Assessing effects of land use on landscape connectivity: loss and fragmentation of western U.S. forests

Effects of land-use change on the conservation of biodiversity have become a concern to conservation scientists and land managers, who have identified loss and fragmentation of natural areas as a high-priority issue. Despite urgent calls to inform national, regional, and state planning efforts, there remains a critical need to develop practical approaches to identify where important lands are for landscape connectivity (i.e., linkages), where land use constrains connectivity, and which linkages are most important to maintain network-wide connectivity extents. Our overall goal in this paper was to develop an approach that provides comprehensive, quantitative estimates of the effects of land-use change on landscape connectivity and illustrate its use on a broad, regional expanse of the western United States. We quantified loss of habitat and landscape connectivity for western forested systems due to land uses associated with residential development, roads, and highway traffic. We examined how these land-use changes likely increase the resistance to movement of forest species in non-forested land cover types and, therefore, reduce the connectivity among forested habitat patches. To do so, we applied a graph-theoretic approach that incorporates ecological aspects within a geographic representation of a network. We found that roughly one-quarter of the forested lands in the western United States were integral to a network of forested patches, though the lands outside of patches remain critical for habitat and overall connectivity. Using remotely sensed land cover data (ca. 2000), we found 1.7 million km2 of forested lands. We estimate that land uses associated with residential development, roads, and highway traffic have caused roughly a 4.5% loss in area (20 000 km2) of these forested patches, and continued expansion of residential land will likely reduce forested patches by another 1.2% by 2030. We also identify linkages among forest patches that are critical for landscape connectivity. Our approach can be readily modified to examine connectivity for other habitats/ecological systems and for other geographic areas, as well as to address more specific requirements for particular conservation planning applications.     

Use of linkage mapping and centrality analysis across habitat gradients to conserve connectivity of gray wolf populations in western North America

Centrality metrics evaluate paths between all possible pairwise combinations of sites on a landscape to rank the contribution of each site to facilitating ecological flows across the network of sites. Computational advances now allow application of centrality metrics to landscapes represented as continuous gradients of habitat quality. This avoids the binary classification of landscapes into patch and matrix required by patch-based graph analyses of connectivity. It also avoids the focus on delineating paths between individual pairs of core areas characteristic of most corridor- or linkage-mapping methods of connectivity analysis. Conservation of regional habitat connectivity has the potential to facilitate recovery of the gray wolf (Canis lupus), a species currently recolonizing portions of its historic range in the western United States. We applied 3 contrasting linkage-mapping methods (shortest path, current flow, and minimum-cost-maximum-flow) to spatial data representing wolf habitat to analyze connectivity between wolf populations in central Idaho and Yellowstone National Park (Wyoming). We then applied 3 analogous betweenness centrality metrics to analyze connectivity of wolf habitat throughout the northwestern United States and southwestern Canada to determine where it might be possible to facilitate range expansion and interpopulation dispersal. We developed software to facilitate application of centrality metrics. Shortest-path betweenness centrality identified a minimal network of linkages analogous to those identified by least-cost-path corridor mapping. Current flow and minimum-cost-maximum-flow betweenness centrality identified diffuse networks that included alternative linkages, which will allow greater flexibility in planning. Minimum-cost-maximum-flow betweenness centrality, by integrating both land cost and habitat capacity, allows connectivity to be considered within planning processes that seek to maximize species protection at minimum cost. Centrality analysis is relevant to conservation and landscape genetics at a range of spatial extents, but it may be most broadly applicable within single- and multispecies planning efforts to conserve regional habitat connectivity.     

