Neutral models for testing landscape hypotheses

Neutral landscape models were originally developed to test the hypothesis that human-induced fragmentation produces patterns distinctly different from those associated with random processes. Other uses for neutral models have become apparent, including the development and testing of landscape metrics to characterize landscape pattern. Although metric development proved to be significant, the focus on metrics obscured the need for iterative hypothesis testing fundamental to the advancement of the discipline. We present here an example of an alternative neutral model and hypothesis designed to relate the process of landscape change to observed landscape patterns. The methods and program, QRULE, are described and options for statistical testing outlined. The results show that human fragmentation of landscapes results in a non-random association of land-cover types that can be describe by simple statistical methods. Options for additional landscape studies are discussed and access to QRULE described in the hope that these methods will be employed to advance our understanding of the processes that affect the structure and function in human dominated landscapes.     

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.     

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.     

Evaluation of patch isolation metrics in mosaic landscapes for specialist vs. generalist dispersers

We examined the effects of matrix structure and movement responses of organisms on the relationships between 7 patch isolation metrics and patch immigration. Our analysis was based on simulating movement behaviour of two generic disperser types (specialist and generalist) across mosaic landscapes containing three landcover types: habitat, hospitable matrix and inhospitable matrix. Movement, mortality and boundary crossing probabilities of simulated individuals were linked to the landcover and boundary types in the landscapes. The results indicated that area-based isolation metrics generally predict patch immigration more reliably than distance-based isolation metrics. Relationships between patch isolation metrics and patch immigration varied between the two generic disperser types and were affected by matrix composition or matrix fragmentation. Patch immigration was always affected by matrix composition but not by matrix fragmentation. Our results do not encourage the generic use of patch isolation metrics as a substitute for patch immigration, in particular in metapopulation models where generic use may result in wrong projections of the survival probability of metapopulations. However, our examination of the factors affecting the predictive potential of patch isolation metrics should facilitate interpretation and comparison of existing patch isolation studies. Future patch isolation studies should include information about landscape structure and the dispersal distance and dispersal behaviour of the organism of interest.This revised version was published online in May 2005 with corrections to the Cover Date.     

Landscape cohesion: an index for the conservation potential of landscapes for biodiversity

In urbanising landscapes, planning for sustainable biodiversity occurs in a context of multifunctional land use. Important conditions for species persistence are habitat quality, the amount and configuration of habitat and the permeability of the landscape matrix. For planning purposes, these determinants should be integrated into simple indicators for spatial conditions of persistence probability. We propose a framework of three related indices. The cohesion index is based on the ecology of metapopulations in a habitat network. We discuss how an indicator for species persistence in such a network could be developed. To translate this network index into an area index, we propose the concept of spatial cohesion. Habitat cohesion and spatial cohesion are defined and measured for single species or, at best, for species profiles. Since species differ in their perception of the same landscape, different species will rate different values of these indices for the same landscape. Because landscapes are rarely planned for single species, we further propose the index of landscape cohesion, which integrates the spatial cohesion indices of different species. Indices based on these concepts can be built into GIS tools for landscape assessment. We illustrate different applications of these indices, and emphasise the distinction between ecological and political decisions in developing and applying such tools. This revised version was published online in August 2006 with corrections to the Cover Date.     

Implications of rescaling rules for multi-scaled habitat models

Recent work in landscape ecology suggests that organisms use environmental cues at a variety of scales to select habitat. As a result, habitat studies that evaluate environmental conditions at multiple spatial scales have become increasingly common. We examined whether the way in which data are rescaled influences inferred relationships between organisms and habitat features. Using a habitat model developed at fine scales, we systematically rescaled habitat (canopy density, slope, and cover type) and distribution maps according to a variety of different rescaling rules, including spatial averaging, thresholding, presence/absence, and majority. We found that the spatial autocorrelation of habitat data interacts with rescaling rules to alter the correspondence between species presence and habitat across scale. Different rules lead to substantially divergent and sometimes opposite correlations among the species and habitat features on the landscape. Such differences in interpretation due to variation in methodology can lead to very different interpretations of a species habitat requirements and thus have important implications for both ecology and conservation.     

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

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

Spatial and non-spatial factors: When do they affect landscape indicators of watershed loading?

