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.     

Effects of changing spatial scale on acoustic observations of patchiness in the Gulf Stream

We examine the influence of spatial scale on studies of nektonic patchiness at the north edge of the Gulf Stream by altering the grain size of acoustic cross sections and applying a patch-finding algorithm. From original pictures of 180 pixels deep by 540–1260 pixels long, we averaged depth and length, to give 9 scales ranging from fine grain (1 m vertical × 25 m horizontal sized pixels) to coarse grain (12 m × 300 m sized pixels). Measures of overall echo intensity within patches were the most predictable and showed little change with scale. Measures of variability of the echo within patches showed simple negative trends with scale and were best measured at fine spatial scales. Patch size and shape related variables have a more complex relationship with scale with differences between day and night transects more pronounced at intermediate scales. This suggests behavioral patch forming mechanisms within the nekton occur at a specific grain size (4 m vertical × 100 m horizontal) within the Gulf Stream front.     

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.     

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.     

Foraging in a fractal environment: Spatial patterns in a marine predator-prey system

Spatial relationships between predators and prey have important implications for landscape processes and patterns. Highly mobile oceanic birds and their patchily distributed prey constitute an accessible model system for studying these relationships. High-frequency echosounders can be used together with simultaneous direct visual observations to quantitatively describe the distributions of seabird consumers and their resources over a wide range of spatial scales, yielding information which is rarely available in terrestrial systems.Recent fine-scale investigations which have used acoustics to study the distribution of foraging marine birds have reported weak or ephemeral spatial associations between the birds and their prey. These results are inconsistent with predictions of optimal foraging, but several considerations suggest that traditional foraging models do not adequately describe resource acquisition in marine environments. Relative to their terrestrial counterparts, oceanic landscapes are structurally very simple, but they generally lack visual cues about resource availability.An emerging view assumes that perceptually constrained organisms searching for food in multiscale environments should respond to patterns of resource abundance over a continuum of scales. We explore fractal geometry as a possible tool for quantifying this view and for describing spatial dispersion patterns that result from foraging behavior. Data on an Alaskan seabird (least auklet [Aethia pusilla]) and its zooplanktonic food resources suggest that fractal approaches can yield new ecological insights into complex spatial patterns deriving from animal movements.     

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.     

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.     

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.     

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.     

Pseudoreplication: a sine qua non for regional ecology

We question whether classical experimentation is adequate for real progress in landscape or regional ecology. One cannot do classical experimentation unless one can replicate the treatment. There is conflict between the need to replicate and the need to study processes at appropriately large scales.Because of the difficulties in doing controlled field experiments at regional scales, we propose that landscape ecologists take greater advantage of natural field experiments. Natural experiments must be coordinated, standardized, and synchronized over space and through time, and will require the cooperation of multiple investigators. Distributed computer networks can help provide the automated region-wide monitoring which will supply natural experiments with pre-treatment data.Regions or landscapes need not be replicated, and indeed, cannot be. One can achieve a relational understanding between a system's response and environmental characteristics. This understanding is not definitive, but allows for the development of testable hypotheses, in the classical sense. The confounding of space, time, and/or other environmental factors in pseudoreplicated natural experiments only allows for the development of hypotheses - how-possibly explanations. Discrimination among competing hypotheses can be done at smaller scales and used to infer processes occurring at larger scales. Use of natural and controlled field experiments in complementary roles is a more promising approach than views of one or the other as methodologically inferior.     

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.     

