Behavior of class-level landscape metrics across gradients of class aggregation and area

Habitat loss and fragmentation processes strongly affect biodiversity conservation in landscapes undergoing anthropogenic land use changes. Many attempts have been made to use landscape structure metrics to quantify the independent and joint effects of these processes. Unfortunately, ecological interpretation of those metrics has been plagued by lack of thorough understanding of their theoretical behavior. We explored behavior of 50 metrics in neutral landscapes across a 21-step gradient in aggregation and a 19-step gradient in area using a full factorial design with 100 replicates of each of the 399 combinations of the two factors to assess how well metrics reflected changes in landscape structure. Metric values from real landscapes were used to determine the extent of neutral landscape space that is represented in real landscapes. We grouped metrics into three major behavioral classes: strongly related to focal class area (n=15), strongly related to aggregation (n=7), and jointly responding to area and aggregation (n=28). Metrics strongly related to class area exhibited a variety of distinct behaviors, and many of these metrics have unique interpretations that make each of them particularly useful in certain applications. Metrics strongly related to aggregation, independent of class area, are particularly useful in assessing effects of fragmentation. Moreover, metrics in this group exhibited a range of specific behaviors, highlighting subtle but different aspects of landscape aggregation even though we controlled only one aspect of aggregation. The non-linear behavior exhibited by many metrics renders interpretation difficult and use of linear analytical techniques inappropriate under many circumstances. Ultimately, comprehensive characterization of landscapes undergoing habitat loss and fragmentation will require using several metrics distributed across behavioral groups.This revised version was published online in May 2005 with corrections to the Cover Date.     

Species Dynamics in Disturbed Landscapes: When does a Shifting Habitat Mosaic Enhance Connectivity?

Although landscape ecology emphasizes the effects of spatial pattern on ecological processes, most neutral models of species–habitat relationships have treated habitat as a static constraint. Do the working hypotheses derived from these models extend to real landscapes where disturbances create a shifting mosaic? A spatial landscape simulator incorporating vegetation dynamics and a metapopulation model was used to compare species in static and dynamic landscapes with identical habitat amounts and spatial patterns. The main drivers of vegetation dynamics were stand-replacing disturbances, followed by gradual change from early-successional to old-growth habitats. Species dynamics were based on a simple occupancy model, with dispersal simulated as a random walk. As the proportion of available habitat (p) decreased from 1.0, species occupancy generally declined more rapidly and reached extinction at higher habitat levels in dynamic than in static landscapes. However, habitat occupancy was sometimes actually higher in dynamic landscapes than in static landscapes with similar habitat amounts and patterns. This effect was most pronounced at intermediate amounts of habitat (p = 0.3?0.6) for mobile species that had high colonization rates, but were unable to cross non-habitat patches. Differences between static and dynamic landscapes were contingent upon the initial metapopulation size and the shapes of disturbances and the resulting habitat patterns. Overall, the results demonstrate that dispersal-limited species exhibit more pronounced critical behavior in dynamic landscapes than is predicted by simple neutral models based on static landscapes. Thus, caution should be exercised in extending generalizations derived from static landscape models to disturbance-driven landscape mosaics.     

Effects of area and isolation of woodland patches on herbaceous plant species richness across Great Britain

Richness of Ancient Woodland Indicator plant species was analysed in 308 woodland patches that were surveyed during the Countryside Survey of Great Britain carried out in 1998. The Countryside Survey recorded vegetation plots and landscape structure in 569 stratified 1 km sample squares and developed a remotely-sensed land cover map of the UK. Using these datasets, we tested the hypothesis that Ancient Woodland Indicator species richness in woodland fragments was limited by patch area, shape and spatial isolation and that woodland patches located in the lowland region of Great Britain would respond differently than those in the upland region. The variation in Ancient Woodland Indicator species richness in the British lowlands (n = 218) was mainly explained by patch area and two measures of connectivity, the length of hedgerows and lines of trees in the 1 km square and the area of woodland within 500 m of the vegetation plot. By contrast, variation in Ancient Woodland Indicator species richness in the British uplands (n = 90) was related to Ellenberg scores of the vegetation communities sampled – a surrogate for habitat quality – and no significant effect of spatial structure was detected. It therefore appears that the degree of fragmentation of woodland in the British lowlands limits the distribution of Ancient Woodland Indicator species, while in the uplands, failed colonisation is a matter of habitat quality rather than a result of landscape structure.This revised version was published online in May 2005 with corrections to the Cover Date.     