Consequences of ignoring dispersal variation in network models for landscape connectivity

Habitat loss and fragmentation can negatively influence population persistence and biodiversity, but the effects can be mitigated if species successfully disperse between isolated habitat patches. Network models are the primary tool for quantifying landscape connectivity, yet in practice, an overly simplistic view of species dispersal is applied. These models often ignore individual variation in dispersal ability under the assumption that all individuals move the same fixed distance with equal probability. We developed a modeling approach to address this problem. We incorporated dispersal kernels into network models to determine how individual variation in dispersal alters understanding of landscape-level connectivity and implemented our approach on a fragmented grassland landscape in Minnesota. Ignoring dispersal variation consistently overestimated a population's robustness to local extinctions and underestimated its robustness to local habitat loss. Furthermore, a simplified view of dispersal underestimated the amount of habitat substructure for small populations but overestimated habitat substructure for large populations. Our results demonstrate that considering biologically realistic dispersal alters understanding of landscape connectivity in ecological theory and conservation practice.     

A Graph-Theory Framework for Evaluating Landscape Connectivity and Conservation Planning

Abstract:  Connectivity of habitat patches is thought to be important for movement of genes, individuals, populations, and species over multiple temporal and spatial scales. We used graph theory to characterize multiple aspects of landscape connectivity in a habitat network in the North Carolina Piedmont (U.S.A).^. We compared this landscape with simulated networks with known topology, resistance to disturbance, and rate of movement. We introduced graph measures such as compartmentalization and clustering, which can be used to identify locations on the landscape that may be especially resilient to human development or areas that may be most suitable for conservation. Our analyses indicated that for songbirds the Piedmont habitat network was well connected. Furthermore, the habitat network had commonalities with planar networks, which exhibit slow movement, and scale-free networks, which are resistant to random disturbances. These results suggest that connectivity in the habitat network was high enough to prevent the negative consequences of isolation but not so high as to allow rapid spread of disease. Our graph-theory framework provided insight into regional and emergent global network properties in an intuitive and visual way and allowed us to make inferences about rates and paths of species movements and vulnerability to disturbance. This approach can be applied easily to assessing habitat connectivity in any fragmented or patchy landscape.     

Coupling urban 3-D information and circuit theory to advance the development of urban ecological networks

Ongoing, rapid urban growth accompanied by habitat fragmentation and loss challenges biodiversity conservation and leads to decreases in ecosystem services. Application of the concept of ecological networks in the preservation and restoration of connections among isolated patches of natural areas is a powerful conservation strategy. However, previous approaches often failed to objectively consider the impacts of complex 3-D city environments on ecological niches. We used airborne lidar-derived information on the 3-D structure of the built environment and vegetation and detailed land use and cover data to characterize habitat quality, niche diversity, and human disturbance and to predict habitat connectivity among 38 identified habitat core areas (HCAs) in Nanjing, China. We used circuit theory and Linkage Mapper to create a landscape resistance layer, simulate habitat connectivity, and identify and prioritize important corridors. We mapped 64 links by using current flow centrality to evaluate each HCA's contribution and the links that facilitate intact connectivity. Values were highest for HCA links located in the west, south, and northeast of the study area, where natural forests with complex 3-D structures predominate. Two smaller HCA areas had high centrality scores relative to their extents, which means they could act as important stepping stones in connectivity planning. The mapped pinch-point regions had narrow and fragile links among the HCAs, suggesting they require special protection. The barriers with the highest impact scores were mainly located at the HCA connections to Purple Mountain and, based on these high scores, are more likely to indicate important locations that can be restored to improve potential connections. Our novel framework allowed us to sufficiently convey spatially explicit information to identify targets for habitat restoration and potential pathways for species movement and dispersal. Such information is critical for assessing existing or potential habitats and corridors and developing strategic plans to balance habitat conservation and other land uses based on scientifically informed connectivity planning and implementation.     

Resource distributions among habitats determine solitary bee offspring production in a mosaic landscape.

