The protean relationship between boreal forest landscape structure and red squirrel distribution at multiple spatial scales

This paper investigates two fundamental questions in landscape ecology: what influence does landscape context, or the composition of the matrix, have on an animals’ response to landscape structure, and how does this relationship extrapolate between landscapes? We investigate how the distribution of North American red squirrels (Tamiasciurus hudsonicus) in the boreal mixedwood forest is influenced by anthropogenically (forest harvest) and naturally (forest fire) derived landscape structure. We studied the presence and absence of red squirrels over two years in three landscape types: one managed for timber harvest, one recently burned by wildfire, and a third unburned unmanaged landscape. Landscape composition and configuration, measured at several spatial scales, predicted red squirrel’s distribution in all three landscapes, but the significant landscape variables changed across spatial scales, across time, and across landscapes. These findings emphasize the variability in landscape structure/animal distribution relationships, and enforce the need to link pattern-finding studies, such as this one, with searches for the mechanisms behind the observed pattern.     

The structure of boreal old-growth forests changes at multiple spatial scales over decades

Landscape Ecology - Changes in the structure of boreal old-growth forests are typically studied at a specific spatial scale. Consequently, little is known about forest development across different...     

Landscape effects on mallard habitat selection at multiple spatial scales during the non-breeding period

Previous studies that evaluated effects of landscape-scale habitat heterogeneity on migratory waterbird distributions were spatially limited and temporally restricted to one major life-history phase. However, effects of landscape-scale habitat heterogeneity on long-distance migratory waterbirds can be studied across the annual cycle using new technologies, including global positioning system satellite transmitters. We used Bayesian discrete choice models to examine the influence of local habitats and landscape composition on habitat selection by a generalist dabbling duck, the mallard (Anas platyrhynchos), in the midcontinent of North America during the non-breeding period. Using a previously published empirical movement metric, we separated the non-breeding period into three seasons, including autumn migration, winter, and spring migration. We defined spatial scales based on movement patterns such that movements >0.25 and <30.00 km were classified as local scale and movements >30.00 km were classified as relocation scale. Habitat selection at the local scale was generally influenced by local and landscape-level variables across all seasons. Variables in top models at the local scale included proximities to cropland, emergent wetland, open water, and woody wetland. Similarly, variables associated with area of cropland, emergent wetland, open water, and woody wetland were also included at the local scale. At the relocation scale, mallards selected resource units based on more generalized variables, including proximity to wetlands and total wetland area. Our results emphasize the role of landscape composition in waterbird habitat selection and provide further support for local wetland landscapes to be considered functional units of waterbird conservation and management.     

Responses of Chilean forest birds to anthropogenic habitat fragmentation across spatial scales

Although it is recognized that anthropogenic forest fragmentation affects habitat use by organisms across multiple spatial scales, there is uncertainty about these effects. We used a hierarchical sampling design spanning three spatial scales of habitat variability (landscape > patch > within-patch) and generalized mixed-effect models to assess the scale-dependent responses of bird species to fragmentation in temperate forests of southern Chile. The abundances of nine of 20 bird species were affected by interactions across spatial scales. These interactions resulted in a limited effect of within-patch habitat structure on the abundance of birds in landscapes with low forest cover, suggesting that suitable local habitats, such as sites with dense understory cover or large trees, are underutilized or remain unused in highly fragmented landscapes. Habitat specialists and cavity-nesters, such as tree-trunk foragers and tapaculos, were most likely to exhibit interactions across spatial scales. Because providing additional sites with dense understory vegetation or large habitat trees does not compensate the negative effect of the loss of forest area on bird species, conservation strategies should ensure the retention of native forest patches in the mixed-use landscapes.     