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

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

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.     

A new measure of longitudinal connectivity for stream networks

Habitat connectivity is a central factor in shaping aquatic biological communities, but few tools exist to describe and quantify this attribute at a network scale in riverine systems. Here, we develop a new index to quantify longitudinal connectivity of river networks based on the expected probability of an organism being able to move freely between two random points of the network. We apply this index to two fish life histories and evaluate the effects of the number, passability, and placement of barriers on river network connectivity through the use of simulated dendritic ecological networks. We then extend the index to a real world dendritic river system in Newfoundland, Canada. Our results indicate that connectivity in river systems, as represented by our index, is most impacted by the first few barriers added to the system. This is in contrast to terrestrial systems, which are more resilient to low levels of connectivity. The results show a curvilinear relationship between barrier passability and structural connectivity. This suggests that an incremental improvement in passability would result in a greater improvement to river network connectivity for more permeable barriers than for less permeable barriers. Our analysis of the index in simulated and real river networks also showed that barrier placement played an important role in connectivity. Not surprisingly, barriers located near the river mouth have the greatest impact on fish with diadromous life histories while those located near the center of the river network have the most impact on fish with potadromous life histories. The proposed index is conceptually simple and sufficiently flexible to deal with variations in river structure and biological communities. The index will enable researchers to account for connectivity in habitat studies and will also allow resource managers to characterize watersheds, assess cumulative impacts of multiple barriers and determine priorities for restoration.     

Neutral model analysis of landscape patterns from mathematical morphology

Mathematical morphology encompasses methods for characterizing land-cover patterns in ecological research and biodiversity assessments. This paper reports a neutral model analysis of patterns in the absence of a structuring ecological process, to help set standards for comparing and interpreting patterns identified by mathematical morphology on real land-cover maps. We considered six structural classes (core, perforated, edge, connector, branch, and patch) on randomly generated binary (forest, non-forest) maps in which the percent occupancy (P) of forest varied from 1% to 99%. The maps were dominated by the patch class for low P, by the branch and connector classes for intermediate P, and by the edge, perforated, and core classes for high P. Two types of pattern phase changes were signaled by abrupt transitions among the six structural classes, at critical P thresholds that were indicated by increased variance among maps for the same P. A phase change from maps dominated by the patch class to maps dominated by the branch and connector classes was related to the existence of a percolating cluster of forest, and the P threshold varied depending on the co-existence of the core class. A second phase change from the edge class to the perforated class was related to the existence of a percolating cluster of non-core (including non-forest) and represents a change of context from exterior to interior. Our results appear to be the first demonstration of multiple phase changes controlling different aspects of landscape pattern on random neutral maps. Potential applications of the results are illustrated by an analysis of ten real forest maps. The U.S. Government's right to retain a non-exclusive, royalty-free license in and to any copyright is acknowledged.     

Regional assessment on influence of landscape configuration and connectivity on range size of white-tailed deer

Variation in the size of home range of white-tailed deer (Odocoileus virginianus) has broad implications for managing populations, agricultural damage, and disease spread and transmission. Size of home range of deer also varies seasonally because plant phenology dictates the vegetation types that are used as foraging or resting sites. Knowledge of the landscape configuration and connectivity that contributes to variation in size of home range of deer for the region is needed to fully understand differences and similarities of deer ecology throughout the Midwest. We developed a research team from four Midwestern states to investigate how size of home range of deer in agro-forested landscapes is influenced by variations in landscape characteristics that provide essential habitat components. We found that for resident female deer, annual size of home range in Illinois (mean = 0.99 km²), Michigan (mean = 1.34 km²), Nebraska (mean = 1.20 km²), and Wisconsin (mean = 1.47 km²) did not differ across the region (F _3,175 = 0.42, P = 0.737), but differences between agricultural growing and nongrowing periods were apparent. Variables influencing size of home range included: distance to forests, roads, and urban development from the centroid of deer home range, and percent of crop as well as four landscape pattern indices (contrast-weighted edge density, mean nearest neighbor, area-weighted mean shape index, and patch size coefficient of variation). We also identified differences in model selection for four landscapes created hierarchically to reflect levels of landscape connectivity determined from perceived ability of deer to traverse the landscape. Connectivity of selected forested regions within agro-forested ecosystems across the Midwest plays a greater role in understanding the size of home ranges than traditional definitions of deer habitat conditions and landscape configuration.     

