Modelling multi-species response to landscape dynamics: mosaic cycles support urban biodiversity

The importance of the spatial as well as the temporal structure of habitat patches for urban biodiversity has been recognised, but rarely quantified. In dynamic environments the rate of habitat destruction and recreation (i.e. the landscape turnover rate), the minimum amount of potential habitat, its spatial configuration as well as the environmental conditions determining habitat quality are crucial factors for species occurrence. We analysed species responses to environmental parameters and to the spatio-temporal configuration of urban brownfield habitats in a multi-species approach (37 plant and 43 insect species). Species presence/absence data and soil parameters, site age, vegetation structure and landscape context were recorded by random stratified sampling at 133 study plots in industrial areas in the city of Bremen (Germany). Based on the field data, we predicted species occurrences by species distribution models using a multi-model inference approach. Predicted species communities were driven by successional age both at the scale of a single building lot and at the landscape scale. Minimum average succession time of brownfield habitats required to support all and especially regionally rare species depended on the proportion of available open space; the larger the potential habitat area the faster the acceptable turnover. Most plant, grasshopper, and leafhopper species modelled could be maintained at an intermediate turnover rate (mean age of 10–15 years) and a proportion of open sites of at least 40%. Our modelling approach provides the opportunity of inferring optimal spatio-temporal landscape configurations for urban conservation management from patch scale species-environment relationships. The results indicate that urban planning should incorporate land use dynamics into the management of urban biodiversity. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Green Veining: Landscape Determinants of Biodiversity in European Agricultural Landscapes

Many semi-natural landscape elements, the so-called green veining, are disappearing from the intensively used agricultural landscapes of Europe. In order to develop or restore biodiversity in these networks, it is necessary to quantify the relation between biodiversity and amount, spatial arrangement and management intensity of green veining elements. In this review, we investigate whether biodiversity increases with the amount of green veining in an agricultural landscape following the species–area relationship, and whether a certain level of biodiversity can be reached at lower densities of green veining if green veining elements are better connected (higher spatial connectivity) or if they are managed less intensively (lower management intensity). We reviewed studies on aboveground biodiversity in green veining structures in 39 scientific papers on field and experimental studies within Europe. More of these studies focussed on management intensity than on amount or spatial configuration of green veining. Also more studies focussed on the spatial scale of individual landscape elements than on the farm or landscape scale, which may be caused by the large number of studies focussing on plant or invertebrate species. Species living at larger spatial scales, e.g. mammals and birds were not often studied at the level of green veining elements as they also use agricultural fields as part of their habitat. We could not verify the species–area relation for green veining, nor the effect of amount, spatial configuration or management intensity on this relation, because only few studies quantified the found effects and no studies were found on the effect of management intensity or spatial configuration on the species–area curve in green veining. We addressed the most important challenges for future field and model research in order to fill the identified gaps in knowledge.     

Impact of scale on morphological spatial pattern of forest

Assessing and monitoring landscape pattern structure from multi-scale land-cover maps can utilize morphological spatial pattern analysis (MSPA), only if various influences of scale are known and taken into account. This paper lays part of the foundation for applying MSPA analysis in landscape monitoring by quantifying scale effects on six classes of spatial patterns called: core, edge, perforation, branch, connector and islet. Four forest maps were selected with different forest composition and configuration. The sensitivity of MSPA to scale was studied by comparing frequencies of pattern classes in total forest area for various combinations of pixel size (P) and size parameter (S). It was found that the quantification of forest pattern with MSPA is sensitive to scale. Differences in initial composition and configuration influence the amount but not the general tendencies of the variations of morphological spatial pattern (MSP) class proportions with scale. Increase of P led to data generalization resulting in either a removal of the small size features or their potential transformation into other non-core MSP classes, while an increase of S decreases the MSP core area and this process may transform small core areas into the MSP class islet. We established that the behavior of the MSPA classes with changing scale can be categorized as consistent and robust scaling relations in the forms of linear, power, or logarithmic functions over a range of scales.     