Temporal change in forest fragmentation at multiple scales

Previous studies of temporal changes in fragmentation have focused almost exclusively on patch and edge statistics, which might not detect changes in the spatial scale at which forest occurs in or dominates the landscape. We used temporal land-cover data for the Chesapeake Bay region and the state of New Jersey to compare patch-based and area–density scaling measures of fragmentation for detecting changes in the spatial scale of forest that may result from forest loss. For the patch-based analysis, we examined changes in the cumulative distribution of patch sizes. For area–density scaling, we used moving windows to examine changes in dominant forest. We defined dominant forest as a forest parcel (pixel) surrounded by a neighborhood in which forest occupied the majority of pixels. We used >50% and ≥60% as thresholds to define majority. Moving window sizes ranged from 2.25 to 5,314.41 hectares (ha). Patch size cumulative distributions changed very little over time, providing no indication that forest loss was changing the spatial scale of forest. Area–density scaling showed that dominant forest was sensitive to forest loss, and the sensitivity increased nonlinearly as the spatial scale increased. The ratio of dominant forest loss to forest loss increased nonlinearly from 1.4 to 1.8 at the smallest spatial scale to 8.3 to 11.5 at the largest spatial scale. The nonlinear relationship between dominant forest loss and forest loss in these regions suggests that continued forest loss will cause abrupt transitions in the scale at which forest dominates the landscape. In comparison to the Chesapeake Bay region, dominant forest loss in New Jersey was less sensitive to forest loss, which may be attributable the protected status of the New Jersey Pine Barrens.     

Site fidelity of female caribou at multiple spatial scales

Studies of site fidelity have been hampered by arbitrary designations of spatial scale and the lack of null models for comparison. We generated null expectations of fidelity at different scales from the distribution of radio-tracked animals in a population. We applied the models to space use of satellite-tracked caribou (Rangifer tarandus caribou), the most vagile nonvolant terrestrial animal, from populations representing sedentary and migratory ecotypes. We compared distances between consecutive-year locations of adult females to expectations based on the total range and seasonal range of each population. At the scale of the total range, sedentary and migratory caribou displayed remarkably similar philopatry, despite a 30-fold difference in size of their population ranges, from time of calving (late May) to breeding (late October). The most intense fidelity occurred during post-calving when, on average, sedentary and migratory females returned to as near as 6.7 km and 123 km, respectively, of locations occupied the previous year. At the scale of the seasonal range, the ecotypes differed. Sedentary caribou still displayed fidelity from calving to breeding; migratory caribou exhibited fidelity only during late autumn. For migratory, but not sedentary caribou, inter-year distances during winter were negatively correlated with age, implying that older females were more philopatric. We conclude that reproductive activities delimit the season of fidelity of female caribou of both ecotypes, and that scale-dependent ecotypic differences in fidelity may reflect different factors of population limitation. A spatially-explicit approach to site fidelity is essential for synthesizing patterns across studies.     

Behavioral switching in Magellanic woodpeckers reveals perception of habitat quality at different spatial scales

Context

The switching pattern between behavioral modes provides a mechanistic basis for understanding how animals perceive and memorize the habitat quality in their home ranges.

Objectives

We assessed if Magellanic woodpeckers (Campephilus magellanicus) move based on habitat quality at local (neighboring trees) and home range scales.

Methods

We used state-space models to examine the relationship between remotely-sensed estimates of habitat quality (tree decay) and movement of adult woodpeckers tracked with GPS telemetry in southern Chile.

Results

Woodpeckers spent most time (>?80%) in the area-restricted search (ARS) mode in contrast to the exploratory transient mode, characterized by frequent directional displacements (>?50 m). The extent to which individuals switched between behavioral modes was related to habitat quality at different scales. Woodpeckers switched to and remained in the ARS mode when encountering moderate levels of heterogeneity in habitat quality. At very low or high heterogeneity levels, however, individuals switched to and remained in the transient mode, respectively. Likewise, as habitat quality declined locally and across home range, woodpeckers were more likely to adopt a transient mode.

Conclusions

Although woodpeckers seemed to easily perceive and memorize habitat quality at different spatial scales, our results suggest that spatial memory will less effective under extreme levels of habitat heterogeneity.

     

Projecting land use transitions at forest fringes in the Philippines at two spatial scales

This paper presents two applications of a spatially explicit model of land use change at two spatial scales: a nation-wide application for the Philippines at relatively coarse resolution and an application with high spatial resolution for one island of the Philippines: Sibuyan island, Romblon province. The model is based on integrated analysis of socio-economic and biophysical factors that determine the allocation of land use change in combination with the simulation of the temporal dynamics (path-dependence and reversibility of changes), spatial policies and land requirements. Different scenarios of near-future developments in land use pattern are simulated illustrating the effects of implementing spatial policies. Results from the coarse scale model with national extent mainly serve to identify the overall pattern of land use change and hot zones of deforestation. The detailed application provides more insight in the pattern of land use change and its consequences for ecological processes. The use of the results for environmental assessments is illustrated by calculating spatial indices to assess the impact of land use change on forest fragmentation. It is concluded that spatially explicit modeling of land use change yields important information for environmental management and land use planning. The applications illustrate that the scale of analysis is an important determinant of the model configuration, the interpretation of the results and the potential use by stakeholders. There is no single, optimal, scale for land use change assessments. Each scale enables different types of analysis and assessment: applications at multiple scales therefore give complementary information needed for environmental management.This revised version was published online in May 2005 with corrections to the Cover Date.     

