Neutral models for the analysis of broad-scale landscape pattern

The relationship between a landscape process and observed patterns can be rigorously tested only if the expected pattern in the absence of the process is known. We used methods derived from percolation theory to construct neutral landscape models,i.e., models lacking effects due to topography, contagion, disturbance history, and related ecological processes. This paper analyzes the patterns generated by these models, and compares the results with observed landscape patterns. The analysis shows that number, size, and shape of patches changes as a function of p, the fraction of the landscape occupied by the habitat type of interest, and m, the linear dimension of the map. The adaptation of percolation theory to finite scales provides a baseline for statistical comparison with landscape data. When USGS land use data (LUDA) maps are compared to random maps produced by percolation models, significant differences in the number, size distribution, and the area/perimeter (fractal dimension) indices of patches were found. These results make it possible to define the appropriate scales at which disturbance and landscape processes interact to affect landscape patterns.     

Relationships between landcover proportion and indices of landscape spatial pattern

Recent studies have related percolation theory and critical phenomena to the spatial pattern of landscapes. We generated simulated landscapes of forest and non-forest landcover to investigate the relationship between the proportion of forest (P_i) and indices of patch spatial pattern. One set of landscapes was generated by randomly assigning each pixel independently of other pixels, and a second set was generated by randomly assigning rectilinear clumps of pixels. Indices of spatial pattern were calculated and plotted against P_i. The random-clump landscapes were also compared with real agricultural landscapes. The results support the use of percolation models as neutral models in landscape ecology, and the performance of the indices studied with these neutral models can be used to help interpret those indices calculated for real landscapes.     

Effects of changing spatial scale on the analysis of landscape pattern

The purpose of this study was to observe the effects of changing the grain (the first level of spatial resolution possible with a given data set) and extent (the total area of the study) of landscape data on observed spatial patterns and to identify some general rules for comparing measures obtained at different scales. Simple random maps, maps with contagion (i.e., clusters of the same land cover type), and actual landscape data from USGS land use (LUDA) data maps were used in the analyses. Landscape patterns were compared using indices measuring diversity (H), dominance (D) and contagion (C). Rare land cover types were lost as grain became coarser. This loss could be predicted analytically for random maps with two land cover types, and it was observed in actual landscapes as grain was increased experimentally. However, the rate of loss was influenced by the spatial pattern. Land cover types that were clumped disappeared slowly or were retained with increasing grain, whereas cover types that were dispersed were lost rapidly. The diversity index decreased linearly with increasing grain size, but dominance and contagion did not show a linear relationship. The indices D and C increased with increasing extent, but H exhibited a variable response. The indices were sensitive to the number (m) of cover types observed in the data set and the fraction of the landscape occupied by each cover type (P _k); both m and P _kvaried with grain and extent. Qualitative and quantitative changes in measurements across spatial scales will differ depending on how scale is defined. Characterizing the relationships between ecological measurements and the grain or extent of the data may make it possible to predict or correct for the loss of information with changes in spatial scale.     

Theory and language in landscape analysis,planning, and evaluation

A soft paradigm for landscape analysis is presented. This paradigm focuses on the analysis of function first, and then on structure. The objective is to determine which factors are operationally significant, how these factors bring about change, and how they define the spatial characteristics of landscapes.     

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.     

A novel use of the lacunarity index to discern landscape function

Discerning the function of a landscape involves comparing landscape use with spatial patterns. To do this requires both quantification of landscape use and landscape pattern and a means of comparing the two. An index of lacunarity has been used to quantify spatial pattern (specifically, habitat contagion). We demonstrate a new way of using the lacunarity index to quantify landscape function as well. We calculated lacunarity to describe landscape patchiness of experimental landscapes with respect to patterns of habitat and non-habitat areas (the previous use of lacunarity) as well as to describe patterns of patch use by animals in those landscapes, irrespective of habitat-patch patterns (a novel application of lacunarity). We demonstrate a disparity between landscape pattern and landscape use. This finding suggests that drawing generalizations of, and making predictions about, how animals respond to landscape spatial structure may not be straightforward.     

Transitions in forest fragmentation: implications for restoration opportunities at regional scales

Where the potential natural vegetation is continuous forest (e.g., eastern US), a region can be divided into smaller units (e.g., counties, watersheds), and a graph of the proportion of forest in the largest patch versus the proportion in anthropogenic cover can be used as an index of forest fragmentation. If forests are not fragmented beyond that converted to anthropogenic cover, there would be only one patch in the unit and its proportional size would equal 1 minus the percentage of anthropogenic cover. For a set of 130 watersheds in the mid-Atlantic region, there was a transition in forest fragmentation between 15 and 20% anthropogenic cover. The potential for mitigating fragmentation by connecting two or more disjunct forest patches was low when percent anthropogenic cover was low, highest at moderate proportions of anthropogenic cover, and again low as the proportion of anthropogenic cover increased toward 100%. This fragmentation index could be used to prioritize locations for restoration by targeting watersheds where there would be the greatest increase in the size of the largest forest patch.     