Modeling multiscale effects of light limitations and edge-induced mortality on carbon stores in forest landscapes

Analyses of carbon (C) dynamics at broad scales usually do not consider spatial interactions. The assumption is that C dynamics can be modeled within homogenous (i.e., even-aged) patches and then summed to predict broad-scale dynamics (an additive approach). The goal of this paper is to elucidate the scales over which this additive approach is sufficient to explain observed C dynamics at broad scales. We define emergent behaviors (vs. emergent properties) as those behaviors that cannot be predicted solely from the additive properties of units at a finer scale. We used a forest process model to check for possible emergent behaviors due to pattern-process interactions at multiple levels, from the patch to the landscape. Specifically, using artificial forest landscapes with various spatial structures, we estimated the relative effects of edge-induced, tree mortality (mainly due to wind) and light limitations on C dynamics. Emergent behaviors were observed at all levels examined, indicating that emergent behaviors did not cease as one proceeded from the patch to the landscape level, as we had expected. However, the magnitude of the emergent behaviors depended on the level of spatial interaction considered as well as the type and intensity of the processes included. In all simulations, interactions of light and wind processes resulted in significant emergent behaviors only when parameters controlling wind mortality were set to the highest levels observed in the literature. In one simulation, the magnitude of emergent behaviors differed among the landscapes, indicating that interactions among patches may not be accounted for by an additive correction for edge effects unless spatial interactions are addressed. The implication is that some C dynamics in fragmented landscapes may not be captured at broad-scales using an additive approach, whereas in other cases spatial interactions are small enough to be ignored.This revised version was published online in May 2005 with corrections to the Cover Date.     

Land-use changes in Illinois,ASA: The influence of landscape attributes on current and historic land use

The Illinois Geographic Information System was used to compare the soil and landscape attributes of the State with its historic vegetation, current land use, and patterns of land-use change over the past 160 years. Patch structural characteristics among land types in four geographic zones were also compared. The assessment of patch characteristics revealed a highly modified State with most land patches controlled by human influences and relatively few by topographic and hydrologic features. Correlation and regression analyses determined the relationships of land type and abundance within each of 50 general soil associations to properties of the soil associations - typically slope, texture, organic matter, productivity index, and available waterholding capacity. The distribution of the historic vegetation of the State and its current deciduous forests and nonforested wetlands related moderately (r² 0.44) to various landscape attributes. Urban and other highly modified land types were less closely related.     

Resource depletion versus landscape complementation: habitat selection by a multiple central place forager

Context

The spatial distribution of non-substitutable resources implies diverging predictions for animal movement patterns. At broad scales, animals should respond to landscape complementation by selecting areas where resource patches are close-by to minimize movement costs. Yet at fine scales, central place effects lead to the depletion of patches that are close to one another and that should ultimately be avoided by consumers.

Objectives

We developed a multi-scale resource selection framework to test whether animal movement is driven by landscape complementation or resource depletion and identify at which spatial scale these processes are relevant from an animal’s perspective.

Methods

During the dry season, surface water and forage are non-substitutable resources for African elephants. Eight family herds were tracked using GPS loggers in Hwange National Park, Zimbabwe. We explained habitat selection during foraging trips by mapping surface water at two scales with gaussian kernels of varying widths placed over each waterhole.

Results

Unexpectedly, elephants select areas with low waterhole density at both fine scales (< 1 km) and broad scales (5–7 km). Selection is stronger when elephants forage far away from water, even more so as the dry season progresses.

Conclusions

Elephant selection of low waterhole density areas suggests that resource depletion around multiple central places is the main driver of their habitat selection. By identifying the scale at which animals respond to waterhole distribution we provide a template for water management in arid and semi-arid landscapes that can be tailored to match the requirements and mobility of free ranging wild or domestic species.

     

Modeling the effects of forest harvesting on landscape structure and the spatial distribution of cowbird brood parasitism