Quantifying edges as gradients at multiple scales improves habitat selection models for northern spotted owl

Context

Testing the influence of edges on animal distributions depends on our capacity to quantify ‘edge’, particularly in heterogeneous landscapes. Habitat quality is likely to differ in instances where edges are abrupt and anthropogenic in origin, versus diffuse, disturbance-created edges.

Objectives

We tested whether or not structurally distinct edge types influence northern spotted owl habitat selection and whether the relationship between edge type and use varied across spatial scales relevant to owl foraging (<3 ha) and home range selection (50–800 ha).

Methods

We used remotely sensed disturbance severity data to define two distinct edge types, ‘hard’ and ‘diffuse’, following a 11,000 ha fire and subsequent salvage logging in southern Oregon. The approach quantifies the steepness of gradients directly by measuring the ‘slope’ of change in disturbance severity. We tested the degree to which 23 radio-collared spotted owls responded to edge characteristics caused by fire and logging.

Results

Spotted owls showed a strong negative association with hard edge, even after accounting for habitat suitability and other confounding variables. However, this negative relationship was highly scale-dependent; spotted owls were resilient to hard edges at broad scales, but avoided the same feature at fine scales. On the other hand, spotted owls showed a positive association with diffuse edge, especially at broader scales.

Conclusions

Differential use of edge types indicates that owls favor disturbances that create diffuse edge habitat (e.g. low and mixed-severity fire) and rather than abrupt boundaries created by high severity disturbance.

     

Landscape influences on stream biotic integrity assessed at multiple spatial scales

The biological integrity of stream ecosystems depends critically on human activities that affect land use/cover along stream margins and possibly throughout the catchment. We evaluated stream condition using an Index of Biotic Integrity (IBI) and a habitat index (HI), and compared these measures to landscape and riparian conditions assessed at different spatial scales in a largely agricultural Midwestern watershed. Our goal was to determine whether land use/cover was an effective predictor of stream integrity, and if so, at what spatial scale. Twenty-three sites in first-through third-order headwater streams were surveyed by electrofishing and site IBIs were calculated based on ten metrics of the fish collection. Habitat features were characterized through field observation, and site HIs calculated from nine instream and bank metrics. Field surveys, aerial photograph interpretation, and geographic information system (GIS) analyses provided assessments of forested land and other vegetation covers at the local, reach, and regional (catchment) scales. The range of conditions among the 23 sites varied from poor to very good based on IBI and HI scores, and habitat and fish assemblage measures were highly correlated. Stream biotic integrity and habitat quality were negatively correlated with the extent of agriculture and positively correlated with extent of wetlands and forest. Correlations were strongest at the catchment scale (IBI with % area as agriculture, r²=0.50, HI with agriculture, r²=0.76), and tended to become weak and non-significant at local scales. Local riparian vegetation was a weak secondary predictor of stream integrity. In this watershed, regional land use is the primary determinant of stream conditions, able to overwhelm the ability of local site vegetation to support high-quality habitat and biotic communities.     

Using remote-sensing data to assess habitat selection of a declining passerine at two spatial scales

Context

Detailed information on habitat needs is integral to identify conservation measures for declining species. However, field data on habitat structure is typically limited in extent. Remote sensing has the potential to overcome these limitations of field-based studies.

Objective

We aimed to assess abiotic and biotic characteristics of territories used by the declining wood warbler (Phylloscopus sibilatrix), a forest-interior migratory passerine, at two spatial scales by evaluating a priori expectations of habitat selection patterns.

Methods

First, territories established by males before pairing, referred to as pre-breeding territories, were compared to pseudo-absence control areas located in the wider forested landscape (first spatial scale, N_territories = 66, N_controls = 66). Second, breeding territories of paired wood warblers were compared to true-absence control areas located immediately close-by in the forest (second spatial scale, N_territories = 78, N_controls = 78). Habitat variables predominantly described forest structure and were mainly based on first and last pulse lidar (light detection and ranging) data.

Results

Occurrence of pre-breeding territories was related to vegetation height, vertical diversity and stratification, canopy cover, inclination and solar radiation. Occurrence of breeding territories was associated to vegetation height, vertical diversity and inclination.

