Scale of heterogeneity of forage production and winter foraging by elk and bison

The relationship between fine-scale spatial patterns of forage abundance and the feeding patterns of large ungulates is not well known. We compared these patterns for areas grazed in winter by elk and bison in a sagebrush-grassland landscape in northern Yellowstone National Park. At a fine scale, the spatial distribution of mapped feeding stations in 30 m × 30 m sites was found to be random where there were no large patches devoid of vegetation. In areas similar to the mapped sites, the underlying spatial distribution pattern of biomass was also determined to be random. At a broad scale, forage biomass differed among communities across the northern range but forage quality did not. These results suggest that ungulates are feeding randomly within forage patches (fine scale) but may select feeding sites based upon forage abundance at broader, landscape scales. Contrary to what has been suggested in other systems, ungulates were not overmatching at finer scales.     

Deciduous forest and resident birds: the problem of fragmentation within a coniferous forest landscape

Six species of resident birds were censused in patches of deciduous forest within a coniferous forest landscape in south central Sweden. Here, the forests have been subjected to active forestry for a long time, but with recently increased intensity. Although the forest cover is more or less continuous in this landscape, mature deciduous forest is now a rare element compared with the untouched forest.All censused patches were similar with regards to size, proportion and amount of deciduous trees, but were either isolated in the coniferous forest (isolated patches) or near to other deciduous patches (aggregated patches). We concentrated on six species of resident birds, with moderate area requirements, that are tied to deciduous forest and whose ecology is well-known. The Nuthatch and the Marsh tit, which both show strict year-round territoriality and have a restricted dispersal phase, were significantly more likely to be found in aggregated than in isolated patches. No effect was found for the Great tit and the Blue tit, which are less territorial outside the breeding season and have a longer dispersal phase. Moreover, the Great tit is less specialized on deciduous forest than the other species. Also, the Long-tailed tit was negatively affected by isolation, which may be due to restricted dispersal and to larger area requirements of this flock-territorial species. The Hazel grouse, finally, was not affected, but this larger bird probably uses the forest in a different way from the smaller species.Our study clearly shows that fragmentation of one type of forest (deciduous) within another can have serious detrimental effects on forest-living species and raises important issues for forest management practices and conservation within a forest landscape.     

Using fractal analysis to assess how species perceive landscape structure

To develop a species-centered definition of landscapes, I suggest using a fractal analysis of movement patterns to identify the scales at which organisms are interacting with the patch structure of the landscape. Significant differences in the fractal dimensions of movement patterns of two species indicate that the species may be interacting with the patch structure at different scales. Fractal analysis therefore permits comparisons of landscape perceptions of different species within the same environment.I tested the utility of this fractal application by analyzing the movement patterns of three species of acridid grasshoppers (Orthoptera) in a grassland mosaic. The largest species moved up to 6 times faster than the two smaller species, and species exhibited different responses to microlandscape structure within 25-m² plots. Further, the largest species exhibited different responses to microlandscape structure in two pastures subjected to different intensities of cattle grazing. This species thus is able to integrate information on landscape structure at broad spatial scales. Fractal analysis of movement patterns revealed that the two small species had significantly more tortuous patterns than the larger species, which suggests that these species are interacting with patch structure at a finer scale of resolution than the large species. Fractal analysis can be used to identify the perceptive resolution of a species; that is, the spatial grain and extent at which they are able to perceive and respond to heterogeneity. Analysis of movement patterns across a range of spatial scale may reveal shifts in fractal dimension that reflect transitions in how species respond to the patch structure of the landscape at different scales.     

Fire-driven dynamic mosaics in the Great Victoria Desert,Australia – II. A spatial and temporal landscape model

An explicitly spatial, large scale, high resolution model of fire driven landscape dynamics in the Great Victoria Desert is constructed and parameterized to simulate frequency distributions of fire size and shape obtained from previous analyses of satellite chronosequences. We conclude that probabilities of fire spread cannot be constant over time, and that realistic distributions of fire size and plausible rates of fire spread can be obtained by assuming that fire spread is conditional on observed durations of windy conditions. Landscapes subject to this form of disturbance show large scale correlation structure many times greater than the average dimensions of single fires, and exhibit low frequency quasi-periodic stochastically driven oscillations in proportions of the landscape at different successional states over spatial scales exceeding 100,000 km². Average fire return intervals are 30 yrs. Analysis of patch structure suggests that this landscape is composed of few large younger patches, embedded in a mature sea of surrounding habitat. Intermediate and late successional habitat must exist in more abundant patches somewhat smaller than young habitat. Numerous small patches of mature habitat are likely to be scattered throughout this younger habitat. The model predicts that fire size frequency distributions are relatively insensitive to changes of as much as ±50% of observed fire ignition frequency.     