Independent effects of connectivity predict homing success by northern flying squirrel in a forest mosaic

Landscape composition and configuration, often termed as habitat loss and fragmentation, are predicted to reduce species population viability, partly due to the restriction of movement in the landscape. Unfortunately, measuring the effects of habitat loss and fragmentation on functional connectivity is challenging because these variables are confounded, and often the motivation for movement by target species is unknown. Our objective was to determine the independent effects of landscape connectivity from the perspective of a mature forest specialist—the northern flying squirrel (Glaucomys sabrinus). To standardize movement motivation, we translocated 119 squirrels, at varying distances (0.18–3.8 km) from their home range across landscapes representing gradients in both habitat loss and fragmentation. We measured the physical connectedness of mature forest using an index of connectivity (landscape coincidence probability). Patches were considered connected if they were within the mean gliding distance of a flying squirrel. Homing success increased in landscapes with a higher connectivity index. However, homing time was not strongly predicted by habitat amount, connectivity index, or mean nearest neighbour and was best explained as a simple function of sex and distance translocated. Our study shows support for the independent effects of landscape configuration on animal movement at a spatial scale that encompasses several home ranges. We conclude that connectivity of mature forest should be considered for the conservation of some mature forest specialists, even in forest mosaics where the distinction between habitat and movement corridors are less distinct.     

Functional connectivity defined through cost-distance and genetic analyses: a case study for the rock-dwelling mountain vizcacha (<Emphasis Type="Italic">Lagidium viscacia</Emphasis>) in Patagonia,Argentina

Landscape connectivity can have profound consequences for distribution and persistence of populations and metapopulations. Evaluating functional connectivity of a landscape for a species requires a measure of dispersal rates through landscape elements at a spatial scale sufficient to encompass movement capabilities of individuals over the entire landscape. We evaluated functional connectivity for a rock-dwelling mammal, the mountain vizcacha (Lagidium viscacia), in northern Patagonia. Because of the strict association of mountain vizcachas with rocks, we hypothesized that connectivity for this species would be influenced by geology. We used molecular genetic estimates of gene flow to test spatially explicit models of connectivity created with GIS cost-distance analysis of landscape resistance to movement. We analyzed the spatial arrangement of cliffs with join counts and local k-function analyses. We did not capture and genotype individuals, but sampled at the population level through non-invasive collection of feces of mountain vizcachas. The model of landscape connectivity for mountain vizcachas based on geology was corroborated by the pattern of genetic structure, supporting the hypothesis that functional connectivity for mountain vizcachas is influenced by geology, particularly by the distribution of appropriate volcanic rocks. Analysis of spatial arrangement of cliffs indicated that occupied cliffs are clustered and confirmed that rivers act as barriers to dispersal for mountain vizcachas. Our methods could be used, within certain constraints, to study functional landscape connectivity in other organisms, and may be particularly useful for cryptic or endangered species, or those that are difficult or expensive to capture.     

Habitat suitability modelling to correlate gene flow with landscape connectivity

Landscape connectivity is important in designing corridor and reserve networks. Combining genetic distances among individuals with least-cost path (LCP) modelling helps to correlate indirect measures of gene flow with landscape connectivity. Applicability of LCP modelling, however, is reduced if knowledge on dispersal pathways or routes is lacking. Therefore, we integrated habitat suitability modelling into LCP analysis to avoid the subjectivity common in LCP analyses lacking knowledge on dispersal pathways or routes. We used presence-only data and ecological niche factor analysis to model habitat suitability for the spiny rat, Niviventer coninga, in a fragmented landscape of western Taiwan. We adapted the resultant habitat suitability map for incorporation into LCP analyses. Slightly increased Mantel correlations indicated that a class-weighted suitability map better explained genetic distances among individuals than did geographical distances. The integration of habitat suitability modelling into LCP analysis can thus generate information on distribution of suitable habitats, on potential routes of dispersal, for placement of corridors, and evaluate landscape connectivity. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Landscape patterns from mathematical morphology on maps with contagion

The perceived realism of simulated maps with contagion (spatial autocorrelation) has led to their use for comparing landscape pattern metrics and as habitat maps for modeling organism movement across landscapes. The objective of this study was to conduct a neutral model analysis of pattern metrics defined by morphological spatial pattern analysis (MSPA) on maps with contagion, with comparisons to phase transitions (abrupt changes) of patterns on simple random maps. Using MSPA, each focal class pixel on a neutral map was assigned to one of six pattern classes—core, edge, perforated, connector, branch, or islet—depending on MSPA rules for connectivity and edge width. As the density of the focal class (P) was increased on simple random maps, the proportions of pixels in different pattern classes exhibited two types of phase transitions at threshold densities (0.41 ≤ P ≤ 0.99) that were predicted by percolation theory after taking into account the MSPA rules for connectivity and edge width. While there was no evidence of phase transitions on maps with contagion, the general trends of pattern metrics in relation to P were similar to simple random maps. Using an index P for comparisons, the effect of increasing contagion was opposite that of increasing edge width.     