Timber harvesting affects both composition and structure of the landscape and has important consequences for organisms using forest habitats. A timber harvest allocation model was constructed that allows the input of specific rules to allocate forest stands for clearcutting to generate landscape patterns reflecting the look and feel of managed landscapes. Various harvest strategies were simulated on four 237 km² study areas in Indiana, USA. For each study area, the model was applied to simulate 80 years of management activity. The resulting landscape spatial patterns were quantified using a suite of landscape pattern metrics and plotted as a function of mean harvest size and total area of forest harvested per decade to produce response surfaces. When the mean clearcut size was 1 ha, the area of forest interior remaining on the landscape was dramatically reduced and the amount of forest edge on the landscape increased dramatically. The potential consequences of the patterns produced by the model were assessed for a generalized neotropical migrant forest bird using a GIS model that generates maps showing the spatial distribution of the relative vulnerability of forest birds to brood parasitism by brown-headed cowbirds. The model incorporates the location and relative quality of cowbird feeding sites, and the relation between parasitism rates and distance of forest from edge. The response surface relating mean harvest size and total area harvested to the mean value of vulnerability to cowbird brood parasitism had a shape similar to the response surfaces showing forest edge. The results of our study suggest that it is more difficult to maintain large contiguous blocks of undisturbed forest interior when harvests are small and dispersed, especially when producing high timber volumes is a management goal. The application of the cowbird model to landscapes managed under different strategies could help managers in deciding where harvest activity will produce the least negative impact on breeding forest birds.     

Scale detection in real and artificial landscapes using semivariance analysis

Semivariance analysis is potentially useful to landscape ecologists for detecting scales of variability in spatial data. We used semivariance analysis to compare spatial patterns of winter foraging by large ungulates with those of environmental variables that influence forage availability in northern Yellowstone National Park, Wyoming. In addition, we evaluated (1) the ability of semivariograms to detect known scales of variability in artificial maps with one or more distinct scales of pattern, and (2) the influence of the amount and spatial distribution of absent data on semivariogram results and interpretation. Semivariograms of environmental data sets (aspect, elevation, habitat type, and slope) for the entire northern Yellowstone landscape clearly identified the dominant scale of variability in each map layer, while semivariograms of ungulate foraging data from discontinuous study areas were difficult to interpret. Semivariograms of binary maps composed of a single scale of pattern showed clear and interpretable results: the range accurately reflected the size of the blocks of which the maps were constructed. Semivariograms of multiple scale maps and hierarchical maps exhibited pronounced inflections which could be used to distinguish two or three distinct scales of pattern. To assess the sensitivity of semivariance analysis to absent data, often the product of cloud interference or incomplete data collection, we deliberately masked (deleted) portions of continuous northern Yellowstone map layers, using single scale artificial maps as masks. The sensitivity of semivariance analysis to random deletions from the data was related to both the size of the deleted blocks, and the total proportion of the original data set that was removed. Small blocks could be deleted in very high proportions without degrading the semivariogram results. When the size of deleted blocks was large relative to the size of the map, the corresponding variograms became sensitive to the total proportion of data removed: variograms were difficult or impossible to interpret when the proportion of data deleted was high. Despite success with artificial maps, standard semivariance analysis is unlikely to detect multiple scales of pattern in real ecological data. Semivariance analysis is recommended as an effective technique for quantifying some spatial characteristics of ecological data, and may provide insight into the scales of processes that structure landscapes.     

Streambed landscapes: evidence that stream invertebrates respond to the type and spatial arrangement of patches

The availability and spatial arrangement of habitat patches are known to strongly influence fauna in terrestrial ecosystems. The importance of patch arrangement is not well-studied within running-water systems where flow-induced movements of patches and of fauna could decouple habitat characteristics and faunal habitat preferences. Using small, stream-dwelling invertebrates, we asked if fauna in such systems can distinguish among patch types and if patch arrangement at their `landscape scale' (i.e., within a streambed across which they move and forage) can be linked to faunal abundance. We quantified the spatial distribution of sand and leaf patches at multiple sites on a streambed at regular intervals over a 1 yr period, estimated faunal abundance in the two patch types, and experimentally determined if faunal colonization varied among leaf patches that were similar structurally but differed in their potential microbial food resources. We show that despite their small size and limited swimming abilities, these stream invertebrates did respond to patch type, that specific characteristics of an individual patch influenced faunal colonization, and that the spatial arrangement of patches on the streambed was linked to field abundances. Larval chironomids and adult copepods were more abundant in leaves than in sand and preferentially colonized leaf patches made with rapidly decomposing leaves that harbored higher microbial (bacteria and fungi) abundances over leaf patches with more refractory leaves and lower microbial abundances. Further, statistical models that included spatially-explicit data on patch arrangement (e.g., patch contagion, distance between patches) explained significantly more variation in faunal abundance, than models that included only nonspatial information (e.g., date, time since last flood). Despite the fact that these fauna live in a highly dynamic environment with variable flow rates during the year, unstable patch configurations, and seasonal changes in total abundance, our findings suggest a need for aquatic ecologists to test the hypothesis that small-scale landscape attributes within streams (e.g., leaf patch aggregation) may be important to faunal dynamics. If patch aggregation has negative consequences for stream biota, streambed `landscapes' may be fundamentally different from many terrestrial landscapes due to the inherent connectivity provided by the water and the over-riding importance of patch edges. Regardless of these differences, our findings suggest that the spatial configuration of patches in a landscape may have consequences for fauna even in highly dynamic systems, in which patches move and fauna periodically experience high levels of passive dispersal.     