Conclusions

Territory selection at the two spatial scales addressed was governed by similar factors. With respect to conservation, habitat suitability for wood warblers could be retained by maintaining a shifting mosaic of stand ages and structures at large spatial scales. Moreover, leaf-off lidar variables have the potential to contribute to understanding the ecological niche of species in predominantly deciduous forests.

     

Quantifying patch distribution at multiple spatial scales: Applications to wildlife-habitat models

Multiscale analyses are widely employed for wildlife-habitat studies. In most cases, however, each scale is considered discrete and little emphasis is placed on incorporating or measuring the responses of wildlife to resources across multiple scales. We modeled the responses of three Arctic wildlife species to vegetative resources distributed at two spatial scales: patches and collections of patches aggregated across a regional area. We defined a patch as a single or homogeneous collection of pixels representing 1 of 10 unique vegetation types. We employed a spatial pattern technique, three-term local quadrat variance, to quantify the distribution of patches at a larger regional scale. We used the distance at which the variance for each of 10 vegetation types peaked to define a moving window for calculating the density of patches. When measures of vegetation patch and density were applied to resource selection functions, the most parsimonious models for wolves and grizzly bears included covariates recorded at both scales. Seasonal resource selection by caribou was best described using a model consisting of only regional scale covariates. Our results suggest that for some species and environments simple patch-scale models may not capture the full range of spatial variation in resources to which wildlife may respond. For mobile animals that range across heterogeneous areas we recommend selection models that integrate resources occurring at a number of spatial scales. Patch density is a simple technique for representing such higher-order spatial patterns.     

Quantifying patch distribution at multiple spatial scales: Applications to wildlife-habitat models

Multiscale analyses are widely employed for wildlife-habitat studies. In most cases, however, each scale is considered discrete and little emphasis is placed on incorporating or measuring the responses of wildlife to resources across multiple scales. We modeled the responses of three Arctic wildlife species to vegetative resources distributed at two spatial scales: patches and collections of patches aggregated across a regional area. We defined a patch as a single or homogeneous collection of pixels representing 1 of 10 unique vegetation types. We employed a spatial pattern technique, three-term local quadrat variance, to quantify the distribution of patches at a larger regional scale. We used the distance at which the variance for each of 10 vegetation types peaked to define a moving window for calculating the density of patches. When measures of vegetation patch and density were applied to resource selection functions, the most parsimonious models for wolves and grizzly bears included covariates recorded at both scales. Seasonal resource selection by caribou was best described using a model consisting of only regional scale covariates. Our results suggest that for some species and environments simple patch-scale models may not capture the full range of spatial variation in resources to which wildlife may respond. For mobile animals that range across heterogeneous areas we recommend selection models that integrate resources occurring at a number of spatial scales. Patch density is a simple technique for representing such higher-order spatial patterns.     

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.     

Modeling and field-testing of Ovenbird (<Emphasis Type="Italic">Seiurus aurocapillus</Emphasis>) responses to boreal forest dissection by energy sector development at multiple spatial scales

Although the area disturbed by linear features in forested systems is small relative to many other human disturbances, linear features create significantly more amounts of edge per unit area. In the boreal plains of Alberta, Canada, energy sector exploration has resulted in extensive dissection of the landscape through 8 m wide seismic lines. A spatially explicit model was developed to test how bird abundance might change in response to increasing seismic line density if individuals use seismic lines as territory boundaries or actively avoid these edges. Assuming birds had fixed territory shape and size, increasing seismic line density from 0 to 8 km/km² resulted in a 38% decline and an 82% decline in bird abundance when individuals used lines as territory boundaries or avoided edges by 50 m, respectively. We tested the assumptions of our model using the Ovenbird (Seiurus aurocapillus). Based on radio-telemetry (n = 12), all Ovenbirds crossed seismic lines at some point during the breeding season. However, male Ovenbirds showed a distinct use of one side of the seismic line, suggesting lines acted as territory boundaries. In 12.25 ha plots (n = 24) spot-mapping detected no change in Ovenbird density as linear feature density increased from 0 to 8.6 km/km². In 4 km² landscapes (n = 62) sampled using a grid of nine point-counts, we also detected no changes in Ovenbird numbers across the same range of seismic line densities. Ovenbirds declined with seismic line density at the level of the individual point-count station (12 ha scale), but only when a threshold seismic line density of 8.5 km/km² was reached. Above the threshold, Ovenbirds declined 19% for each 1 km/km² increase in seismic line density. While relatively few places in Alberta’s boreal forest have local seismic line densities of 8.5 km/km², forest dissection could increasingly become an issue if current energy exploration practices continue.     