Within mosaic landscapes, many organisms depend on attributes of the environment that operate over scales ranging from a single habitat patch to the entire landscape. One such attribute is resource distribution. Organisms' reliance on resources from within a local patch vs. those found among habitats throughout the landscape will depend on local habitat quality, patch quality, and landscape composition. The ability of individuals to move among complementary habitat types to obtain various resources may be a critical mechanism underlying the dynamics of animal populations and ultimately the level of biodiversity at different spatial scales. We examined the effects that local habitat type and landscape composition had on offspring production and survival of the solitary bee Osmia lignaria in an agri-natural landscape in California (U.S.A.). Female bees were placed on farms that did not use pesticides (organic farms), on farms that did use pesticides (conventional farms), or in seminatural riparian habitats. We identified pollens collected by bees nesting in different habitat types and matched these to pollens of flowering plants from throughout the landscape. These data enabled us to determine the importance of different plant species and habitat types in providing food for offspring, and how this importance changed with landscape and local nesting-site characteristics. We found that increasing isolation from natural habitat significantly decreased offspring production and survival for bees nesting at conventional farms, had weaker effects on bees in patches of seminatural habitat, and had little impact on those at organic farm sites. Pollen sampled from nests showed that females nesting in both farm and seminatural habitats relied on pollen from principally native plant species growing in seminatural habitat. Thus connectivity among habitats was critical for offspring production. Females nesting on organic farms were buffered to isolation effects by switching to floral resources growing at the farm site when seminatural areas were too distant. Overall local habitat conditions (farm management practices) can help bolster pollinators, but maintaining functional connectivity among habitats will likely be critical for persistence of pollinator populations as natural habitats are increasingly fragmented by human activities.     

Graph theory as a proxy for spatially explicit population models in conservation planning.

Spatially explicit population models (SEPMs) are often considered the best way to predict and manage species distributions in spatially heterogeneous landscapes. However, they are computationally intensive and require extensive knowledge of species' biology and behavior, limiting their application in many cases. An alternative to SEPMs is graph theory, which has minimal data requirements and efficient algorithms. Although only recently introduced to landscape ecology, graph theory is well suited to ecological applications concerned with connectivity or movement. This paper compares the performance of graph theory to a SEPM in selecting important habitat patches for Wood Thrush (Hylocichla mustelina) conservation. We use both models to identify habitat patches that act as population sources and persistent patches and also use graph theory to identify patches that act as stepping stones for dispersal. Correlations of patch rankings were very high between the two models. In addition, graph theory offers the ability to identify patches that are very important to habitat connectivity and thus long-term population persistence across the landscape. We show that graph theory makes very similar predictions in most cases and in other cases offers insight not available from the SEPM, and we conclude that graph theory is a suitable and possibly preferable alternative to SEPMs for species conservation in heterogeneous landscapes.     

Ranking individual habitat patches as connectivity providers: Integrating network analysis and patch removal experiments

Here we propose an integrated framework for modeling connectivity that can help ecologists, conservation planners and managers to identify patches that, more than others, contribute to uphold species dispersal and other ecological flows in a landscape context. We elaborate, extend and partly integrate recent network-based approaches for modeling and supporting the management of fragmented landscapes. In doing so, experimental patch removal techniques and network analytical approaches are merged into one integrated modeling framework for assessing the role of individual patches as connectivity providers. In particular, we focus the analyses on the habitat availability metrics PC and IIC and on the network metric Betweenness Centrality. The combination and extension of these metrics jointly assess both the immediate connectivity impacts of the loss of a particular patch and the resulting increased vulnerability of the network to subsequent disruptions. In using the framework to analyze the connectivity of two real landscapes in Madagascar and Catalonia (NE Spain), we suggest a procedure that can be used to rank individual habitat patches and show that the combined metrics reveal relevant and non-redundant information valuable to assert and quantify distinctive connectivity aspects of any given patch in the landscape. Hence, we argue that the proposed framework could facilitate more ecologically informed decision-making in managing fragmented landscapes. Finally, we discuss and highlight some of the advantages, limitations and key differences between the considered metrics.     

