The Importance of Spatial Scale in Habitat Models: Capercaillie in the Swiss Alps

The role of scale in ecology is widely recognized as being of vital importance for understanding ecological patterns and processes. The capercaillie (Tetrao urogallus) is a forest grouse species with large spatial requirements and highly specialized habitat preferences. Habitat models at the forest stand scale can only partly explain capercaillie occurrence, and some studies at the landscape scale have emphasized the role of large-scale effects. We hypothesized that both the ability of single variables and multivariate models to explain capercaillie occurrence would vary with the spatial scale of the analysis. To test this hypothesis, we varied the grain size of our analysis from 1 to just over 1100 hectares and built univariate and multivariate habitat suitability models for capercaillie in the Swiss Alps. The variance explained by the univariate models was found to vary among the predictors and with spatial scale. Within the multivariate models, the best single-scale model (using all predictor variables at the same scale) worked at a scale equivalent to a small annual home range. The multi-scale model, in which each predictor variable was entered at the scale at which it had performed best in the univariate model, did slightly better than the best single-scale model. Our results confirm that habitat variables should be included at different spatial scales when species-habitat relationships are investigated.     

Response of snails and slugs to fragmentation of lowland forests in NW Germany

Habitat fragmentation is a major cause for species loss, but its effect on invertebrates with low active dispersal power, like terrestrial gastropods, has rarely been studied. Such species can not cross a hostile habitat matrix, for which the predictions of island theory, such as positive relations between species richness and patch size, should apply. In order to test this prediction, we studied gastropod species diversity by assessing gastropod assemblage characteristics from 35 sites in 19 fragments of deciduous old-growth forests in the Lower Rhine Embayment, Germany. Assemblages differed between larger (≥700 ha) and smaller forests (<400 ha), those of large forests held a higher percentage of forest species. Although α-diversity was similar between the two forest size classes, small forests often comprised matrix species, resulting in a higher β-diversity. Edge effects on the species richness of matrix species were noticeable up to 250 m into the forest. Hierarchical partitioning revealed that distance to disturbances (external edge, internal edges like roads) explained most assemblage variables, whereas forest size and woodland cover within a 1 km radius from the sites explained only a few assemblage variables. Densities of two forest-associated species, Discus rotundatus and Arion fuscus, decreased with forest size. Yet, forest size was positively correlated with richness of typical forest species and densities of Limax cinereoniger. The latter species seems to need forests of >1,000 ha, i.e., well above the size of most fragments. In conclusion, the prediction is valid only for forest species. The response to fragmentation is species specific and seems to depend on habitat specialization and macroclimatic conditions. Jean-Pierre Maelfait: Deceased.     

Importance of agricultural landscapes to nesting burrowing owls in the Northern Great Plains, USA

Anthropogenic habitat loss and fragmentation are the principle factors causing declines of grassland birds. Declines in burrowing owl (Athene cunicularia) populations have been extensive and have been linked to habitat loss, primarily the decline of black-tailed prairie dog (Cynomys ludovicianus) colonies. Development of habitat use models is a research priority and will aid conservation of owls inhabiting human-altered landscapes. From 2001 to 2004 we located 160 burrowing owl nests on prairie dog colonies on the Little Missouri National Grassland in North Dakota. We used multiple linear regression and Akaike’s Information Criterion to estimate the relationship between cover type characteristics surrounding prairie dog colonies and (1) number of owl pairs per colony and (2) reproductive success. Models were developed for two spatial scales, within 600 m and 2,000 m radii of nests for cropland, crested wheatgrass (Agropyron cristatum), grassland, and prairie dog colonies. We also included number of patches as a metric of landscape fragmentation. Annually, fewer than 30% of prairie dog colonies were occupied by owls. None of the models at the 600 m scale explained variation in number of owl pairs or reproductive success. However, models at the 2,000 m scale did explain number of owl pairs and reproductive success. Models included cropland, crested wheatgrass, and prairie dog colonies. Grasslands were not included in any of the models and had low importance values, although percentage grassland surrounding colonies was high. Management that protects prairie dog colonies bordering cropland and crested wheatgrass should be implemented to maintain nesting habitat of burrowing owls.     

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.     