The ocean ‘landscape’

The ocean has a complex physical structure at all scales in space and time, with peaks at certain wave numbers and frequencies. Pelagic ecosystems show regular progressions in size of organisms, life cycle, spatial ambit, and trophic status. Thus, physiological and ecological parameters are closely coupled to spatial and temporal physical scales.     

Longterm response of disturbance landscapes to human intervention and global change

The structure of landscapes subject to patch-forming catastrophic disturbances, or disturbance landscapes, is controlled by the characteristics of the disturbance regime, including the distribution of disturbance sizes and intervals, and the rotation time. The primary landscape structure in disturbance landscapes is the structure of the mosaic of disturbance patches, which can be described by indices such as patch size and shape.The purpose of this research was to use a geographical information system-based spatial model (DISPATCH) to simulate the effects of the initial density of patches on the rate of response to alteration of a disturbance regime, the effects of global warming and cooling, and the effects of fragmentation and restoration, on the structure of a generalized temperate-zone forested disturbance landscape over a period of 400 yr.The simulations suggest that landscapes require 1/2 to 2 rotations of a new disturbance regime to adjust to that regime regardless of the size and interval distributions. Thus alterations that shorten rotations, as would be the case if global warming increases fire sizes and decreases fire intervals, produce a more rapid response than do alterations that lengthen rotations, such as cooling and fire suppression. Landscape with long rotations may be in perpetual disequilibrium with their disturbance regimes due to a mismatch between their adjustment rate and the rate of climatic change. Landscapes with similar rotation times may have different structures, because size and interval distributions independently affect landscape structure. The response of disturbance landscapes to changing disturbance regimes is governed by both the number and size of patch births.     

Landscape graphs: Ecological modeling with graph theory to detect configurations common to diverse landscapes

In view of the bewildering diversity of landscapes and possible patterns therein, our objectives were to see if a useful modeling method for directly comparing land mosaics could be developed based on graph theory, and whether basic spatial patterns could be identified that are common to diverse landscapes. The models developed were based on the spatial configuration of and interactions between landscape elements (ecosystems, land uses or ecotopes). Nodes represented landscape elements and linkages represented common boundaries between elements. Corridors, corridor intersections, and the matrix were successfully incorporated in the models. Twenty-five landscape graphs were constructed from aerial photographs chosen solely to represent a breadth of climates, land uses and human population densities. Seven distinctive clusters of nodes and linkages were identified and common, three of which, in the forms of a spider, necklace and graph cell, were in >90% of the graphs. These represented respectively the following configurations of patches, corridors and matrix: (1) a matrix area surrounding or adjoining many patches; (2) a corridor bisecting a heterogeneous area; and (3) a unit in a network of intersecting corridors. The models also indicated that the connectivity or number of linkages for several common elements, such as fields and house clearings, was relatively constant across diverse landscapes, and that linear shaped elements such as roads and rivers were the most connected. Several additional uses of this graph modeling, including compatibility with systems dynamics models, are pinpointed. Thus the method is useful in allowing simple direct comparisons of any scale and any landscape to help identify patterns and principles. A focus on the common and uncommon configurations should enhance our understanding of fluxes across landscapes, and consequently the quality of land planning and management.     

Fire-induced changes in northern Patagonian landscapes

In northern Patagonia, Argentina we quantify changes in fire frequency along a gradient from mesic Nothofagus dombeyi forest to xeric woodlands of Austrocedrus chilensis at the steppe ecotone, and we examine patterns of vegetation change coincident with the changes in fire regimes across a range of spatial scales. At a regional scale changes in land cover types are documented by comparing 1:250000 scale cover type maps from 1913 and 1985. Changes in landscape structure are analyzed by comparing vegetation patterns on 1:24000 scale aerial photographs taken in 1940 and 1970. Fire frequency peaked in the late nineteenth-century due to widespread burning and clearing of forests by European settlers late in the century. Subsequently, fire frequency declined dramatically about 1910 due to the cessation of intentional fires and has remained low due to increasingly effective fire exclusion. At a regional scale there has been a dramatic increase during the twentieth century in the proportion of forest cover relative to areas mapped as recent burns or shrublands in 1913. Remnant forest patches that survived the widespread late-nineteenth century burning have coalesced to form more continuous forest covers, and formerly continuous areas of shrublands have become dissected by forest. Under reduced fire frequency there has been a shift in dominance from short-lived resprouting species (mostly shrubs) towards longer-lived species and obligate seed-dispersers such as Austrocedrus chilensis and Nothofagus dombeyi. Due to limited seed dispersal of these tree species, the spatial configuration of remnant forest patches plays a key role in subsequent changes in landscape pattern.     