Multiple structural modifications to dendritic ecological networks produce simple responses

Structural modifications to landscapes affect the ability of organisms to access different habitat patches. There exist, however, very few general methods by which to relate modifications to expectations of effects, and even fewer that enable understanding of how multiple modifications may interact. In the absence of any guiding principles, ecologists have assumed that interactions will result in complex landscape-scale effects. One way of understanding such effects is through rendering a landscape as a graph or network, among the simplest of which are dendritic networks typified by stream systems. Yet even for stream networks, there are no known general principles concerning the nature of interactions between multiple modifications. We developed a model to describe the ability of fish to access and use different habitat patches within dendritic networks. We used mathematical and numerical analyses of the model to investigate how the habitat value of a network is affected by changes in connectivity and habitat quality, and then to examine interactions between multiple modifications. Rather than showing complex interactions, our analytic and simulation-based results show that the combined effect of multiple modifications approximately equals the sum of individually predicted effects. Dendritic networks thus appear to respond far more simply to multiple modifications than has previously been assumed. These results have implications for stream management planning, and offer a firm foundation from which to better understand population processes within dendritic networks.     

Validating graph-based connectivity models with independent presence–absence and genetic data sets

Habitat connectivity is a key objective of current conservation policies and is commonly modeled by landscape graphs (i.e., sets of habitat patches [nodes] connected by potential dispersal paths [links]). These graphs are often built based on expert opinion or species distribution models (SDMs) and therefore lack empirical validation from data more closely reflecting functional connectivity. Accordingly, we tested whether landscape graphs reflect how habitat connectivity influences gene flow, which is one of the main ecoevolutionary processes. To that purpose, we modeled the habitat network of a forest bird (plumbeous warbler [Setophaga plumbea]) on Guadeloupe with graphs based on expert opinion, Jacobs’ specialization indices, and an SDM. We used genetic data (712 birds from 27 populations) to compute local genetic indices and pairwise genetic distances. Finally, we assessed the relationships between genetic distances or indices and cost distances or connectivity metrics with maximum-likelihood population-effects distance models and Spearman correlations between metrics. Overall, the landscape graphs reliably reflected the influence of connectivity on population genetic structure; validation R² was up to 0.30 and correlation coefficients were up to 0.71. Yet, the relationship among graph ecological relevance, data requirements, and construction and analysis methods was not straightforward because the graph based on the most complex construction method (species distribution modeling) sometimes had less ecological relevance than the others. Cross-validation methods and sensitivity analyzes allowed us to make the advantages and limitations of each construction method spatially explicit. We confirmed the relevance of landscape graphs for conservation modeling but recommend a case-specific consideration of the cost-effectiveness of their construction methods. We hope the replication of independent validation approaches across species and landscapes will strengthen the ecological relevance of connectivity models.     

Chinook salmon use of spawning patches: relative roles of habitat quality, size, and connectivity.

Declines in many native fish populations have led to reassessments of management goals and shifted priorities from consumptive uses to species preservation. As management has shifted, relevant environmental characteristics have evolved from traditional metrics that described local habitat quality to characterizations of habitat size and connectivity. Despite the implications this shift has for how habitats may be prioritized for conservation, it has been rare to assess the relative importance of these habitat components. We used an information-theoretic approach to select the best models from sets of logistic regressions that linked habitat quality, size, and connectivity to the occurrence of chinook salmon (Oncorhynchus tshawytscha) nests. Spawning distributions were censused annually from 1995 to 2004, and data were complemented with field measurements that described habitat quality in 43 suitable spawning patches across a stream network that drained 1150 km2 in central Idaho. Results indicated that the most plausible models were dominated by measures of habitat size and connectivity, whereas habitat quality was of minor importance. Connectivity was the strongest predictor of nest occurrence, but connectivity interacted with habitat size, which became relatively more important when populations were reduced. Comparison of observed nest distributions to null model predictions confirmed that the habitat size association was driven by a biological mechanism when populations were small, but this association may have been an area-related sampling artifact at higher abundances. The implications for habitat management are that the size and connectivity of existing habitat networks should be maintained whenever possible. In situations where habitat restoration is occurring, expansion of existing areas or creation of new habitats in key areas that increase connectivity may be beneficial. Information about habitat size and connectivity also could be used to strategically prioritize areas for improvement of local habitat quality, with areas not meeting minimum thresholds being deemed inappropriate for pursuit of restoration activities.     

