Dispersal of annual plants in hierarchically structured landscapes

The scale at which plants utilize spatially distributed resources may be determined by their ability to locate sites that can sustain population growth. We developed a spatially-explicit model of the dispersal of annual plants in landscapes which were hierarchically structured, i.e., the spatial pattern of suitable sites was nested and scale-dependent. Results show that colonizing ability and extinction probability are most sensitive to the mean dispersal distance of the species. Dispersal out of the parental site, but within the immediate neighborhood, was the most efficient means for population expansion. When landscape patterns change with scale then dispersal distances determine the spatial scales of habitat utilization. As a complicating factor, the type of statistical distribution of dispersal distances also influences the colonizing ability. However, the importance of dispersal distance mean and distribution decreased as the number and connectance of suitable sites increased. The results suggest that landscape models which consider the interaction between scale dependent changes in landscape pattern and species dispersal and establishment characteristics are relevant to many issues in community ecology, invasion biology, and conservation biology.     

Epidemiology theory and disturbance spread on landscapes

Abtract Epidemiology models, modified to include landscape pattern, are used to examine the relative importance of landscape geometry and disturbance dynamics in determining the spatial extent of a disturbance, such as a fire. The models indicate that, except for very small values for the probability of spread, a disturbance tends to propagate to all susceptible sites that can be reached. Therefore, spatial pattern, rather than disturbance dynamic, will ordinarily determine the total extent of a single disturbance event. The models also indicate that a single disturbance will seldom become endemic,i.e., always present on the landscape. However, increasing disturbance frequency can lead to a landscape in which the proportion of susceptible, disturbed, and recovering sites are relatively constant. Research supported by Ecological Research Division, Office of Health and Environmental Research, U.S. Department of Energy under contract No. DE-AC05-840R21400 with Martin Marietta Energy Systems, Inc. Contribution No. 10 to the Sevilleta LTER program. Environmental Sciences Division Publication No. 3812, ORNL.     

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.     

Characterizing watershed-delineated landscapes in Pennsylvania using conditional entropy profiles

When the objective is to characterize landscapes with respect to relative degree and type of forest (or other critical habitat) fragmentation, it is difficult to decide which variables to measure and what type of discriminatory analysis to apply. It is also desirable to incorporate multiple measurement scales. In response, a new method has been developed that responds to changes in both the marginal and spatial distributions of land cover in a raster map. Multiscale features of the map are captured in a sequence of successively coarsened resolutions based on the random filter for degrading raster map resolutions. Basically, the entropy of spatial pattern associated with a particular pixel resolution is calculated, conditional on the pattern of the next coarser parent resolution. When the entropy is plotted as a function of changing resolution, we obtain a simple two-dimensional graph called a conditional entropy profile, thus providing a graphical visualization of multi-scale fragmentation patterns.Using eight-category raster maps derived from 30-meter resolution LANDSAT Thematic Mapper images, the conditional entropy profile was obtained for each of 102 watersheds covering the state of Pennsylvania (USA). A suite of more conventional single-resolution landscape measurements was also obtained for each watershed using the FRAGSTATS program. After dividing the watersheds into three major physiographic provinces, cluster analysis was performed within each province using various combinations of the FRAGSTATS variables, land cover proportions and variables describing the conditional entropy profiles. Measurements of both spatial pattern and marginal land cover proportions were necessary to clearly discriminate the watersheds into distinct clusters for most of the state; however, the Piedmont province essentially only required the land cover proportions. In addition to land cover proportions, only the variables describing a conditional entropy profile appeared to be necessary for the Ridge and Valley province, whereas only the FRAGSTATS variables appeared to be necessary for the Appalachian Plateaus province. Meanwhile, the graphical representation of conditional entropy profiles provided a visualization of multi-scale fragmentation that was quite sensitive to changing pattern.     