Role of green roofs in urban connectivity,an exploratory approach using landscape graphs in the city of Paris,France

Green roofs provide many ecosystem services, but little is known about the way they contribute to urban functional connectivity. This paper has the following four objectives: (1) to compare the potential green roofs’ role to connectivity in relation to other urban green spaces, (2) to specify the green roofs contribution’s type, (3) to explore the influence of building height integration method and finally (4) to assess the impact on connectivity of simulated greening new roofs. Using a landscape graph approach, we modeled ecological networks of three species groups with different dispersion capacities in the Paris region (France). Then, we computed several connectivity metrics to assess the potential contribution of green roofs to functional connectivity. At a large scale (metropole scale), our results show that green roofs can slightly improve the global connectivity largely through the connections rather than the addition of habitat area. More than a stepping stone function, green roofs would have a dispersion flux function at a local scale. Furthermore, when the difficulty of crossing movement is exponential to the height of buildings, green roofs over 20 m high are mostly disconnected from the ecological networks. In addition to the green roof’s height, our analysis highlights the very strong role played by buildings’ configuration. This study raises promising directions for the integration of building height into the analysis of urban connectivity. Detailed research and long-term biological data from green roofs and green spaces are needed to confirm our results.     

Effectiveness of corridor vegetation depends on urbanization tolerance of forest birds in central Tokyo,Japan

Vegetation corridors, such as street trees in urban areas, which connect patchy woodland and mitigate habitat isolation, are expected to enhance the persistence of birds in urban landscapes. However, the effectiveness of urban corridors on birds remains equivocal because vegetation corridor is often managed for human use with little consideration of wildlife. Here we compared the effects of three major corridors of varying vegetation structures (trees with a dense understory, trees with a sparse understory, and grassy areas with sparse trees) on the species richness and abundance of birds in 21 wooded patches in the center of Tokyo, Japan, during wintering and breeding seasons. Using generalized linear models and Akaike’s information criterion, we found that the effectiveness of corridors depended on the tolerance of birds to urbanization. Urban avoider species, having low tolerance to urbanization, demonstrated lower species richness and abundance in patches close to the corridor with a sparsely vegetated understory as compared with patches close to the understory-richer corridors during winter, although such an effect disappeared during the breeding season. The corridors did not have a significant effect on suburban adapter species with a high tolerance to urbanization. Our results suggest that corridors with scarce understory vegetation may limit the persistence of birds avoiding urban areas.     

Modeling and field-testing of Ovenbird (<Emphasis Type="Italic">Seiurus aurocapillus</Emphasis>) responses to boreal forest dissection by energy sector development at multiple spatial scales

Although the area disturbed by linear features in forested systems is small relative to many other human disturbances, linear features create significantly more amounts of edge per unit area. In the boreal plains of Alberta, Canada, energy sector exploration has resulted in extensive dissection of the landscape through 8 m wide seismic lines. A spatially explicit model was developed to test how bird abundance might change in response to increasing seismic line density if individuals use seismic lines as territory boundaries or actively avoid these edges. Assuming birds had fixed territory shape and size, increasing seismic line density from 0 to 8 km/km² resulted in a 38% decline and an 82% decline in bird abundance when individuals used lines as territory boundaries or avoided edges by 50 m, respectively. We tested the assumptions of our model using the Ovenbird (Seiurus aurocapillus). Based on radio-telemetry (n = 12), all Ovenbirds crossed seismic lines at some point during the breeding season. However, male Ovenbirds showed a distinct use of one side of the seismic line, suggesting lines acted as territory boundaries. In 12.25 ha plots (n = 24) spot-mapping detected no change in Ovenbird density as linear feature density increased from 0 to 8.6 km/km². In 4 km² landscapes (n = 62) sampled using a grid of nine point-counts, we also detected no changes in Ovenbird numbers across the same range of seismic line densities. Ovenbirds declined with seismic line density at the level of the individual point-count station (12 ha scale), but only when a threshold seismic line density of 8.5 km/km² was reached. Above the threshold, Ovenbirds declined 19% for each 1 km/km² increase in seismic line density. While relatively few places in Alberta’s boreal forest have local seismic line densities of 8.5 km/km², forest dissection could increasingly become an issue if current energy exploration practices continue.     