Connectivity in an Agricultural Landscape as Reflected by Interpond Movements of a Freshwater Turtle

Abstract:  Connectivity is a measure of how landscape features facilitate movement and thus is an important factor in species persistence in a fragmented landscape. The scarcity of empirical studies that directly quantify species movement and determine subsequent effects on population density have, however, limited the utility of connectivity measures in conservation planning. We undertook a 4-year study to calculate connectivity based on observed movement rates and movement probabilities for five age-sex classes of painted turtles ( Chrysemys picta ) inhabiting a pond complex in an agricultural landscape in northern Virginia (U.S.A.). We determined which variables influenced connectivity and the relationship between connectivity and subpopulation density. Interpatch distance and quality of habitat patches influenced connectivity but characteristics of the intervening matrix did not. Adult female turtles were more influenced by the habitat quality of recipient ponds than other age-sex classes. The importance of connectivity on spatial population dynamics was most apparent during a drought. Population density and connectivity were low for one pond in a wet year but dramatically increased as other ponds dried. Connectivity is an important component of species persistence in a heterogeneous landscape and is strongly dependent on the movement behavior of the species. Connectivity may reflect active selection or avoidance of particular habitat patches. The influence of habitat quality on connectivity has often been ignored, but our findings highlight its importance. Conservation planners seeking to incorporate connectivity measures into reserve design should not ignore behavior in favor of purely structural estimates of connectivity.     

An Experimental Test of Matrix Permeability and Corridor Use by an Endemic Understory Bird

Abstract:  Because of widespread habitat fragmentation, maintenance of landscape connectivity has become a major focus of conservation planning, but empirical tests of animal movement in fragmented landscapes remain scarce. We conducted a translocation experiment to test the relative permeability of three landscape elements (open habitat, shrubby secondary vegetation, and wooded corridors) to movement by the Chucao Tapaculo ( Scelorchilus rubecula ), a forest understory bird endemic to South American temperate rainforest. Forty-one radio-tagged subjects were translocated (individually) to three landscape treatments consisting of small release patches that were either entirely surrounded by open habitat (pasture), entirely surrounded by dense shrubs, or linked to other patches by wooded corridors that were otherwise surrounded by open matrix. The number of days subjects remained in release patches before dispersal (a measure of habitat resistance) was significantly longer for patches surrounded by open habitat than for patches adjoining corridors or surrounded by dense shrubs. These results indicate that open habitat significantly constrains Chucao dispersal, in accord with expectation, but dispersal occurs equally well through wooded corridors and shrub-dominated matrix. Thus, corridor protection or restoration and management of vegetation in the matrix (to encourage animal movement) may be equally feasible alternatives for maintaining connectivity.     

Conservation of Fragmented Populations

In this paper we argue that landscape spatial structure is of central importance in understanding the effects of fragmentation on population survival. Landscape spatial structure is the spatial relationships among habitat patches and the matrix in which they are embedded. Many general models of subdivided populations make the assumptions that (1) all habitat patches are equivalent in size and quality and (2) all local populations (in the patches) are equally accessible by dispersers. Models that gloss over spatial details of landscape structure can be useful for theoretical developments but will almost always be misleading when applied to real-world conservation problems. We show that local extinctions of fragmented populations are common. From this it follows that recolonization of local extinctions is critical for regional survival of fragmented populations. The probability of recolonization depends on (1) spatial relationships among landscape elements used by the population, including habitat patches for breeding and elements of the inter-patch matrix through which dispersers move, (2) dispersal characteristics of the organism of interest, and (3) temporal changes in the landscape structure. For endangered species, which are typically restricted in their dispersal range and in the kinds of habitat through which they can disperse, these factors are of primary importance and must be explicitly considered in management decisions.     

