resolutions of these layers to create our suite of plausible landscape definitions. We assessed how PathSF model performance (based on AIC) was affected by landscape definition and examined variability among the predicted probability of movement surfaces, connectivity models, and road crossing locations.

Results

We found model performance was extremely sensitive to landscape definition and identified only seven top models out of our suite of definitions (<1%). Spatial grain and the number of geospatial layers selected for a landscape definition significantly affected model performance measures, with finer grains and greater numbers of layers increasing model performance.

Conclusions

Given the sensitivity of habitat use inference, predicted probability surfaces, and connectivity models to landscape definition, out results indicate the need for increased attention to landscape definition in future studies.

     

Host functional connectivity and the spread potential of Lyme disease

Context

In the ecology of Lyme disease emergence, it remains unclear to what extent spread of the tick vector (Ixodes scapularis) and the pathogen (Borrelia burgdorferi) are dependent upon the dispersal of vertebrate hosts in spatially heterogeneous landscapes. Yet, empirical measure of these complex ecologically driven spread processes present conceptual and methodological challenges despite important public health implications.

Objectives

To examine the relationship between landscape characteristics and tick-borne disease spread, we modeled the influence of landscape connectivity for a simplified vertebrate host community (white-footed mouse—Peromyscus leucopus, American robin—Turdus migratorius, white-tailed deer—Odocoileus virginianus) on the potential spread of the tick population compared to the pathogen in a spatially-structured landscape.

Methods

We parameterized a hybrid demographic-dispersal connectivity model by combining a series of reported host dispersal and tick burden estimates with empirically-measured tick abundance and pathogen prevalence sampled from a Lyme-endemic island landscape in Thousand Islands National Park (Ontario, Canada) and simulated several tick- and pathogen-spread scenarios.

Results

The extent of tick spread by mice [amount of reachable habitat (ARH)?=?18.0%] is considerably similar to that of robins (ARH?=?18.7%), while deer support the greatest tick spread extent (ARH?=?82.0%). Infected mice carrying ticks support the highest pathogen spread (ARH?=?19.8%). Short-distance pathogen spread and long-distance tick spread were facilitated by intermediate stepping stone habitat fragments.

Conclusions

We provide evidence that host functional connectivity mediates tick spread differently than pathogen spread, and depends strongly on landscape configuration. Our study therefore emphasizes the importance of landscape spatial heterogeneity on the ecological processes that influence regional tick-borne disease spread.

     

Estimation of metademographic rates and landscape connectivity for a conservation-reliant anuran

Landscape Ecology - Amphibian conservation efforts commonly assume populations are tied to waterbodies that collectively function as a metapopulation. This assumption is rarely evaluated, and there...     

Temporal shifts in landscape connectivity for an ecosystem engineer,the roe deer,across a multiple-use landscape

Context

Routine movements of large herbivores, often considered as ecosystem engineers, impact key ecological processes. Functional landscape connectivity for such species influences the spatial distribution of associated ecological services and disservices.

Objectives

We studied how spatio-temporal variation in the risk-resource trade-off, generated by fluctuations in human activities and environmental conditions, influences the routine movements of roe deer across a heterogeneous landscape, generating shifts in functional connectivity at daily and seasonal time scales.

Methods

We used GPS locations of 172 adult roe deer and step selection functions to infer landscape connectivity. In particular, we assessed the influence of six habitat features on fine scale movements across four biological seasons and three daily periods, based on variations in the risk-resource trade-off.

Results

The influence of habitat features on roe deer movements was strongly dependent on proximity to refuge habitat, i.e. woodlands. Roe deer confined their movements to safe habitats during daytime and during the hunting season, when human activity is high. However, they exploited exposed open habitats more freely during night-time. Consequently, we observed marked temporal shifts in landscape connectivity, which was highest at night in summer and lowest during daytime in autumn. In particular, the onset of the autumn hunting season induced an abrupt decrease in landscape connectivity.

Conclusions

Human disturbance had a strong impact on roe deer movements, generating pronounced spatio-temporal variation in landscape connectivity. However, high connectivity at night across all seasons implies that Europe’s most abundant and widespread large herbivore potentially plays a key role in transporting ticks, seeds and nutrients among habitats.

     

On the relative importance of landscape variables to plant diversity and phylogenetic community structure on uninhabited islands,South Korea

Landscape Ecology - Although there have been methodological advances in the analysis of biodiversity and landscape properties, most studies in landscape ecology primarily explore whether certain...     

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.     

Effects of landscape composition and connectivity on the distribution of an endangered parrot in agricultural landscapes

The extent and connectivity of individual habitat types strongly affects the distribution and abundance of organisms. However, little is known of how the level of connectivity and the interactions between different habitat types influences the distribution of species. Here, we used the geographically restricted and endangered regent parrot Polytelis anthopeplus monarchoides as a case study to examine the importance of composition and connectivity between different elements in 39 complex landscape mosaics (each 10 km radius). We compiled a database of 674 regent parrot nesting records, regional vegetation maps and measures of multipath connectivity between core vegetation types under different scenarios of resistance to movement provided by landscape elements. The occurrence of regent parrot nests was strongly affected by landscape composition, being positively related to the extent Eucalyptus camaldulensis riverine forest, but negatively related to the extent of semi-arid woodlands dominated by Eucalyptus largiflorens. Connectivity between E. camaldulensis forest (principal nesting habitat) and mallee (preferred feeding habitat) was a strong predictor of nest locations. Our study shows that the suitability of fragmented agricultural landscapes for supporting species can be greatly affected by connectivity and interactions between preferred and non-preferred habitats. For species that require complementary habitats such as the regent parrot, conservation management activities may be ineffective if they simply focus on a single core habitat type or the impacts of human land uses without regard to the interrelationships among landscape elements. While increasing the amount of primary preferred habitat should remain a cornerstone goal, increasing the extent and improving connectivity with alternative landscape elements also should be priority management objectives.