Effects of sensor spatial resolution on landscape structure parameters

We examined the effects of increasing grain size from 20 m to 1100 m on landscape parameters characterizing spatial structure in the northern Wisconsin lake district. We examined whether structural parameters remain relatively constant over this range and whether aggregation algorithms permit extrapolation within this range. Images from three different satellite sensors were employed in this study: (1) the SPOT multispectral high resolution visible (HRV), (2) the Landsat Thematic Mapper (TM), and (3) the NOAA Advanced Very High Resolution Radiometer (AVHRR). Each scene was classified as patches of water in a matrix of land. Spatial structure was quantified using several landscape parameters: percent water, number of lakes (patches), average lake area and perimeter, fractal dimension, and three measures of texture (homogeneity, contrast, and entropy). Results indicate that most measures were sensitive to changes in grain size. As grain size increased from 20 m using HRV image data to 1100 m (AVHRR), the percent water and the number of lakes decreased while the average lake area, perimeter, the fractal dimension, and contrast increased. The other two texture measures were relatively invariant with grain size. Although examination of texture at various angles of adjacency was performed to investigate features which vary systematically with angle, the angle did not have an important effect on the texture parameter values. An aggregation algorithm was used to simulate additional grain sizes. Grain was increased successively by a factor of two from 20 m (the HRV image) to 1280 m. We then calculated landscape parameter values at each grain size. Extrapolated values closely approximated the actual sensor values. Because the grain size has an important effect on most landscape parameters, the choice of satellite sensor must be appropriate for the research question asked. Interpolation between the grain sizes of different satellite sensors is possible with an approach involving aggregation of pixels.     

Grain-dependent functional responses in habitat selection

Context

Spatial scale is an important consideration for understanding how animals select habitat, and multi-scalar designs in resource selection studies have become increasingly common. Despite this, examination of functional responses in habitat selection at multiple scales is rare. The perceptual range of an animal changes as a function of vegetation association, suggesting that use, selection and functional responses may all be habitat- and scale-dependent.

Objectives

Our objective was to determine how varying grain size affects our interpretation of functional response in habitat selection and to elucidate scalar and landscape effects on habitat selection.

Methods

We quantified the functional response of GPS-collared, female white-tailed deer (Odocoileus virginianus, n = 18) in Riding Mountain National Park, Canada, to different habitat types. Functional responses were quantified at multiple spatial scales by regressing proportion of habitat used against proportion of habitat available at different buffer radii (ranging from 75–1000 m radius) surrounding used (telemetry) locations and available points within the individual’s seasonal home range. We examined how functional responses changed as a function of grain by plotting grain size against the slope of the functional response.

Results

We detected functional responses in most habitat types. As expected, functional responses tended to converge towards 1 (use proportional to availability) at large buffer sizes; however, the relationship between scale and functional response was typically non-linear and depended on habitat type.

Conclusions

We conclude that a multi-scalar approach to modelling animal functional responses in habitat selection is important for understanding patterns in animal behaviour and resource use.

     

Quantifying the effect of grazing and shrub-clearing on small scale spatial pattern of vegetation

Disturbances such as grazing, invading species, and clear-cutting, often act at small spatial scales, and means for quantifying their impact on fine scale vegetation patterns are generally lacking. Here we adopt a set of landscape metrics, commonly used for quantifying coarse scale fragmentation, to quantify fine scale fragmentation, namely the fine scale vegetation structure. At this scale, patches often consist of individual plants smaller than 1 m², requiring the grain of the analysis to be much smaller. We used balloon aerial photographs to map fine details of Mediterranean vegetation (pixel size <0.04 m) in experimental plots subjected to grazing and clear-cutting and in undisturbed plots. Landscape metrics are sensitive to scale. Therefore, we aggregated the vegetation map into four coarser scales, up to a resolution of 1 m, and analyzed the effect of scale on the metrics and their ability to distinguish between different disturbances. At the finest scale, six of the seven landscape metrics we evaluated revealed significant differences between treated and undisturbed plots. Four metrics revealed differences between grazed and control plots, and six metrics revealed differences between cleared and control plots. The majority of metrics exhibited scaling relations. Aggregation had mixed effects on the differences between metric values for different disturbances. The control plots were the most sensitive to scale, followed by grazing and clearing. We conclude that landscape metrics are useful for quantifying the very fine scale impact of disturbance on woody vegetation, assuming that the analysis is based on sufficiently high spatial resolution data.     