Use of Empirically Derived Source-Destination Models to Map Regional Conservation Corridors

Abstract:  The ability of populations to be connected across large landscapes via dispersal is critical to long-term viability for many species. One means to mitigate population isolation is the protection of movement corridors among habitat patches. Nevertheless, the utility of small, narrow, linear features as habitat corridors has been hotly debated. Here, we argue that analysis of movement across continuously resistant landscapes allows a shift to a broader consideration of how landscape patterns influence connectivity at scales relevant to conservation. We further argue that this change in scale and definition of the connectivity problem improves one's ability to find solutions and may help resolve long-standing disputes regarding scale and definition of movement corridors and their importance to population connectivity. We used a new method that combines empirically derived landscape-resistance maps and least-cost path analysis between multiple source and destination locations to assess habitat isolation and identify corridors and barriers to organism movement. Specifically, we used a genetically based landscape resistance model for American black bears ( Ursus americanus ) to identify major movement corridors and barriers to population connectivity between Yellowstone National Park and the Canadian border. Even though western Montana and northern Idaho contain abundant public lands and the largest wilderness areas in the contiguous United States, moving from the Canadian border to Yellowstone Park along those paths indicated by modeled gene flow required bears to cross at least 6 potential barriers. Our methods are generic and can be applied to virtually any species for which reliable maps of landscape resistance can be developed.     

Long-term drivers of persistence and colonization dynamics in spatially structured amphibian populations

Many organisms live in networks of local populations connected by dispersing individuals, called spatially structured populations (SSPs), where the long-term persistence of the entire network is determined by the balance between 2 processes acting at the scale of local populations: extinction and colonization. When multiple threats act on an SSP, a comparison of the different factors determining local extinctions and colonizations is essential to plan sound conservation actions. We assessed the drivers of long-term population dynamics of multiple amphibian species at the regional scale. We used dynamic occupancy models within a Bayesian framework to identify the factors determining persistence and colonization of local populations. Because connectivity among patches is fundamental to SSPs dynamics, we considered 2 measures of connectivity acting on each focal patch: incidence of the focal species and incidence of invasive crayfish. We used meta-analysis to summarize the effect of different drivers at the community level. Persistence and colonization of local populations were jointly determined by factors acting at different scales. Persistence probability was positively related to the area and the permanence of wetlands, whereas it was negatively related to occurrence of fish. Colonization probability was highest in semipermanent wetlands and in sites with a high incidence of the focal species in nearby sites, whereas it showed a negative relationship with the incidence of invasive crayfish in the landscape. By analyzing long-term data on amphibian population dynamics, we found a strong effect of some classic features commonly used in SSP studies, such as patch area and focal species incidence. The presence of an invasive non-native species at the landscape scale emerged as one of the strongest drivers of colonization dynamics, suggesting that studies on SSPs should consider different connectivity measures more frequently, such as the incidence of predators, especially when dealing with biological invasions.     

Connecting science,policy, and implementation for landscape‐scale habitat connectivity

We examined the links between the science and policy of habitat corridors to better understand how corridors can be implemented effectively. As a case study, we focused on a suite of landscape‐scale connectivity plans in tropical and subtropical Asia (Malaysia, Singapore, and Bhutan). The process of corridor designation may be more efficient if the scientific determination of optimal corridor locations and arrangement is synchronized in time with political buy‐in and establishment of policies to create corridors. Land tenure and the intactness of existing habitat in the region are also important to consider because optimal connectivity strategies may be very different if there are few, versus many, political jurisdictions (including commercial and traditional land tenures) and intact versus degraded habitat between patches. Novel financing mechanisms for corridors include bed taxes, payments for ecosystem services, and strategic forest certifications. Gaps in knowledge of effective corridor design include an understanding of how corridors, particularly those managed by local communities, can be protected from degradation and unsustainable hunting. There is a critical need for quantitative, data‐driven models that can be used to prioritize potential corridors or multicorridor networks based on their relative contributions to long‐term metacommunity persistence.     