The percentage of a watershed occupied by agricultural areas is widely used to predict nutrient loadings and in-stream water chemistry because water quality is often linked to non-point sources in a watershed. Measures of the spatial location of source areas have generally not been incorporated into such landscape indicators although empirical evidence and watershed loading models suggest that spatially explicit information is useful for predicting loadings. I created a heuristic grid-based surface-flow model to address the discrepancies between spatially explicit and non-spatial approaches to understanding watershed loading. The mean and variance in loading were compared among thousands of simulated watersheds with varying percentages of randomly located source and sinks. The variability in loading among replicate landscapes was greatest for those landscapes with ~65% source areas. This variance peak suggests that considering the spatial arrangement of cover types is most important for watersheds with intermediate relative abundances of sources and sinks as the wide variety of different spatial configurations can lead to either very high or very low loading. Increasing the output from source pixels (relative to the amount absorbed by sink pixels) among different landscapes moved the peak in variance to landscapes with lower percentages of sources. A final scenario examined both broad- and fine-scale heterogeneity in source output to disentangle the relative contributions of spatial configuration, percentage of source covers, and heterogeneity of sources in governing variability in loading. In landscapes with high percentages of source pixels, fine-scale heterogeneity in source output was responsible for a greater portion of the total variability in loading among different watersheds than was spatial arrangement. These results provide several testable hypotheses for when spatial and non-spatial approaches might be most useful in relating land cover to water chemistry and suggest improvements for the spatial sensitivity analyses of eco-hydrologic watershed models.     

The Importance of Spatial Scale for Conservation and Assessment of Anuran Populations in Coastal Wetlands of the Western Great Lakes,USA

Distributions of pond-breeding amphibians may be influenced by habitat factors at different spatial scales. We used anuran calling surveys to investigate the association between 5 anuran species and habitat variables measured within 100, 500, 1000, and 3000 m of sampling points at 63 coastal wetlands along the US shores of Lake Michigan and Lake Huron. Stepwise logistic regression was used to create predictive models for each species at each spatial scale. Our results confirm the view that habitat variables at multiple scales influence frog distributions, but the strength of predictive models was significantly affected by the spatial scale at which habitat variables were derived. Remotely sensed habitat variables within a 3000 m radius were among the most effective predictors of occurrence for American toad (Bufo americanus), eastern gray treefrog (Hyla versicolor), spring peeper (Pseudacris crucifer), and green frog (Rana clamitans melanota). The western chorus frog (Pseudacris triseriata) was predicted most effectively by variables derived within a 500 m radius. For the most part, these anurans exhibited species-specific responses to habitat variables; however the suite of landscape-scale variables associated with urban land use appeared in all species’ regression models. Associations with landscape-scale variables coupled with well-documented habitat needs at local breeding sites suggest that conservation and assessment of frogs and toads in coastal wetlands should consider the influence of habitat variables at multiple spatial scales.     

Long-term decline and local extinction of Clethrionomys rufocanus in boreal Sweden

Over the past three decades in boreal Sweden, there has been a long-term decline of cyclic sympatric voles, leading to local extinctions of the most affected species, the grey-sided vole (Clethrionomys rufocanus). We monitored this decline by snap-trapping on 58 permanent plots spread over 100 km² in spring and fall from fall 1971–2003. The reason for the decline is largely unknown, although a common major factor is likely to be involved in the decline of C. rufocanus and of the coexisting voles. However, here we deal with the reasonability of one complementary hypothesis, the habitat fragmentation hypothesis, which assumes that part of the decline of C. rufocanus is caused by habitat (forest) destruction. There was considerable local variation in the decline among the 58 1-ha sampling plots, with respect to both density and timing of the decline; however, all declines ended up with local extinction almost without exception. Local declines were not associated with habitat destruction by clear-cutting within sampling-plots, as declines started about equally often before as after clear-cutting, which suggested that habitat destruction outside sampling plots could be involved. In a multiple regression analysis, local habitat preference (LHP; expressed as a ratio of observed to expected number of voles trapped per habitat) together with two habitat variables in the surrounding (2.5×2.5 km²) landscape matrix explained 56% of the variation among local cumulated densities of C. rufocanus and hence of local time-series. LHP was positively correlated and explained 31% of the variation, while connectivity among clear-cuts was negatively correlated and proximity among xeric-mesic mires was positively correlated and explained additional 16% and 9%, respectively. Even if the overall decline cannot be connected to local clear-cutting on sampling-plots, clear-cutting and hence habitat fragmentation/destruction in the surrounding landscapes potentially influenced grey-sided vole numbers negatively.     