Forest gradient response in Sierran landscapes: the physical template

Vegetation pattern on landscapes is the manifestation of physical gradients, biotic response to these gradients, and disturbances. Here we focus on the physical template as it governs the distribution of mixed-conifer forests in California's Sierra Nevada. We extended a forest simulation model to examine montane environmental gradients, emphasizing factors affecting the water balance in these summer-dry landscapes. The model simulates the soil moisture regime in terms of the interaction of water supply and demand: supply depends on precipitation and water storage, while evapotranspirational demand varies with solar radiation and temperature. The forest cover itself can affect the water balance via canopy interception and evapotranspiration. We simulated Sierran forests as slope facets, defined as gridded stands of homogeneous topographic exposure, and verified simulated gradient response against sample quadrats distributed across Sequoia National Park. We then performed a modified sensitivity analysis of abiotic factors governing the physical gradient. Importantly, the model's sensitivity to temperature, precipitation, and soil depth varies considerably over the physical template, particularly relative to elevation. The physical drivers of the water balance have characteristic spatial scales that differ by orders of magnitude. Across large spatial extents, temperature and precipitation as defined by elevation primarily govern the location of the mixed conifer zone. If the analysis is constrained to elevations within the mixed-conifer zone, local topography comes into play as it influences drainage. Soil depth varies considerably at all measured scales, and is especially dominant at fine (within-stand) scales. Physical site variables can influence soil moisture deficit either by affecting water supply or water demand; these effects have qualitatively different implications for forest response. These results have clear implications about purely inferential approaches to gradient analysis, and bear strongly on our ability to use correlative approaches in assessing the potential responses of montane forests to anthropogenic climatic change.     

Temporal patterns of ecosystem processes on simulated landscapes in Glacier National Park,Montana, USA

The mechanistic, spatially-explicit fire succession model, Fire-BGC (a Fire BioGeoChemical succession model) was used to investigate long-term trends in landscape pattern under historical and future fire regimes and present and future climate regimes for two 46000 ha landscapes in Glacier National Park, Montana, USA. Fire-BGC has two spatial and temporal resolutions in the simulation architecture where ecological processes that act at a landscape level, such as fire, are simulated annually from information contained in spatial data layers, while stand-level processes such as photosynthesis, transpiration, and decomposition are simulated both daily and annually. Fire is spread across the landscape using the FARSITE fire growth model and subsequent fire effects are simulated at the stand-level. Fire-BGC was used to simulate changes in landscape pattern over 250 years under four scenarios: (1) complete fire exclusion under current climate, (2) historical wildfire occurrence and current climate, (3) complete fire exclusion under a possible future climate, (4) future wildfire occurrence and future climate. Simulated maps of dominant tree species, aboveground standing crop, leaf area index, and net primary productivity (NPP) were contrasted across scenarios using the metrics of patch density, edge density, evenness, contagion, and interspersion. Simulation results indicate that fire influences landscape pattern metrics more that climate alone by creating more diverse, fragmented, and disconnected landscapes. Fires were more frequent, larger, and more intense under a future climate regime. Landscape metrics showed different trends for the process-based NPP map when compared to the cover type map. It may be important to augment landscape analyses with process-based layers as well as structural and compositional layers.     

Are similar landscapes the result of similar histories?

This landscape study was based on the sampling of 20 replicated landscape sites (1 km² each) that were located within the floodplain of the river Seine. For each site, 13 landscape variables were measured at three dates (1963–1985–2000). The aim of this study was to investigate the overall landscape variability through its different dimensions (space vs. time) and to assess the relative importance of each dimension. We used a new statistical method, i.e., partial triadic analysis (PTA), which allowed us to assess both (1) the spatial variability of the floodplain landscape and its dynamics in time and (2) the dynamic trajectories of the landscape variables for each site. The results showed, at the floodplain scale, the same landscape pattern has emerged since 1963, although a major trend was observed which consisted in a decrease in meadows resulting from an increase in arable crops. At the site scale, landscape sites, even if they were all influenced by this general trend during the 40-year period, showed contrasting trajectories. These results suggest that similar sites in 2000 do not necessarily share common histories and that contrasting sites in 2000 may have originated from similar patterns in 1963. The issue of biodiversity surrogates is then discussed, suggesting that new landscape metrics should be developed, emphasising spatial variability and (or) temporal dynamics.     