Effects of Habitat Suitability and Landscape Patterns on Tick (Acarina) Metapopulation Processes

Tick density and population dynamics are important factors in the ecological processes involved in pathogen circulation in a habitat. These characteristics of tick populations are closely linked to habitat suitability, which reflects the limiting ecological factors and landscape features affecting tick populations; however, little work has been done on the regional assessment of habitat suitability. In this study, a regional model for the distribution and abundance of the tick Ixodes ricinus in central Spain is developed. An occurrence and an abundance model were constructed; climate and vegetation variables were found to be the main predictors of both occurrence and density in a relatively homogeneous matrix of habitat patches, whereas topographical variables were found to have small contributions and were therefore discarded. The residuals of the abundance model showed good correlation with the isolation of each patch. The predictive power of the abundance model was greatly enhanced by inclusion of the traversability (a measure of the permeability of each patch to the propagules of the metapopulation) and recruitment (an index of the relative importance of each patch to the traffic through the entire habitat network). The removal from the landscape of the patches whose recruitment values were in the top 10% has a critical effect on tick density, an effect not observed when patches are removed at random. These results indicate that permanent tick populations can be sustained only in landscapes containing a minimum network of viable sites. Graph theory and measurements of patch isolation should prove to be important elements in the forecasting of tick abundance and the management of the features underlying the landscape ecology of tick populations and pathogen circulation in the field.     

Utilization of a habitat mosaic by cotton rats during a population decline

This study describes the demographic features of a population of Sigmodon hispidus utilizing the habitat mosaic provided by a Carolina Bay on the Atlantic coastal plain of South Carolina. A total of 71 cotton rats were captured 160 times on a 4 ha grid during a winter decline from 25/ha to less that 1/ha. Body weights of adults declined until early February and then increased; those of subadults grew very slowly until February followed by a spurt in growth. Weight gain did not differ between survivors and non-survivors for males, but female survivors gained 1.5 g per week more than non-survivors. Female subadults exhibited higher mortality early in the decline and males later.Adult females were randomly distributed across 8 microhabitats, whereas adult males were almost exclusively confined to heavy Rubus cover. Subadult males used wet sites more than any other cohort; subadult females were widely distributed using drier sites most frequently. By the end of the decline, all survivors were localized in Rubus-dominated patches. No statistically significant changes in electromorph genotypes or allele frequencies were detected, but survivors had a higher frequency of the F-allele at the adenylate kinase locus than did non-survivors (42.3% vs. 16.7%).Our findings affirm the importance of a landscape perspective in understanding the population dynamics of cotton rats, and show how a habitat mosaic influences survival differentially among sex-age cohorts.     

Multi-scale landscape and seascape patterns associated with marbled murrelet nesting areas on the U.S. west coast

Habitat for wide-ranging species should be addressed at multiple scales to fully understand factors that limit populations. The marbled murrelet (Brachyramphus marmoratus), a threatened seabird, forages on the ocean and nests inland in large trees. We developed statistical relationships between murrelet use (occupancy and abundance) and habitat variables quantified across many spatial scales (statewide to local) and two time periods in California and southern Oregon, USA. We also addressed (1) if old-growth forest fragmentation was negatively associated with murrelet use, and (2) if some nesting areas are more important than others due to their proximity to high quality marine habitat. Most landscapes used for nesting were restricted to low elevation areas with frequent fog. Birds were most abundant in unfragmented old-growth forests located within a matrix of mature second-growth forest. Murrelets were less likely to occupy old-growth habitat if it was isolated (> 5 km) from other nesting murrelets. We found a time lag in response to fragmentation, where at least a few years were required before birds abandoned fragmented forests. Compared to landscapes with little tono murrelet use, landscapes with many murrelets were closer to the ocean's bays, river mouths, sandy shores, submarine canyons, and marine waters with consistently high primary productivity. Within local landscapes (≤ 800ha), inland factors limited bird abundance, but at the broadest landscape scale studied (3200 ha), proximity to marine habitat was most limiting. Management should focus on protecting or creating large, contiguous old-growth forest stands, especially in low-elevation areas near productive marine habitat. This revised version was published online in July 2006 with corrections to the Cover Date.     