The importance of landscape context for songbirds on migration: body mass gain is related to habitat cover

Landscape context influences many aspects of songbird ecology during the breeding season. The importance of landscape context at stopover sites for migrating songbirds, however, has received less attention. In particular, landscape context may affect the availability and quality of food for refueling during stopovers, which is critical for successful migration. We evaluated the influence of woody habitat cover in the surroundings of stopover sites at several spatial extents on the hourly changes of body mass in two species of European-African forest-dwelling songbird migrants (Willow Warbler, Phylloscopus trochilus, and the Eurasian Redstart, Phoenicurus phoenicurus). Data were sampled by standardized methods from a network of ringing stations throughout Europe during the falls of 1994–1996. In both species, hourly body mass gain calculated for first captures increased with woody habitat cover. We found a similar logarithmic relationship for both species, although for Willow Warblers mass gain was more strongly related to the habitat cover within 5 km, in contrast to 3 km for Redstarts. For Willow Warblers, where sufficient data are available for each year, the relationship is consistent over the years. The shape of the relationship suggests existence of a threshold of landscape suitability for refueling at stopover sites: in sites with less than 10% of woody habitat cover, birds tend to lose body mass or to gain mass at a lower rate. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Evolving core-periphery interactions in a rapidly expanding urban landscape: The case of Beijing

We characterized and analyzed the dynamics of a rapidly expanding urban landscape of Beijing Municipality, based on the Hierarchical Regional Space (HRS) model. We focused on ecological processes such as flows of energy, materials and population between the urban core and its periphery, and how these processes co-evolved with urbanization. We treated the HRS as an alternative to the cellular automata (CA) approach to characterizing and modeling of landscape dynamics. With LANDSAT data, we showed that the urban area of Beijing expanded from 269 km² to 901 km² in the period from 1975 to 1997, an increase of 2.35 times in 22 years. Meanwhile, a number of secondary urban centers formed on areas that used to be sparsely populated around the city. These secondary centers quickly expanded and ultimately merged with each other and with the urban core. The changes in spatial pattern and organization were accompanied by evolution of urban functions and particularly the interactions between the urban core and its periphery. We demonstrated a dramatic increase in dependence of the urban core on the periphery as well as the cores influence on the periphery with a case analysis of the vegetable supply to Beijing. The tightening link between the city and its periphery reinforces the urbanization process and further drives the transformation of the regions landscape. We conclude that the HRS model is capable of characterizing the patterns and processes of complex and dynamic landscapes such as the case of Beijing, and this model has great potential for quantitative modeling of human dominated landscapes as well.This revised version was published online in May 2005 with corrections to the Cover Date.     

A multi-scaled analysis of avian response to habitat amount and fragmentation in the Canadian dry mixed-grass prairie

Previous research has suggested that ducks and songbirds may benefit from prairie landscapes that consist primarily of contiguous grasslands. However, the relative importance of landscape-level vs. local characteristics on mechanisms underlying observed patterns is unclear. We measured effects of grassland amount and fragmentation on upland and wetland songbird and duck density and nest success, and on some nest predators, across 16 landscapes in southern Alberta, Canada. We compared these landscape-level effects with local-scale responses, including distance to various edges and vegetation characteristics. We also evaluated several statistical approaches to comparing effects of habitat characteristics at multiple spatial scales. Few species were influenced by grassland amount or fragmentation. In contrast, distance to edge and local vegetation characteristics had significant effects on densities and nest success of many species. Previous studies that reported effects of landscape characteristics may have detected patterns driven by local mechanisms. As a corollary, results were very sensitive to statistical model structure; landscape level effects were much less apparent when local characteristics were included in the models.     

Effect of matrix habitat on the spread of flea beetle introductions for biological control of leafy spurge

Matrix habitats are known to influence the movement patterns of a variety of species but it is less well known whether these effects have strong implications for spatial population dynamics, including the spread of biological introductions. Using a spatially explicit simulation model parameterized with empirical data, we examine how grass and shrub matrix habitats, each offering different resistance to dispersal, influence the spread and impact of a biocontrol agent, Aphthona lacertosa, on the invasive weed, leafy spurge. Model predictions indicate that differential responses to matrix habitat have little effect on the agent’s spread over the study landscape and this is supported by statistical models fit to observed A. lacertosa incidence on the same landscape. Subsequent experimentation with the simulation model suggested that A. lacertosa colonization rates were largely unaffected by increases in amount of the more restrictive shrub matrix. However, simulations of an hypothetical species with greater overall dispersal ability but reduced dispersal rate through shrub matrix showed that colonization rates were noticeably reduced when the percentage of shrub matrix on the landscape approached 50%. Combined these results suggest that some tailoring of release strategies may be required to accommodate the unique dispersal capabilities of different biocontrol agents on particular release landscapes, but for A. lacertosa there appears to be little effect of matrix habitat structure on rates of spread.