Spatial dynamics of coastal forest bird assemblages: the influence of landscape context,forest type,and structural connectivity

Context

Complex structural connectivity patterns can influence the distribution of animals in coastal landscapes, particularly those with relatively large home ranges, such as birds. To understand the nuanced nature of coastal forest avifauna, where there may be considerable overlap in assemblages of adjacent forest types, the concerted influence of regional landscape context and vegetative structural connectivity at multiple spatial scales warrants investigation.

Objectives

This study determined whether species compositions of coastal forest bird assemblages differ with regional landscape context or with forest type, and if this is influenced by structural connectivity patterns measured at multiple spatial scales.

Methods

Three replicate bird surveys were conducted in four coastal forest types at ten survey locations across two regional landscape contexts in northeast Australia. Structural connectivity patterns of 11 vegetation types were quantified at 3, 6, and 12 km spatial scales surrounding each survey location, and differences in bird species composition were evaluated using multivariate ordination analysis.

Results

Bird assemblages differed between regional landscape contexts and most coastal forest types, although Melaleuca woodland bird assemblages were similar to those of eucalypt woodlands and rainforests. Structural connectivity was primarily correlated with differences in bird species composition between regional landscape contexts, and correlation depended on vegetation type and spatial scale.

Conclusions

Spatial scale, landscape context, and structural connectivity have a combined influence on bird species composition. This suggests that effective management of coastal landscapes requires a holistic strategy that considers the size, shape, and configuration of all vegetative components at multiple spatial scales.

     

Functional rather than structural connectivity explains grassland plant diversity patterns following landscape scale habitat loss

Context

Functional connectivity is vital for plant species dispersal, but little is known about how habitat loss and the presence of green infrastructure interact to affect both functional and structural connectivity, and the impacts of each on species groups.

Objectives

We investigate how changes in the spatial configuration of species-rich grasslands and related green infrastructure such as road verges, hedgerows and forest borders in three European countries have influenced landscape connectivity, and the effects on grassland plant biodiversity.

Methods

We mapped past and present land use for 36 landscapes in Belgium, Germany and Sweden, to estimate connectivity based on simple habitat spatial configuration (structural connectivity) and accounting for effective dispersal and establishment (functional connectivity) around focal grasslands. We used the resulting measures of landscape change to interpret patterns in plant communities.

Results

Increased presence of landscape connecting elements could not compensate for large scale losses of grassland area resulting in substantial declines in structural and functional connectivity. Generalist species were negatively affected by connectivity, and responded most strongly to structural connectivity, while functional connectivity determined the occurrence of grassland specialists in focal grasslands. Restored patches had more generalist species, and a lower density of grassland specialist species than ancient patches.

Conclusions

Protecting both species rich grasslands and dispersal pathways within landscapes is essential for maintaining grassland biodiversity. Our results show that increases in green infrastructure have not been sufficient to offset loss of semi-natural habitat, and that landscape links must be functionally effective in order to contribute to grassland diversity.

     

Spatial spread of an alien tree species in a heterogeneous forest landscape: a spatially realistic simulation model

The effect of environmental heterogeneity on spatial spread of invasive species has received little attention in the literature. Altering landscape heterogeneity may be a suitable strategy to control invaders in man-made landscapes. We use a population-based, spatially realistic matrix model to explore mechanisms underlying the observed invasion patterns of an alien tree species, Prunus serotina Ehrh., in a heterogeneous managed forest. By altering several parameters in the simulation, we test for various hypotheses regarding the role of several mechanisms on invasion dynamics, including spatial heterogeneity, seed dispersers, site of first introduction, large-scale natural disturbances, and forest management. We observe that landscape heterogeneity makes the invasion highly directional resulting from two mechanisms: (1) irregular jumps, which occur rarely via long-distance dispersers and create new founder populations in distant suitable areas, and (2) regular, continuous diffusion toward adjacent cells via short- and mid-distance vectors. At the landscape scale, spatial heterogeneity increases the invasion speed but decreases the final invasion extent. Hence, natural disturbances (such as severe storms) appear to facilitate invasion spread, while forest management can have contrasting effects such as decreasing invasibility at the stand scale by increasing the proportion of light interception at the canopy level. The site of initial introduction influences the invasion process but without altering the final outcome. Our model represents the real landscape and incorporates the range of dispersal modes, making it a powerful tool to explore the interactions between environmental heterogeneity and invasion dynamics, as well as for managing plant invaders. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Effects of changing spatial resolution on the results of landscape pattern analysis using spatial autocorrelation indices