Scale-dependent proximity of wildlife habitat in a spatially-neutral Bayesian model

Organisms may be constrained by the energetic costs incurred while obtaining resources in fragmented landscapes. We used a spatially neutral model of deer wintering habitat to evaluate the effects of landscape fragmentation on the aggregation of deer habitat. The spatially neutral model used Bayesian probabilities to predict where deer wintering areas occurred. The probabilities were conditional on 12 landscape variables measured at 22,750 contiguous 0.4 ha locations. The model predicted deer habitat at each location independently, thereby enabling a comparison of habitat aggregation in observed, simulated, and random distributions of deer habitat. The predictions of the neutral model exhibited greater fragmentation than observed in nature, suggesting that suitable, yet isolated, locations were not visited by deer. The most suitable sites for deer were clumped in the neutral model, regardless of scale. The inclusion of less suitable sites preserved significant aggregation at fine scales but not at broad scales. Species operate at different scales within a landscape, so ecologists, nature reserve designers and natural resource planners may benefit from models that focus on the proximity of habitat sites as a function of both spatial scale and habitat quality.     

A landscape-level assessment of understory diversity in upland forests of North-Central Wisconsin,USA

The study measured landscape level diversity of the understory plants of mature, upland forests in north-central Wisconsin USA. Habitat types were used to segregate the landscape along a moisture-nutrient gradient. Forty sites that had closed canopies, had been undisturbed for at least 20 years, and were at least 8 ha in size were used. The percent cover of groundlayer species was ocularly estimated in 12–18 randomly located, one meter square plots in June and August, 1995. Shrub cover was estimated by the line intercept method. Alpha, beta and gamma diversity were determined for early and late summer periods separately. Gamma diversity was quantified using a new method, affinity analysis, which generates a list of modal and outlier sites and calculates mosaic diversity, a measure of landscape complexity. Generally, communities in the middle of the moisture-nutrient gradient were modal, whereas those at the mesic end of the gradient were outlier. Mosaic diversity values were very similar for early summer and late summer (2.88±0.04, 2.95±0.03, respectively), but was much higher for both periods combined (3.95±0.07). Whittaker's Index (beta diversity) revealed varying rates of species turnover along presumed moisture and nutrient gradients, whereas species densities and richness were relatively constant among habitat types. A one-way analysis of variance of Shannon-Weaver values found no significant differences among habitat types (p0.05). Regional diversity mainly resulted from high beta values which appears to be primarily a function of the moisture gradient. The other factors influencing compositional differences among sites are variation in site history, especially disturbance, with niche partitioning and differences in seed dispersal capacity having a minor influence. The affinity analysis method indicated that sampling once per season is inadequate, and that many types of sites are modal. This method for estimating gamma (landscape) diversity shows considerable promise, but information on the processes that produce outlier sites is needed to fully understand and use the results of this method.     

Statistical models of landscape pattern metrics,with applications to regional scale dynamic forest simulations

Forest managers in Canada need to model landscape pattern or spatial configurationoverlarge (100,000 km²) regions. This presents a scalingproblem, as landscape configuration is measured at a high spatial resolution,but a low spatial resolution is indicated for regional simulation. We present astatistical solution to this scaling problem by showing how a wide range oflandscape pattern metrics can be modelled from low resolution data. Our studyarea comprises about 75,000 km² of boreal mixedwoodforest in northeast Alberta, Canada. Within this area we gridded a sample of 84digital forest cover maps, each about 9500 ha in size, to aresolution of 1 ha and used FRAGSTATS to compute a suite oflandscape pattern metrics for each map. We then used multivariate dimensionreduction techniques and canonical correlation analysis to model therelationship between landscape pattern metrics and simpler stand table metricsthat are easily obtained from non-spatial forest inventories. These analyseswere performed on four habitat types common in boreal mixedwood forests: youngdeciduous, old deciduous, white spruce, and mixedwood types. Using only threelandscape variables obtained directly from stand attribute tables (totalhabitatarea, and the mean and standard deviation of habitat patch size), ourstatistical models explained more than 73% of the joint variation in fivelandscape pattern metrics (representing patch shape, forest interior habitat,and patch isolation). By PCA, these five indices captured much of the totalvariability in the rich set of landscape pattern metrics that FRAGSTATS cangenerate. The predictor variables and strengths of association were highlyconsistent across habitat classes. We illustrate the potential use of suchstatistical relationships by simulating the regional, cumulative effects ofwildfire and forest management on the spatial arrangement of forest patches,using non-spatial stand attribute tables.This revised version was published online in May 2005 with corrections to the Cover Date.     