Predicting across scales: Theory development and testing

Landscape ecologists deal with processes that occur at a variety of temporal and spatial scales. The ability to make predictions at more than one level of resolution requires identification of the processes of interest and parameters that affect this process at different scales, the development of rules to translate information across scales, and the ability to test these predictions at the relevant spatial and temporal scales. This paper synthesizes discussions from a workshop on Predicting Across Scales: Theory Development and Testing that was held to discuss current research on scaling and to identify key research issues.     

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

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

The influence of research scale on bald eagle habitat selection along the lower Hudson River,New York (USA)

As the concepts of landscape ecology have been incorporated into otherdisciplines, the influence of spatial patterns on animal abundance anddistribution has attracted considerable attention. However, there remains asignificant gap in the application of landscape ecology theories and techniquesto wildlife research. By combining landscape ecology techniques withtraditionalwildlife habitat analysis methods, we defined an organism-centeredperspectivefor breeding bald eagles (Haliaeetus leucocephalus) alongthe Hudson River, New York, USA. We intensively monitored four pairs ofbreedingeagles during the 1999 and 2000 breeding seasons, and collected detailedinformation on perch and forage locations. Our analysis focused on threecritical habitat elements: available perch trees, access to foraging areas, andfreedom from human disturbance. We hypothesized that eagle habitat selectionrelative to each of these elementswould vary with the spatial scale of analysis, and that these scalingrelationships would vary among habitat elements. We investigated two elementsofspatial scale: grain and local extent. Grain was defined as the minimum mappingunit; local extent was defined by the size of an analysis window placed aroundeach focal point. For each habitat element, we quantified habitat use over arange of spatial scales. Eagles displayed scale-dependent patterns of habitatuse in relation to all habitat features, including multi-scale andthreshold-like patterns. This information supports the existence ofscale-dependant relationships in wildlife habitat use and allowed for a moreaccurate and biologically relevant evaluation of Hudson River breeding eagle habitat.This revised version was published online in May 2005 with corrections to the Cover Date.     

Landscape change with agricultural intensification in a rural watershed,southwestern Ohio,U.S.A.

Specialized cash grain production, emergent in the midwestern United States during the post-WWII era, typifies the Upper Four Mile Creek watershed in southwestern Ohio. This style of agriculture intensifies cropland use, with consequent increases in soil erosion and stream sedimentation - a serious problem in the lower reservoir, Acton Lake. Agricultural statistics and aerial photographs compiled between 1934 and 1984 were used to quantify agricultural dynamics and landscape change in the watershed, including land-use apportionment, diversity, and the structural configuration of forest, woodland, and old-field/brushland patches and corridors. A questionnaire sent to all land owners in the basin documented farm-level characteristics and factors that influence management decisions. Crop diversity (H) in Preble County, Ohio decreased from 1.42 in 1934 to 1.17 in 1982, as corn and soybeans dominated the landscape mosaic. Yields rose, but net profits were reduced by declining prices per bushel and increases in fertilizer and petroleum-based subsidies. Landuse diversity in the county also declined (H = 1.37 in 1934 tot 0.80 in 1982) in response to cropland expansion, whereas forest land in the watershed increased from 1605 to 2603 ha. Fragmentation declined and the landscape became polarized after 1956, with a concentration of agricultural patches in the upper watershed and forest-patch coalescence in stream gullies and state park land in the lower watershed. The questionnaire (~ 29% return) further supported, at the farm-level, observed regional trends toward expansion (farm coalescence and lease contracts) and specialization (conversion toward corn and soybeans). The most important factors influencing farm size and management were better equipment and family traditions. Thus, cultural and technological factors that operate at the farm-level, coupled with meso-scale variation in the physical conditions of a catchment basin, tend to influence landscape-level patterns more than regional socioeconomics and governmental policies.     