Understanding the relationship between pattern and scale is a central issue in landscape ecology. Pattern analysis is necessarily a critical step to achieve this understanding. Pattern and scale are inseparable in theory and in reality. Pattern occurs on different scales, and scale affects pattern to be observed. The objective of our study is to investigate how changing scale might affect the results of landscape pattern analysis using three commonly adopted spatial autocorrelation indices,i.e., Moran Coefficient, Geary Ratio, and Cliff-Ord statistic. The data sets used in this study are spatially referenced digital data sets of topography and biomass in 1972 of Peninsular Malaysia. Our results show that all three autocorrelation indices were scale-dependent. In other words, the degree of spatial autocorrelation measured by these indices vary with the spatial scale on which analysis was performed. While all the data sets show a positive spatial autocorrelation across a range of scales, Moran coefficient and Cliff-Ord statistic decrease and Geary Ratio increases with increasing grain size, indicating an overall decline in the degree of spatial autocorrelation with scale. The effect of changing scale varies in their magnitude and rate of change when different types of landscape data are used. We have also explored why this could happen by examining the formulation of the Moran coefficient. The pattern of change in spatial autocorrelation with scale exhibits threshold behavior,i.e., scale effects fade away after certain spatial scales are reached (for elevation). We recommend that multiple methods be used for pattern analysis whenever feasible, and that scale effects must be taken into account in all spatial analysis.     

Influence of landscape structure and stand age on species density and biomass of a tropical dry forest across spatial scales

Three central related issues in ecology are to identify spatial variation of ecological processes, to understand the relative influence of environmental and spatial variables, and to investigate the response of environmental variables at different spatial scales. These issues are particularly important for tropical dry forests, which have been comparatively less studied and are more threatened than other terrestrial ecosystems. This study aims to characterize relationships between community structure and landscape configuration and habitat type (stand age) considering different spatial scales for a tropical dry forest in Yucatan. Species density and above ground biomass were calculated from 276 sampling sites, while land cover classes were obtained from multi-spectral classification of a Spot 5 satellite imagery. Species density and biomass were related to stand age, landscape metrics of patch types (area, edge, shape, similarity and contrast) and principal coordinate of neighbor matrices (PCNM) variables using regression analysis. PCNM analysis was performed to interpret results in terms of spatial scales as well as to decompose variation into spatial, stand age and landscape structure components. Stand age was the most important variable for biomass, whereas landscape structure and spatial dependence had a comparable or even stronger influence on species density than stand age. At the very broad scale (8,000–10,500 m), stand age contributed most to biomass and landscape structure to species density. At the broad scale (2,000–8,000 m), stand age was the most important variable predicting both species density and biomass. Our results shed light on which landscape configurations could enhance plant diversity and above ground biomass.     

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.     

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

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

Landscape,community, countryside: linking biophysical and social scales in US Corn Belt agricultural landscapes

Understanding the interplay between ecological and social factors across multiple scales is integral to landscape change initiatives in productive agricultural regions such as the rural US Corn Belt. We investigated the cultural context surrounding the use of perennial cover types—such as stream buffers, wetlands, cellulosic bioenergy stocks, and diverse cropping rotations—to restore water quality, biodiversity, and ecosystem function within a Corn Belt agricultural mosaic in Iowa, USA. Through ethnographic techniques and 33 in-depth interviews, we examined what was most important to rural stakeholders about their countryside. We then used photo elicitation to probe how interviewees’ assessments of farm practices involving perennial cover types were related to their sense of place. Our interviewees perceived their rural “countryside” as a linked social and biophysical entity, identifying strongly with the farming lifestyle and with networks of people across the landscape. While most interviewees approved of perennial farm practices on marginal agricultural land, implementation of these practices was neither a priority nor strongly assimilated into rural experience and ethics. We identified three scale boundaries in our interviewees’ perception of place which present key challenges and opportunities for landscape change: landscape-community, individual-community, and community-institution. In all cases, community social norms and networks—exhibited at landscape spatial scales—may be instrumental in bridging these boundaries and enabling networks of perennial cover types that span privately owned and operated farms. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Surface metrics: an alternative to patch metrics for the quantification of landscape structure