A polygon-based spatial (PBS) model for simulating landscape change

A spatial model of long term habitat succession at a degrading Louisiana wetland was constructed based upon simulating exchanges across irregularly shaped polygons. Polygons represented the natural morphology which is indicative of the natural landscape. The PBS model was partially successful in simulating spatial habitat changes over a 28-year period when more than 1000 ha of wetland loss occurred (r²=.56). General landscape trends did, however, emerge from the model development. Areas of high annual water level fluctuations, and high primary productivity were less likely to change from wetlands to open water and were most likely to recover if altered. We discuss the potential for predictive improvement and for integration with polygon-based geographic information systems, and conclude that a PBS model demonstrates the need for spatially explicit landscape management.     

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.     

Barriers as boundaries for metapopulations and demes of Peromyscus leucopus in farm landscapes

Effects of potential barriers (roads and cultivated fields) on both demographic and genetic features of subpopulations of white-footed mice were studied near Ottawa, Canada. Live trapping, colored bait and track registry were used to study animal movements across roads on four 1.44 ha areas each within a small forest bisected by a narrow gravel road. The genetic study was done in 11 other forest fragments separated from each other by cultivated fields. Frequencies of three electrophoretic variants of salivary amylases were established for mice caught in each patch of wood and genetic similarity of subpopulations was calculated. Movements of mice across the roads were very infrequent (quantitative barrier), although movements adjacent to roads were frequent and long enough to cross the roads. Salivary amylase data showed that studied subpopulations were genetically very similar although the sample was intentionally biased toward demographic isolation. Results are discussed in terms of possible hierarchical relationships of metapopulations and genetic demes in the context of landscape ecology, management and conservation practice.     

An aggregation index (AI) to quantify spatial patterns of landscapes

There is often need to measure aggregation levels of spatial patterns within a single map class in landscape ecological studies. The contagion index (CI), shape index (SI), and probability of adjacency of the same class (Qi), all have certain limits when measuring aggregation of spatial patterns. We have developed an aggregation index (AI) that is class specific and independent of landscape composition. AI assumes that a class with the highest level of aggregation (AI =1) is comprised of pixels sharing the most possible edges. A class whose pixels share no edges (completely disaggregated) has the lowest level of aggregation (AI =0). AI is similar to SI and Qi, but it calculates aggregation more precisely than the latter two. We have evaluated the performance of AI under varied levels of (1) aggregation, (2) number of patches, (3) spatial resolutions, and (4) real species distribution maps at various spatial scales. AI was able to produce reasonable results under all these circumstances. Since it is class specific, it is more precise than CI, which measures overall landscape aggregation. Thus, AI provides a quantitative basis to correlate the spatial pattern of a class with a specific process. Since AI is a ratio variable, map units do not affect the calculation. It can be compared between classes from the same or different landscapes, or even the same classes from the same landscape under different resolutions.     

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.     

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.     

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.     

Rates and patterns of landscape change in the Central Sikhote-alin Mountains,Russian Far East

We used Landsat imagery and GIS to quantify the rates and patterns of landscape change between 1972 and 1992 for a 734,126 ha forested study area in the central Sikhote-alin Mountains of the Russian Far East. The study area includes a portion of the Sikhote-alinskiy Biosphere Reserve which is a part of the United Nations international Man and the Biosphere (MAB) reserve network. Wildfire is a major disturbance agent throughout the area and timber harvesting outside the reserve is also important. Maximum likelihood classification of the satellite imagery identified four broad cover types (hardwood, conifer, mixed and non-forest) in 1992 and changes among them between 1972 and 1992. We used multi-temporal principal components analysis to describe the magnitude and direction of landscape change for six watersheds that represent a range of ecological histories and disturbance regimes. Overall, forest cover declined from 90.4% in 1972 to 77.2% in 1992. The disturbance rate was more than twice as high in conifer than in hardwood forests. The rate of disturbance outside the reserve was three times that inside. While the rates of disturbance are not markedly higher than those recorded from other temperate forests, there has recently been a large alteration in the disturbance regime which will lead to a general transformation of forest composition and structure in the study area if the trend continues.     