Carabids (Coleoptera: Carabidae) in a forest patchwork: a connectivity analysis of the Bereg Plain landscape graph

For many species, one important key to persistence is maintaining connectivity among local populations that allow for dispersal and gene flow. This is probably true for carabid species (Coleoptera:Carabidae) living in the fragmented forests of the Bereg Plain (NE Hungary and W Ukraine). Based on field data, we have drafted a landscape graph of the area representing the habitat network of these species. Graph nodes and links represented two kinds of landscape elements: habitat (forest) patches and corridors, respectively. The quality of habitat patches and corridors were ranked (from low (1) to high (4)), reflecting local population sizes in the case of patches and estimated permeability in the case of corridors. We analysed (1) the positional importance of landscape elements in maintaining the connectivity of the intact network, (2) the effect of inserting hypothetical corridors into the network, (3) the effects of improving the quality of the existing corridors, and (4) how to connect every patch in a cost-effective way. Our results set quantitative priorities for conservation practice by identifying important corridors: what to protect, what to build and what to improve. Several network analytical techniques were used to account for the directed (source-sink) and highly fragmented nature of the landscape graph. We provide conservation priority ranks for the landscape elements and discuss the conditions for the use of particular network indices. Our study could be of extreme relevance, since a new highway is being planned through the area.     

The potential of school green areas to improve urban green connectivity and multifunctionality

Urban green infrastructure supports resilience in cities and promotes sustainable resource management. Small green areas, including school green areas (SGAs), are an important component of urban green infrastructure, playing a key role in supplying cities with educational services. This article describes how SGAs can amplify an urban green area's connectivity and multifunctionality. The analysis was performed in Bucharest as a case study. A survey based on questionnaires was used to obtain data regarding green spaces within public schools. A total of 411 administrators from 461 public schools participated in the survey for a response rate of 89.1%. Information from the questionnaires was augmented with spatial data of SGAs and public green spaces, i.e., parks and city gardens. Using parametric and nonparametric statistical analysis, we first identified the variables that determine an SGA's presence and size. Potential connectivity assessment results showed that most of the schools that lack or have small-sized SGAs have the possibility to cover their green space deficit by developing activities within nearby public green spaces. A structural connectivity assessment of SGAs toward other public urban green areas revealed that SGAs are an important element of the urban environment by serving as stepping stones to species flow. The multifunctionality of the SGAs was emphasized through the educational services they provide, being involved in pupils’ daily activities. The increased connectivity and multifunctionality of urban green infrastructure through small, specialized green areas, such as SGAs, is an indicator of the fact that such areas can be used to ameliorate the deficit of green space in major urban areas.     

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.     

Corridors and connectivity: when use and function do not equate

Connectivity, or the integration of populations into a single demographic unit, is an often desired, but largely untested aspect of wildlife corridors. Using a corridor system that was established at least 85 years prior, we investigated the extent of connectivity provided. This was undertaken using a combined ecological and genetic approach with connectivity estimated by gene flow. Vegetation within the corridor was found to be comparable in physical structure and species composition to that within the connected patches and the two target species (Melomys cervinipes and Uromys caudimaculatus) were shown to occur along the corridor but not within the surrounding matrix. These factors indicated that the corridor was suitable for use as a model system. The population structure (weights of individuals, sex ratios and the percentage of juveniles) of both species were also similar within the corridor and the connected patches suggesting that the corridor provided the resources necessary to sustain breeding populations along its length. Despite this, populations in patches linked by the corridor were found to show the same significant levels of genetic differentiation as those in isolated habitats. M. cervinipes, but not U. caudimaculatus, also showed population differentiation within the continuous habitat. Although based on only one corridor system, these results clearly demonstrate that connectivity between connected populations will not always be achieved by the construction or retention of a corridor and that connectivity cannot be inferred solely from the presence of individuals, or breeding populations, within the corridor. C. Wilson: deceased     