Modern landscape ecology is based on the patch mosaic paradigm, in which landscapes are conceptualized and analyzed as mosaics of discrete patches. While this model has been widely successful, there are many situations where it is more meaningful to model landscape structure based on continuous rather than discrete spatial heterogeneity. The growing field of surface metrology offers a variety of surface metrics for quantifying landscape gradients, yet these metrics are largely unknown and/or unused by landscape ecologists. In this paper, we describe a suite of surface metrics with potential for landscape ecological application. We assessed the redundancy among metrics and sought to find groups of similarly behaved metrics by examining metric performance across 264 sample landscapes in western Turkey. For comparative purposes and to evaluate the robustness of the observed patterns, we examined 16 different patch mosaic models and 18 different landscape gradient models of landscape structure. Surface metrics were highly redundant, but less so than patch metrics, and consistently aggregated into four cohesive clusters of similarly behaved metrics representing surface roughness, shape of the surface height distribution, and angular and radial surface texture. While the surface roughness metrics have strong analogs among the patch metrics, the other surface components are largely unique to landscape gradients. We contend that the surface properties we identified are nearly universal and have potential to offer new insights into landscape pattern–process relationships. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Influence of within-field and landscape factors on aphid predator populations in wheat

The influence of prey density, within-field vegetation, and the composition and patchiness of the surrounding landscape on the abundance of insect predators of cereal aphids was studied in wheat fields in eastern South Dakota, USA. Cereal aphids, aphid predators, and within-field vegetation were sampled in 104 fields over a three year period (1988–1990). The composition and patchiness of the landscape surrounding each field were determined from high altitude aerial photographs. Five landscape variables, aggregated at three spatial scales ranging from 2.6 km² to 581 km², were measured from aerial photographs. Regression models incorporating within-field and landscape variables accounted for 27–49% of the variance in aphid predator abundance in wheat fields. Aphid predator species richness and species diversity were also related to within-field and landscape variables. Some predators were strongly influenced by variability in the composition and patchiness of the landscape surrounding a field at a particular spatial scale while others responded to variability at all scales. Overall, predator abundance, species richness, and species diversity increased with increasing vegetational diversity in wheat fields and with increasing amounts of non-cultivated lands and increasing patchiness in the surrounding landscape.     

The cross-level impact of landscape patterns on housing premiums in micro-neighborhoods

This paper attempts to examine the impacts of the spatial structure of landscape on housing premiums in Austin, TX through the years of 2009–2011. Dissimilar to previous studies, this paper highlighted three points. First, landscape patterns was observed—not only the composition but also the configuration in terms of shape, fragmentation, isolation, and connectivity. Second, the definition of neighborhood in this study was closely matched to the nature of a residential neighborhood, which was represented by a residential subdivision that has a name and a high level of homogeneous characteristics (i.e., development age, appraised value, lot size, and housing size). This approach specifically helps understand the impact of landscape configuration at the micro-level neighborhoods. Third, the hierarchical nature of neighborhoods and embedded parcels—nested parcels within the same neighborhood posit a certain level of identity as a whole—was given attention so that multi-level modeling could be employed. Conceptually and theoretically, multi-level modeling is a better approach to examine phenomenon that occur in multi-level units that show a clear hierarchy. The findings indicated that home buyers are willing to pay more to live in neighborhoods with the high ratio of tree cover. This is a finding consistent with previous studies. Meanwhile, the detailed spatial configuration of landscape does not play an important role at the residential neighborhood level. This result urges policy makers to be more scale-sensitive when planning landscape and indicates that micro neighborhoods are not the correct spatial level to discuss the spatial configuration of landscape, greenways, and green network.     

Landscape ecology of <Emphasis Type="Italic">Phragmites australis</Emphasis> invasion in networks of linear wetlands

The interaction between landscape structure and spatial patterns of plant invasion has been little addressed by ecologists despite the new insights it can provide. Because of their spatial configuration as highly connected networks, linear wetlands such as roadside or agricultural ditches, can serve as corridors facilitating invasion at the landscape scale, but species dynamics in these important habitats are not well known. We conducted a landscape scale analysis of Phragmites australis invasion patterns (1985–2002 and 1987–2002) in two periurban areas of southern Québec (Canada) focusing on the interaction between the network of linear wetlands and the adjacent land-uses. Results show that, at the beginning of the reference period, the two landscapes were relatively non-invaded and populations occurred mostly in roadside habitats which then served as invasion foci into other parts of the landscape. The intrinsic rates of increase of P. australis populations in linear anthropogenic habitats were generally higher than those reported for natural wetlands. Riparian habitats along streams and rivers were little invaded compared to anthropogenic linear wetlands, except when they intersected transportation rights-of-way. Bivariate spatial point pattern analysis of colonization events using both Euclidian and network distances generally showed spatial dependence (association) to source populations. An autologistic regression model that included landscape and edaphic variables selected transportation rights-of-way as the best predictor of P. australis occurrence patterns in one of the landscapes. Given the high invasion rates observed, managers of linear wetlands should carefully monitor expansion patterns especially when roads intersect landscapes of conservation or economic value.