Importance of Wetland Landscape Structure to Shorebirds Wintering in an Agricultural Valley

Only recently has the influence of landscape structure on habitat use been a research focus in wetland systems. During non-breeding periods when food can be locally limited, wetland spatial pattern across a landscape may be of great importance in determining wetland use. We studied the influence of landscape structure on abundances of wintering Dunlin (Calidris alpina) and Killdeer (Charadrius vociferus) observed on wetlands in the agricultural Willamette Valley of Oregon, USA, during two winters (1999–2000, 2000–2001) of differing rainfall. We examined (1) shorebird use within a sample of 100 km² regions differing in landscape structure (hectares of shorebird habitat [wet, unvegetated]) and (2) use of sites differing in landscape context (area of shorebird habitat within a species-defined radius). For use of sites, we also assessed the influence of two local characteristics: percent of soil exposed and area of wet habitat. We analyzed data using linear regression and information-theoretic modeling. During the dry winter (2000–2001), Dunlin were attracted to regions with more wetland habitat and their abundances at sites increased with greater area of shorebird habitat within both the site and the surrounding landscape. In contrast, Dunlin abundances at sites were related to availability of habitat at only a local scale during the wet winter (1999–2000). Regional habitat availability was of little importance in predicting Killdeer distributions, and Killdeer site use appeared unrelated to habitat distributions at both landscape and local scales. Results suggest prioritizing sites for conservation that are located in areas with high wetland coverage.     

Disentangling habitat and social drivers of nesting patterns in songbirds

Nest locations of breeding birds are often spatially clustered. This tendency to nest together has generally been related to a patchy distribution of nesting habitat in landscape studies, but behavioral studies of species with clustered breeding patterns draw attention to the importance of social and biotic factors. Indeed, it is becoming increasingly apparent that the breeding system of many territorial, migrant birds may be semi-colonial. The reasons for, and extent of, spatial clustering in their breeding systems are not well understood. Our goal was to tease apart the influence of habitat availability and social drivers of clustered breeding in a neotropical migrant species, the hooded warbler (Wilsonia citrina). To test alternative hypotheses related to clustered habitat or conspecific attraction, we combined a habitat classification based on remote sensing with point pattern analysis of nesting sites. Nest locations (n = 150, 1999–2004), collected in a 1213 ha forested area of Southern Ontario (Canada), were analyzed at multiple spatial scales. Ripley’s K and pair-correlation functions g (uni- and bivariate) were used to test whether nests were clustered merely because potential nesting habitat was also clustered, or whether nests were additionally clustered with respect to conspecifics. Nest locations tended to be significantly clustered at intermediate distances (particularly between 240 and 420 m). Nests were randomly distributed within available habitat at larger distance scales, up to 1500 m. A reasonable hypothesis to explain the detected additional clustering, and one that is consistent with the results of several behavioral studies, is that females pack their nests more tightly than the available habitat requires to be situated closer to their neighbors’ mates. Linking spatially explicit, point pattern analysis with strong inference based on Monte Carlo tests may bring us closer to understanding the generality and reasons behind conspecific attraction at different spatial scales. F. Csillag—deceased.     

Landscape-level movements of North American elk (<Emphasis Type="Italic">Cervus elaphus</Emphasis>): effects of habitat patch structure and topography

We examined movements of North American elk (Cervus elaphus) in northeastern Oregon, USA. Movement vectors at 449 locations over a 7762 ha area were calculated based on 16,724 sequential observations of 94 female elk-year combinations during spring (15 April–14 May) 1993, 1995, 1996. We calculated movement vectors at the start of morning and evening feeding bouts (0500, 1900 h) and during periods of least activity (0100, 1500 h). Here, we measured characteristics of habitat patches (habitat type, mean patch size, coefficient of variation in patch size, edge density, mean shape index, and mean nearest neighbor) at two levels of habitat grain (eight habitat types, two habitat types) and at three spatial scales (250, 500, and 1000 m) around each movement vector. We also measured topographic features around each vector including distance to nearest stream, direction of drainage, elevation, slope, and convexity (a measure of ridge top vs. valley bottom land form). We used mixed models adjusted for positive spatial correlation among vectors to examine the relationship between vector length, or speed of movement, and habitat patch characteristics, and between vector direction and topographic features. Speed of movements by elk were not related to characteristics of habitat patches that we measured. The direction of movement, however, was dependent on topography. Elk were more likely to move parallel to major drainages than perpendicular to them. Furthermore, elk were less likely to move perpendicular to drainages when close to the nearest stream, in valley bottoms vs. ridge tops, and on steep slopes. The dendritic nature of movements by elk with respect to topography may help elucidate ecosystem processes such as nutrient flows, nutrient cycling, and successional trajectories of plant communities.     