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.     

A hierarchical neutral model for landscape analysis

Empirical studies have revealed scaled structure on a variety of landscapes. Understanding processes that produce these structures requires neutral models with hierarchical structure. The present study presents a method for generating random maps possessing a variety of hierarchical structures. The properties of these scaled landscapes are analyzed and compared to patterns on totally random, unstructured landscapes. Hierar-chical structure permits percolation (i.e., continous habitat spanning the landscape) under a greater variety of conditions than found on totally random landscapes. Habitat clusters on structured maps tend to have smaller perimeters. The clusters tend to be less clumped on sparsely occupied landscapes and more clumped in densely occupied conditions. Hierarchical structure changes the expected spatial properties of the landscape, indicating a strong need for this new generation of neutral models. Research supported by the Ecological Research Division, Office to Health and Environmental Research, U.S. Department of Energy under contract No. DE-AC05-840R21400 with Martin Marietta, Energy Systems, Inc. Environmental Sciences Publication No. 3813, ORNL.     

Multiple landscape scales: An intersite comparison

Vegetation transect data from three locations were analyzed to determine if multiple scales of pattern could be detected. The sites included a semiarid grassland in New Mexico, a series of calcareous openings in a deciduous forest in Tennessee, and a shrub-steppe system in Washington. The data were explored with four statistical techniques. A scale of pattern was accepted if detected by more than one analytical method or located by a single method in multiple taxa. The analyses indicated 3–5 scales of pattern on all three sites, as predicted by Hierarchy Theory.     

Some general principles of landscape and regional ecology

A dozen general principles of landscape and regional ecology are delineated to stimulate their evaluation, refinement, and usage. Brief background material and a few references provide entrées into the subjects. The principles are presented in four groups: landscapes and regions; patches and corridors; mosaics; and applications. Most appear useful in solving a wide range of environmental and societal land-use issues.     

Structural dynamic of a hedgerow network landscape in Brittany France

Changes in agricultural systems since the 50's led to considerable changes in rural hedgerow network landscapes. In these landscapes, ecological processes depend on the spatial structure of the network (length of hedgerows, connectedness, grain size). This paper reports on a study of the dynamics of such a landscape at four periods of time (1952, 1961, 1972, 1985) done on 26 contiguous 16 ha quadrats. A correspondence analysis of the data matrix yields a gradient of change from dense highly connected networks to heterogeneous landscapes with few hedgerows. The study of individual trajectories of the quadrats allowed them to be regrouped in various types of changes. It is possible for a quadrat to go through several pathways. Rates of change varied through time, the 1961–1972 period had most changes. The use of supplementary elements in correspondence analysis proves to be a useful way to approach spatial hierarchy and allows a better understanding of the differentiation of landscape units.     

Ownership and ecosystem as sources of spatial heterogeneity in a forested landscape,Wisconsin, USA

The interaction between physical environment and land ownership in creating spatial heterogeneity was studied in largely forested landscapes of northern Wisconsin, USA. A stratified random approach was used in which 2500-ha plots representing two ownerships (National Forest and private non-industrial) were located within two regional ecosystems (extremely well-drained outwash sands and moderately well-drained moraines). Sixteen plots were established, four within each combination of ownership and ecosystem, and the land cover on the plots was classified from aerial photographs using a modified form of the Anderson (U.S. Geological Survey) land use and land cover classification system.Upland deciduous forests dominated by northern hardwoods were common on the moraines for both ownerships. On the outwash, the National Forest was dominated by pine plantations, upland deciduous forests, and upland regenerating forests (as defined by <50% canopy coverage). In contrast, a more even distribution among the classes of upland forest existed on private land/outwash. A strong interaction between ecosystem and ownership was evident for most comparisons of landscape structure. On the moraine, the National Forest ownership had a finer grain pattern with more complex patch shapes compared to private land. On the outwash, in contrast, the National Forest had a coarser grain pattern with less complex patch shapes compared to private land. When patch size and shape were compared between ecosystems within an ownership, statistically significant differences in landscape structure existed on public land but not on private land. On public land, different management practices on the moraine and outwash, primarily related to timber harvesting and road building, created very different landscape patterns. Landscape structure on different ecosystems on private land tended to be similar because ownership was fragmented in both ecosystems and because ownership boundaries often corresponded to patch boundaries on private land. A complex relationship exits between ownership, and related differences in land use, and the physical environment that ultimately constrains land use. Studies that do not consider these interactions may misinterpret the importance of either variable in explaining variation in landscape patterns.     