The influences of patch shape and boundary contrast on insect response to fragmentation in California grasslands

Landscape ecologists typically identify boundaries to demarcate habitatpatches. The boundary between two habitat types may be abrupt, such as thetransition between a grassland and a parking lot, or more gradual, such as theshift between successional forest stages. Two key aspects of landscapeboundaries, their shape and contrast, are predicted to influence movement ofmaterials, plants, and animals. Ecological theory suggests that a patchsperimeter-to-area ratio should strongly influence animal emigration when patchboundaries are relatively permeable, but not when boundaries are more severe.Weinvestigated the interactive effects of patch shape and boundary contrast onmovement of ground-dwelling beetles (Carabidae and Tenebrionidae) in nativegrassland habitat at Jepson Prairie, Solano County, California, USA. Weconducted a field experiment with two patch shape treatments, square andrectangle, that held patch area constant, and two boundary contrast treatmentscreated by mowing grass surrounding each plot at two different heights. Wemonitored the number of beetles leaving each patch over a three-week periodfollowing treatment establishment. We observed a significant effect of boundarycontrast on net movement of beetles, with low contrast boundaries exhibitingnetimmigration and high contrast boundaries experiencing net emigration. Moreover,the importance of patch shape appeared to be greater for low contrast versushigh contrast boundaries, consistent with theoretical expectations. Ourcombinedobservations indicate that these ground-dwelling beetles were more likely tomove into patches that were rectangular and surrounded by a low contrast matrixthan patches that were square or surrounded by a high contrast matrix. Weconclude that net movement of beetles across patch boundaries is stronglyinfluenced by boundary contrast and may be affected by patch shape whenboundarycontrast is low.This revised version was published online in May 2005 with corrections to the Cover Date.     

Landscape factors and regional differences in recovery rates of herb layer richness in Flanders (Belgium)

The recovery of understory plants in recent forests is critical for evaluating the overall capacity of landscapes to maintain plant biodiversity. Here we used a large data set of vegetation plots from the Flemish Forest Inventory in combination with maps of forest history and soil-based Potential Natural Vegetation to evaluate regional differences in the rate of recovery of understory plant diversity in three regions of Flanders, Belgium. We expressed the degree of recovery in recent forests using the species richness of ancient forests as a reference point, and found strong differences among regions in the average level of recovery. These differences appeared to be due to regional variation in average patch connectivity and age (ultimately stemming from differences in land use history) and – to a lesser extent – environmental conditions. We also found an increase in the proportional representation of vertebrate dispersed species and species with short-distance dispersal with increasing levels of recovery. Our results highlight the potential drivers of inter-regional variation in the process of recovery of plant diversity during restoration, and they emphasize the importance of historical and spatial context in the recovery process.     

Graph theory in action: evaluating planned highway tracks based on connectivity measures

Maintaining connectivity among local populations in a fragmented landscape is crucial for the survival of many species. For isolated habitat patches, stochastic fluctuations and reduced gene flow can lead to high risk of extinction. The connectivity of the landscape is especially crucial for the carabid species living in the fragmented forests of the Bereg plain (NE Hungary and W Ukraine) because a highway will be constructed through the plain. Our purpose is to (1) evaluate the impacts of three possible highway tracks, (2) suggest a solution that is realistic with less impact on connectivity than other plans and (3) discuss how to decrease the disadvantageous effects of each track. Our results, based on a network analysis of landscape graph of patches and ecological corridors, indicate that the intended highway could have deleterious consequences on forest-living carabids. Relatively simple actions, like the establishment of stepping stones, could compensate for the loss of habitat connectivity and promote the survival of carabids, or minor modifications in one possible track could diminish its adverse effects. While many other studies would be needed for a comprehensive assessment of the biotic impact of the highway, we provide an example on the usefulness of network analysis for land use management. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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

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