Butterflies in Swedish grasslands benefit from forest and respond to landscape composition at different spatial scales

Context

Loss and fragmentation of semi-natural grasslands has critically affected many butterfly species in Europe. Habitat area and isolation can have strong effects on the local biodiversity but species may also be strongly affected by the surrounding matrix.

Objectives

We explored how different land cover types in the landscape explained the occurrence of butterfly species in semi-natural grasslands.

Methods

Using data from 476 semi-natural grasslands in Sweden, we analysed the effect of matrix composition on species richness and occurrence. Additionally, we analysed at which spatial scales butterflies responded to matrix types (forests, semi-natural grasslands, arable land and water).

Results

Forest cover showed the strongest positive effect on species richness, followed by semi-natural grasslands. Forest also had a positive effect on red-listed species at local scales. Responses to matrix composition were highly species-specific. The majority of the 30 most common species showed strong positive responses to the amount of forest cover within 200–500 m. There was a smaller group of species showing a positive response to arable land cover within 500–2000 m. Thirteen species showed positive responses to the amount of semi-natural grasslands, generally at larger scales (10–30 km).

Conclusions

Our study showed that surrounding forest is beneficial for many grassland butterfly species and that forests might mitigate the negative effects of habitat loss caused by agricultural intensification. Also, semi-natural grasslands were an important factor for species richness at larger spatial scales, indicating that a landscape consisting mainly of supporting habitats (i.e. forests) are insufficient to sustain a rich butterfly fauna.

     

Effects of bird and bat exclusion on coffee pest control at multiple spatial scales

Context

Despite the key role of biological control in agricultural landscapes, we still poorly understand how landscape structure modulates pest control at different spatial scales.

Objectives

Here we take an experimental approach to explore whether bird and bat exclusion affects pest control in sun coffee plantations, and whether this service is consistent at different spatial scales.

Methods

We experimentally excluded flying vertebrates from coffee plants in 32 sites in the Brazilian Atlantic Forest, encompassing a gradient of forest cover at landscape (2 km radius) and local (300 m) spatial scales, and quantified coffee leaf loss, as an indicator of herbivory, and fruit set.

Results

Leaf loss decreased with higher landscape forest cover, but this relation was significantly different between treatment and control plants depending on local forest cover. On the other hand, fruit set responded to the interaction between treatment and local forest cover but was not affected by landscape forest cover. More specifically, fruit set increased significantly with local forest cover in exclusion treatments and showed a non-significant decrease in open controls.

Conclusions

These results suggest that services provided by flying vertebrates are modulated by processes occurring at different spatial scales. We posit that in areas with high local forest cover flying vertebrates may establish negative interactions with predaceous arthropods (i.e. intraguild predation), but this would not be the case in areas with low local forest cover. We highlight the importance of employing a multi-scale analysis in systems where multiple species, which perceive the landscape differently, are providing ecosystem services.

     

Relative importance of habitat area and isolation for bird occurrence patterns in a naturally patchy landscape

There is debate among ecologists about whether total habitat area or patch arrangement contributes most to population and/or community responses to fragmented or patchy landscapes. We tested the relative effects of patch area and isolation for predicting bird occurrence in a naturally patchy landscape in the Bear River Mountains of Northern Utah, USA. We selected focal patches (mountain meadows) ranging in elevation from 1,920 to 2,860 m and in size from 0.6 to 182 ha. Breeding birds were sampled in each focal meadow during the summers of 2003 and 2004 using variable-distance point transects. Logistic regression and likelihood-based model selection were used to determine the relationship between likelihood of occurrence of three bird species (Brewer’s sparrow, vesper sparrow, and white-crowned sparrow) and area, isolation, and proximity metrics. We used model weights and model-averaged confidence intervals to assess the importance of each predictor variable. Plots of area versus isolation were used to evaluate complex relationships between the variables. We found that meadow area was the most important variable for explaining occurrence for two species, and that isolation was the most important for the other. We also found that the absolute distance was more appropriate for evaluating isolation responses than was the species-specific proximity metric. Our findings add clarity to the debate between ecologists regarding the relative importance of area and isolation in species responses to patchy landscapes.     

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

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