A model of arctic tundra vegetation derived from topographic gradients

We present a topographically-derived vegetation model (TVM) that predicts the landscape patterns of arctic vegetation types in the foothills of the Brooks Range in northern Alaska. In the Arctic there is a strong relationship between water and plant structure and function and TVM is based on the relationships between vegetation types and slope (tan ) and discharge (), two independent variables that can be easily derived from digital terrain data. Both slope and discharge relate to hydrological similarity within a landscape: slope determines the gravitational hydrological gradient and hence influences flow velocity, whereas discharge patterns are computed based on upslope area and quantify lateral flow amount. TVM was developed and parameterized based on vegetation data from a small 2.2 km² watershed and its application was tested in a larger 22km² region. For the watershed, TVM performed quite well, having a high spatial resolution and a goodness-of-fit ranging from 71–78&percnt;, depending on the functions used. For the larger region, the strength of the vegetation types predictions drops somewhat to between 56–59&percnt;. We discuss the various sources of error and limitations of the model for purposes of extrapolation.     

Spatial scale and pattern dependences of aboveground biomass estimation from satellite images: a case study of the Sierra National Forest,California

Context

Spatial scale and pattern play important roles in forest aboveground biomass (AGB) estimation in remote sensing. Changes in the accuracy of satellite images-estimated forest AGBs against spatial scales and pixel distribution patterns has not been evaluated, because it requires ground-truth AGBs of fine resolution over a large extent, and such data are difficult to obtain using traditional ground surveying methods.

Objectives

We intend to quantify the accuracy of AGB estimation from satellite images on changing spatial scales and varying pixel distribution patterns, in a typical mixed coniferous forest in Sierra Nevada mountains, California.

Methods

A forest AGB map of a 143 km² area was created using small-footprint light detection and ranging. Landsat Thematic Mapper images were chosen as typical examples of satellite images, and resampled to successively coarser resolutions. At each spatial scale, pixels forming random, uniform, and clustered spatial patterns were then sampled. The accuracies of the AGB estimation based on Landsat images associated with varying spatial scales and patterns were finally quantified.

Results

The changes in the accuracy of AGB estimation from Landsat images are not monotonic, but increase up to 60–90 m in spatial scale, and then decrease. Random and uniform spatial patterns of pixel distributions yield better accuracy for AGB estimation than clustered spatial patterns. The corrected NDVI (NDVIc) was the best predictor of AGB estimation.

Conclusions

A spatial scale of 60–90 m is recommended for forest AGB estimation at the Sierra Nevada mountains using Landsat images and those with similar spectral resolutions.

     

Multi-scale network analysis shows scale-dependency of significance of individual protected areas for connectivity

Context

The problem of how ecological mechanisms create and interact with patterns across different scales is fundamental not only for understanding ecological processes, but also for interpretations of ecological dynamics and the strategies that organisms adopt to cope with variability and cross-scale influences.

Objectives

Our objective was to determine the consistency of the role of individual habitat patches in pattern-process relationships (focusing on the potential for dispersal within a network of patches in a fragmented landscape) across a range of scales.

Methods

Network analysis was used to assess and compare the potential connectivity and spatial distribution of highland fynbos habitat in and between protected areas of the Western Cape of South Africa. Connectivity of fynbos patches was measured using ten maximum threshold distances, ranging from five to 50 km, based on the known average dispersal distances of fynbos endemic bird species.

Results

Network connectivity increased predictably with scale. More interestingly, however, the relative contributions of individual protected areas to network connectivity showed strong scale dependence.

Conclusions

Conservation approaches that rely on single-scale analyses of connectivity and context (e.g., based on data for a single species with a given dispersal distance) are inadequate to identify key land parcels. Landscape planning, and specifically the assessment of the value of individual areas for dispersal, must therefore be undertaken with a multi-scale approach. Developing a better understanding of scaling dependencies in fragmenting landscapes is of high importance for both ecological theory and conservation planning.

     

Scale dependence of tree abundance and richness in a tropical rain forest,Malaysia

Abundance and richness are the two fundamental components of speciesdiversity. They represent two distinct types of variables of which the formerisadditive when aggregated across scales while the latter is nonadditive. Thisstudy investigated the changes in the spatial patterns of abundance andrichnessof tree species across multiple scales in a tropical rain forest of Malaysiaandtheir variations in different regions of the study area. The results showedthatfrom fine to coarse scales abundance had a gradual and systematic change inpattern, whereas the change in richness was much less predictable and ahotspot in richness at one scale may become acoldspot at another. The study also demonstrated that differentmeasures of diversity variation (e.g., variance and coefficient of variation)can result in different or even contradictory results which further complicatedthe interpretation of diversity patterns. Because of scale effect the commonlyused measure of species diversity in terms of unit area (e.g.,species/m²) is misleading and of little use in comparing speciesdiversitybetween different ecosystems. Extra care must be taken if management andconservation of species diversity have to be based on information gathered at asingle scale.This revised version was published online in May 2005 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.