Prehistoric,historic, and present settlement patterns related to ecological hierarchy in the Eastern Upper Peninsula of Michigan,U.S.A.

The distribution of human occupation across a landscape provides informationabout how people use the landscape, about patterns of economic development,and about social interactions of human groups. When the distributions areexamined over several thousand years, we gain an evolutionary understanding,not only of the people and their cultural patterns, but also of physicallandscape development. The focus of this assessment was to examine andcompare settlement patterns of prehistoric, historic, and present timeperiods, based on known cultural sites in the Eastern Upper Peninsula ofMichigan, U.S.A., and to generate hypotheses about the interaction ofsettlement pattern and landscape change at multiple scales. Patterns ofsettlement among the three time periods were compared at three geographicscales: by subregional ecosystems, by landscape ecosystems and by terraincharacteristics. The Michigan Bureau of History database of archaeologicalsites was searched for prehistoric habitation sites of Middle or LateWoodland period (ca. 3000-300 years before present). Historic occupationswere drawn from pre-European settlement landscape data based on General LandOffice survey notes of the 1850s. We extracted urban categories from landcover classified from Landsat Thematic Mapper imagery to measure presentoccupations. Spatial patterns and dynamics of settlement areas in each timeperiod were examined using the ARC/INFO geographic information system (GIS).Results showed a tendency for settlement in all time periods on the bedrockand lowland landscape groups near Great Lakes shorelines, generally occupyingslopes less than two percent. The distribution of present occupations, interms of both slope aspect and geographic subregion (multi-scalar), wassimilar to the distribution of prehistoric occupations. Both prehistoric andpresent sites were primarily south facing and were frequently found alongGreen Bay and Lake Michigan shorelines.     

Quaternary landscape ecology: Relevant scales in space and time

Two primary goals of landscape ecologists are to (1) evaluate changes in ecological pattern and process on natural landscapes through time and (2) determine the ecological consequences of transforming natural land-scapes to cultural ones. Paleoecological techniques can be used to reconstruct past landscapes and their changes through time; use of paleoecological methods of investigation in combination with geomorphic and paleoethnobiological data, historical records, and shorter-term ecological data sets makes it possible to integrate long-term ecological pattern and process on a nested series of temporal and spatial scales. Natural experiments of the past can be used to test alternative hypotheses about the relative influences of environmental change, biological interactions, and human activities in structuring biotic communities within landscape mosaics.On the absolute time scale of the Quaternary Period, spanning the past 1.8 million years, current distributional ranges of the biota have taken shape and modern biotic communities have assembled. Quaternary environmental changes have influenced the development of natural landscapes over time scales of centuries to hundreds of thousands of years; human cultural evolution has resulted in the transformation of much of the biosphere from natural to cultural landscapes over the past 5,000 years. The Quaternary extends to and includes the present and the immediate future. Knowledge of landscape changes on a Quaternary time scale is essential to landscape ecologists who wish to have a context for predicting future trends on local, regional, and global scales.