Determination of ecological scale

We suggest that ecological processes and physical characteristics possess an inherent scale at which the processes or characteristics occur over the landscape. We propose a conceptual spatial response model that describes the nature of this ecological scale. Based on the proposed spatial model, we suggest methods for estimating the size of study plots or transects and the distance between replicate plots needed to approach statistical independence. Using data on percent cover for Agropyron spicatum, a common arid-land bunchgrass, we demonstrated four relationships that should hold if the spatial response model is appropriate. These relationships are sample variance increases as functions of (1) transect segment length and (2) intersegment length (transect segment dispersal), and correlation decreases as functions of (3) intersegment length and (4) transect segment length. Based on evaluation of these four relationships, cover for A. spicatum is correlated over the landscape on a scale of 400 to 700 m, and a segment length of 64 to 128 m is most appropriate for measuring cover for this species.     

A framework for environmental management planning — A landscape-ecological approach

Framework, concepts, and methods of Environmental Management Planning (EMP) are discussed. A landscape-ecological approach was taken to integrate the environmental indices. EMP focuses on regional factors- natural, social, amenity related — and becomes more sensitive as the scale of study increases. The processes of EMP include a vertical aspect, dealing with pollution, conservation, and amenities, and a more general horizontal component which involves zoning and land use planning. Environmental impacts may be assessed by modeling exercises using all available data and considering all land use options. To keep up with the rapid change of environment and its perception, EMP should be process-oriented rather than purpose-oriented. The concepts of EMP were applied to the middle basin of the Tamagawa River and it was shown that mulvariate analysis is useful for the regional subdivision and environmental modeling.     

Modeling the optimal ecological security pattern for guiding the urban constructed land expansions

Rapid urbanization has induced numerous ecological and environmental issues seriously threatening the ecological security. The ecological security pattern (ESP), an effective way for protecting the ecological security, is becoming increasingly important in reconciling the rapid urbanization and ecology protection in urban planning practices. Based on the cost-distance analysis method, we constructed a three-rank (basic, moderate and strict-rank) composite ESP of Gaoming (Guangdong, China) aiming at protecting the survivals and habitat securities of rare vegetations, wild animals and human beings. The proposed composite ESP is established on five equal-weighted individual ESPs (namely Geology-ESP, Hydrology-ESP, Atmosphere-ESP, Biodiversity-ESP and Farmland-ESP) for geologic disasters prevention, flood prevention and drinking water protection, air pollution prevention, biodiversity conservation and farmland protection, respectively. Our results show that under the basic, moderate, and strict- rank ESPs, the integration and connectivity of the ecological components are constantly improved, but the connectivity between neighboring urban patches decline gradually. The moderate-rank ESP proves to be the optimal spatial pattern for balancing the conflicts between urban development and ecological protection. Notably, the ESP that considers the security of atmosphere and farmland securities, which protects the regional farmlands better and well balance the expansions of industrial and residual lands, proves to be much more reasonable.     

Changes in land-use/land-cover patterns in Italy and their implications for biodiversity conservation

Land-use/land-cover change is the most important factor in causing biodiversity loss. The Mediterranean region has been affected by antropic disturbance for thousands of years, and is, nowadays, one of the most significantly altered hotspots in the world. However, in the last years a significant increase in forest cover has been measured. These new patterns are independent from planned conservation strategies and appear to have a substantial impact on landscapes and biodiversity. We used three land-use/land-cover maps (from 1960 to 2000) covering the Italian peninsula to analyze the pattern of land-use/land-cover change. We measured an increase in forests, especially in mountains, an increase in artificial areas, especially in coastal zones, and a decrease in pastures. Intensively cultivated areas showed a limited decrease while extensively cultivated ones showed a marked decrease. In the same period mammal and bird species followed a similar pattern, with forest birds, ungulates and carnivores increasing, and typically Mediterranean species decreasing. We suggest that our results may provide important information, which could be useful for conservation planning in the entire Mediterranean hotspot. We suggest that an increasing conservation effort should be made to protect the Mediterranean-type forests and scrublands, as well as traditional agricultural practices. Moreover, future conservation efforts should consider the broad socio-political and ecological processes that are most likely to occur across the whole hotspot, especially along coastal areas, and the network of protected areas should be functionally integrated in a conservation strategy that includes the human-dominated landscape.