Avian response to landscape pattern: The role of species' life histories

We suggest that the life histories of species within communities may differ among geographic locations and that communities from distinct biomes may respond uniquely to a given trajectory of landscape change. This paper presents initial tests relevant to these hypotheses. First, the representation of various life-history guilds in avifaunas from the Eastern Deciduous (EDF) and Pacific Northwest (PNW) forests were compared. Three guilds contained more species in the EDF community (large patch and/or habitat interior guild, small patch and/or edge guild, and fragmentation-sensitive guild). The guild of predators requiring large forest tracts was better represented in the PNW. Next, the relative sensitivity of each community to habitat change was ranked based on the life-history traits of their species. The EDF avifauna had a significantly higher index of sensitivity to both forest fragmentation and to landscape change in general. Among the birds with high scores for sensitivity to landscape change were several species that have received little conservation attention thus far including some associated with open-canopy habitats. Lastly, the validity of using life histories to predict community response to landscape change was supported by the fact that the sensitivity scores for PNW species correlated significantly with independent data on species population trends. While more rigorous analyses are suggested, we conclude that knowledge of life histories is useful for predicting community response to landscape change and that conservation strategies should be uniquely tailored to local communities.     

The effect of spatial scale on Konza landscape classification using textural analysis

Spatial scale is inherent in the definition of landscape heterogeneity and diversity. For example, a landscape may appear heterogeneous at one scale but quite homogeneous at another scale. In assessing the impact of burning and grazing on the Konza Prairie Research Natural Area (a tallgrass prairie), spatial scale is extremely important. Textural contrast algorithms were applied to various scales of remote sensing data and related to landscape units for assessment of heterogeneity under a variety of burning treatments. Acquired data sets included Landsat multispectral scanner (MSS), with 80 m resolution, Landsat thematic mapper (TM), with 30 m resolution, and high resolution density sliced aerial photography (with a 5 m resolution). Results suggest that heterogeneous areas of dense patchiness (e.g., unburned areas) must be analyzed at a finer scale than more homogeneous areas which are burned at least every four years.     

Mapping and analyzing landscape patterns

Landscapes were mapped as clusters of 2 or 3 land cover^** types, based on their pattern within the clusters and tendency for a single type to dominate. These landscapes, called Landscape Pattern Types (LPTs), were combined with other earth surface feature data in a Geographic Information System (GIS) to test their utility as analysis units. Road segment density increased significantly as residential and urbanized land cover components increased from absent, to present as patch, to present as matrix (i.e., the dominant land cover type). Stream segment density was significantly lower in LPTs with an urbanized or residential matrix than in LPTs with either a forest or agriculture matrix, suggesting an inverse relationship between stream network density and the prevalence of human development other than agriculture in the landscape. The ratio of average forest patch size to total forest in the LPT unit decreased as agriculture replaced forest, then increased as residential and urban components dominated. Wetland fractal dimension increased as agriculture and residential land cover components of LPTs increased. Comparison of LPT and LUDA land cover area statistics in ecoregions suggested that land cover data alone does not provide information as to its spatial arrangement.     

Agricultural soil redistribution and landscape complexity

A number of hypotheses and conceptual models, particularly those emphasizing nonlinear dynamics and self-organization, postulate increases or decreases in complexity in the evolution of drainage basins, topography, soils, ecosystems, and other earth surface systems. Accordingly, it is important to determine under what circumstances and at what scales either trend might occur. This paper is concerned with changes in soil landscape complexity due to redistribution of sediment by fluvial, aeolian, and tillage processes at historical time scales in an agricultural field system near Grifton, North Carolina. Soil mapping and soil stratigraphic investigations were used to identify and map soil changes associated with erosion and deposition by water, wind, and tillage; reconstruct the pre-agricultural soil pattern; and identify transformations between soil types. The Kolmogorov entropy of the pre- and post- agricultural landscapes was then compared. The soil transformations associated with erosion and deposition created four distinct new soils and made possible new transformations among soil series, increasing the number of soil types from seven to 11 and the number of possible transformations from 14 to 22. However, the entropy and complexity of the soil landscape decreased, with associated increases in information and redundancy. The mass redistributions created a lower-entropy landscape by concentrating particular soils and soil transformations in specific landscape settings. This result is contrary to studies showing a trend toward increasing pedological complexity at comparable spatial scales, but over much longer time scales. These results point to the importance of temporal scale, and to the fact that environmental complexity is influenced by factors other than the number of different landscape units present.     

A new contagion index to quantify spatial patterns of landscapes

A contagion index was proposed by O'Neill et al. (1988) to quantify spatial patterns of landscapes. However, this index is insensitive to changes in spatial pattern. We present a new contagion index that corrects an error in the mathematical formulation of the original contagion index. The error is identified mathematically. The contagion indices (both original and new) are then evaluated against simulated landscapes.