Role of habitat and landscape in structuring small mammal assemblages in hedgerow networks of contrasted farming landscapes in Brittany,France

In this study, we investigated the environmental factors driving small mammal (rodents and shrews) assemblages in permanent habitat patches in response to a gradient of agricultural intensification. Small mammals were sampled using a trapping standard method in the hedgerow networks of three contrasted landscapes differing by their level of land-use intensity and hedgerow network density (BOC1: slightly intensified; BOC2: moderately intensified and POL: highly intensified). We hypothesized that habitat and landscape characteristics have to be considered to understand the structure of local community. In that way, we carried out a multi-scale study using environmental variables ranging from local habitat (structure and composition of the hedgerows) to hedgerows neighbourhoods in a radius of 300 m (land cover and connectivity around hedges) and to landscape units (three sites). During 1 year, 24 hedgerows were sampled seven times, representing a total of 1,379 captures (86% of rodents and 14% of shrews) and eight species, dominated by the wood mouse (Apodemus sylvaticus) and the bank vole (Clethrionomys glareolus). Inter-site variability was significant and accounted for 18% of total variation in small mammal species abundances. But intra-site variability was also highlighted: species abundance profiles may differ greatly among hedgerows within a site. The more explanatory variables were identified at the different scales of the study: the landscape unit POL was shown to be an important factor in structuring the community, but the predominant factors explaining differences of abundances among hedgerows were about local habitat. In fact, the width of hedges and the tree species richness appeared to be significant and explaining the greatest part of the total variation of the small mammal community composition.     

Predicting carnivore occurrence with noninvasive surveys and occupancy modeling

Terrestrial carnivores typically have large home ranges and exist at low population densities, thus presenting challenges to wildlife researchers. We employed multiple, noninvasive survey methods—scat detection dogs, remote cameras, and hair snares—to collect detection–nondetection data for elusive American black bears (Ursus americanus), fishers (Martes pennanti), and bobcats (Lynx rufus) throughout the rugged Vermont landscape. We analyzed these data using occupancy modeling that explicitly incorporated detectability as well as habitat and landscape variables. For black bears, percentage of forested land within 5 km of survey sites was an important positive predictor of occupancy, and percentage of human developed land within 5 km was a negative predictor. Although the relationship was less clear for bobcats, occupancy appeared positively related to the percentage of both mixed forest and forested wetland habitat within 1 km of survey sites. The relationship between specific covariates and fisher occupancy was unclear, with no specific habitat or landscape variables directly related to occupancy. For all species, we used model averaging to predict occurrence across the study area. Receiver operating characteristic (ROC) analyses of our black bear and fisher models suggested that occupancy modeling efforts with data from noninvasive surveys could be useful for carnivore conservation and management, as they provide insights into habitat use at the regional and landscape scale without requiring capture or direct observation of study species.     

Habitat variables explain Loggerhead Shrike occurrence in the northern Chihuahuan Desert,but are poor correlates of fitness measures

Conservation efforts should be based on habitat models that identify areas of high quality and that are built at spatial scales that are ecologically relevant. In this study, we developed habitat models for the Loggerhead Shrike (Lanius ludovicianus) in the Chihuahuan Desert of New Mexico to answer two questions: (1) are highly used habitats of high quality for shrikes in terms of individual fitness? and (2) what are the spatial scales of habitat associations relevant to this species? Our study area was Fort Bliss Army Reserve (New Mexico). Bird abundance was obtained from 10 min point counts conducted at forty-two 108 ha plots during a 3-year period. Measures of fitness were obtained by tracking a total of 73 nests over the 3 years. Habitat variables were measured at spatial scales ranging from broad to intermediate to local. We related habitat use and measures of fitness to habitat variables using Bayesian model averaging. We found a significant relationship between bird abundance and measures of fitness averaged across nesting birds in each plot (correlation up to 0.61). This suggests that measures of habitat use are indicative of habitat quality in the vicinity of Fort Bliss. Local- and intermediate-scale variables best explained shrike occurrence. Habitat variables were not related to any measures of fitness. A better understanding of the factors that limit individual bird fitness is therefore necessary to identify areas of high conservation value for this species.     