A Resistant-Kernel Model of Connectivity for Amphibians that Breed in Vernal Pools

Abstract:  Pool-breeding amphibian populations operate at multiple scales, from the individual pool to surrounding upland habitat to clusters of pools. When metapopulation dynamics play a role in long-term viability, conservation efforts limited to the protection of individual pools or even pools with associated upland habitat may be ineffective over the long term if connectivity among pools is not maintained. Connectivity becomes especially important and difficult to assess in regions where suburban sprawl is rapidly increasing land development, road density, and traffic rates. We developed a model of connectivity among vernal pools for the four ambystomatid salamanders that occur in Massachusetts and applied it to the nearly 30,000 potential ephemeral wetlands across the state. The model was based on a modification of the kernel estimator (a density estimator commonly used in home range studies) that takes landscape resistance into account. The model was parameterized with empirical migration distances for spotted salamanders ( Ambystoma maculatum ), dispersal distances for marbled salamanders ( A. opacum ), and expert-derived estimates of landscape resistance. The model ranked vernal pools in Massachusetts by local, neighborhood, and regional connectivity and by an integrated measure of connectivity, both statewide and within ecoregions. The most functionally connected pool complexes occurred in southeastern and northeastern Massachusetts, areas with rapidly increasing suburban development. In a sensitivity analysis estimates of pool connectivity were relatively insensitive to uncertainty in parameter estimates, especially at the local and neighborhood scales. Our connectivity model could be used to prioritize conservation efforts for vernal-pool amphibian populations at broader scales than traditional pool-based approaches.     

Fencerows, Edges, and Implications of Changing Connectivity Illustrated by Two Contiguous Ohio Landscapes

We evaluated the amounts and implications of changes in habitat connectivity on rural landscapes by modeling the colonization success and subsequent habitat colonization of a model edge organism within real landscapes. We first inventoried the changes in the fencerow and forest-edge network of two contiguous Ohio (U.S.A.) landscapes, an agriculturally dominated till plain and a more diverse and dynamic moraine landscape, from 1940 to 1988. On the moraine the number of fencerows changed little from 1940 to 1971. The number increased by 86 during 1971–1988 as marginal farms were subdivided. The total length of fencerows on the moraine increased 2.5-fold over 48 years. On the till plain the number and total length of fencerows remained relatively constant through the study period. The sum of fencerows and forest edges was used as a measure of total ecotonal edge. On the moraine total edge increased through the study period, whereas on the till plain it decreased. We selected two levels of landscape connectivity, low and high, to model animal habitat colonization success. As connectivity increased the earliest successful colonists preempted an increasingly large proportion of the total suitable habitat, and the probability of successful colonization by later-arriving individuals decreased. The changes in connectivity that resulted from changes in both the fencerow network and the proportion of forested land have resulted in contiguous landscapes that present very different colonization potentials to organisms with long-distance dispersal capability. Given the current uncertainty of the effects of corridors on species-preservation efforts, we suggest that further modeling of this type prior to field testing will add useful insights, especially if conducted using specific species and landscape types.     

Spatial clustering of habitat structure effects patterns of community composition and diversity

Natural ecosystems often show highly productive habitats that are clustered in space. Environmental disturbances are also often nonrandomly distributed in space and are either intrinsically linked to habitat quality or independent in occurrence. Theoretical studies predict that configuration and aggregation of habitat patch quality and disturbances can affect metacommunity composition and diversity, but experimental evidence is largely lacking. In a metacommunity experiment, we tested the effects of spatially autocorrelated disturbance and spatial aggregation of patch quality on regional and local richness, among-community dissimilarity, and community composition. We found that spatial aggregation of patch quality generally increased among-community dissimilarity (based on two measures of beta diversity) of communities containing protozoa and rotifers in microcosms. There were significant interacting effects of landscape structure and location of disturbances on beta diversity, which depended in part on the specific beta diversity measures used. Effects of disturbance on composition and richness in aggregated landscapes were generally dependent on distance and connectivity among habitat patches of different types. Our results also show that effects of disturbances in single patches cannot directly be extrapolated to the landscape scale: the predictions may be correct when only species richness is considered, but important changes in beta diversity may be overlooked. There is a need for biodiversity and conservation studies to consider the spatial aggregation of habitat quality and disturbance, as well as connectivity among spatial aggregations.