Influence of patch and landscape characteristics on the distribution of the subterranean rodent <Emphasis Type="Italic">Ctenomys porteousi</Emphasis>

The understanding and prediction of the responses of animal populations to habitat fragmentation is a central issue in applied ecology. The identification of habitat variables associated to patch occupancy is particularly important when habitat quality is affected by human activities. Here, we analyze the influence of patch and landscape characteristics on patch occupancy by the subterranean herbivorous rodent Ctenomys porteousi. Patch occupancy was monitored in a network of 63 habitat patches identified by satellite imagery analysis which extends along almost the whole distributional range for C. porteousi. Suitable habitat for the occurrence of C. porteousi is highly fragmented and represents <10% of the total area in its distributional range. The distribution of C. porteousi in the patch network is affected not only by characteristics of the habitat patches, but also by those of the surrounding landscape matrix. Significant differences between occupied and empty patches were found in several environmental variables. Overall, occupied patches were larger, less vegetated, more connected, and had larger neighbor patches than empty patches. A stepwise procedure on a generalized linear model selected four habitat variables that explain patch occupancy in C. porteousi; it included the effects of habitat quality in the matrix surrounding the patch, average vegetation cover in the patch, minimum vegetation cover in the matrix surrounding the patch, and the area of the nearest neighbor patch. These results indicate that patch occupancy in C. porteousi is strongly influenced by the availability and quality of habitat both in the patch and in the surrounding landscape matrix.     

Effects of spatial habitat heterogeneity on habitat selection and annual fecundity for a migratory forest songbird

Understanding how spatial habitat patterns influence abundance and dynamics of animal populations is a primary goal in landscape ecology. We used an information-theoretic approach to investigate the association between habitat patterns at multiple spatial scales and demographic patterns for black-throated blue warblers (Dendroica caerulescens) at 20 study sites in west-central Vermont, USA from 2002 to 2005. Sites were characterized by: (1) territory-scale shrub density, (2) patch-scale shrub density occurring within 25 ha of territories, and (3) landscape-scale habitat patterns occurring within 5 km radius extents of territories. We considered multiple population parameters including abundance, age ratios, and annual fecundity. Territory-scale shrub density was most important for determining abundance and age ratios, but landscape-scale habitat structure strongly influenced reproductive output. Sites with higher territory-scale shrub density had higher abundance, and were more likely to be occupied by older, more experienced individuals compared to sites with lower shrub density. However, annual fecundity was higher on sites located in contiguously forested landscapes where shrub density was lower than the fragmented sites. Further, effects of habitat pattern at one spatial scale depended on habitat conditions at different scales. For example, abundance increased with increasing territory-scale shrub density, but this effect was much stronger in fragmented landscapes than in contiguously forested landscapes. These results suggest that habitat pattern at different spatial scales affect demographic parameters in different ways, and that effects of habitat patterns at one spatial scale depends on habitat conditions at other scales.     

Response of mountain plovers to plague-driven dynamics of black-tailed prairie dog colonies

Sylvatic plague is a major factor influencing the dynamics of black-tailed prairie dog (Cynomys ludovicianus) colonies in the western Great Plains. We studied the nesting response of the mountain plover (Charadrius montanus), a grassland bird that nests on prairie dog colonies, to plague-driven dynamics of prairie dog colonies at three sites in the western Great Plains. First, we examined plover nest distribution on colonies that were previously affected by plague, but that had been recovering (expanding) for at least 6 years. Plovers consistently nested in both young (colonized in the past 1–2 years) and old (colonized for 6 or more years) portions of prairie dog colonies in proportion to their availability. Second, we examined changes in plover nest frequency at two sites following plague epizootics, and found that mountain plover nest numbers declined relatively rapidly (≤2 years) on plague-affected colonies. Taken together, our findings indicate that available plover nesting habitat associated with prairie dog colonies closely tracks the area actively occupied by prairie dogs each year. Given the presence of plague throughout most of the mountain plover’s breeding range in the western Great Plains, important factors affecting plover populations likely include landscape features that determine the scale of plague outbreaks, the distance that plovers move in response to changing breeding habitat conditions, and the availability and quality of alternate breeding habitat within the landscape.