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1.

Context

In heterogeneous landscapes, habitat complementation is a key process underlying the distribution of mobile species able to exploit non-substitutable resources over large home ranges. For instance, insectivorous bats need to forage in a diversity of habitat patches offering varied compositions and structures within forest landscape mosaics to fulfill their life cycle requirements.

Objectives

We aimed at analyzing the effects of forest structure and composition measured at the stand and landscape scales on bat species richness, abundance and community composition in pine plantation forests of south-western France.

Methods

We sampled bat communities at different periods of the summer season using automatic ultrasound recorders along a tree composition gradient from pine monocultures to pure oak stands. We analyzed bat species activity (as a proxy for bat abundance) and species richness with linear mixed models. Distance-based constrained ordinations were used to partition the spatio-temporal variation in bat communities.

Results

Deciduous tree cover increased bat activity and modified community composition at both stand and landscape scales. Changes in bat communities were mostly driven by landscape-scale variables while bat activity responded more to stand-scale predictors.

Conclusions

The maintenance of deciduous trees at both stand and landscape scales is likely critical for bat communities living in fast-growing conifer plantations, by increasing the availability and diversity of prey and roosting sites. Our study suggests that bats respond to forest composition at both stand and landscape scales in mosaic plantation landscapes, mainly through a resource complementation process.
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2.

Context

The patch-mosaic model is lauded for its conceptual simplicity and ease with which conventional landscape metrics can be computed from categorical maps, yet many argue it is inconsistent with ecological theory. Gradient surface models (GSMs) are an alternative for representing landscapes, but adoption of surface metrics for analyzing spatial patterns in GSMs is hindered by several factors including a lack of meaningful interpretations.

Objectives

We investigate the performance and applicability of surface metrics across a range of ecoregions and scales to strengthen theoretical foundations for their adoption in landscape ecology.

Methods

We examine metric clustering across scales and ecoregions, test correlations with patch-based metrics, and provide ecological interpretations for a variety of surface metrics with respect to forest cover to support the basis for selecting surface metrics for ecological analyses.

Results

We identify several factors complicating the interpretation of surface metrics from a landscape perspective. First, not all surface metrics are appropriate for landscape analyses. Second, true analogs between surface metrics and patch-based, landscape metrics are rare. Researchers should focus instead on how surface measures can uniquely measure spatial patterns. Lastly, scale dependencies exist for surface metrics, but relationships between metrics do not appear to change considerably with scale.

Conclusions

Incorporating gradient surfaces into landscape ecological analyses is challenging, and many surface metrics may not have patch analogs or be ecologically relevant. For this reason, surface metrics should be considered in terms of the set of pattern elements they represent that can then be linked to landscape characteristics.
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3.

Context

The roosting habits of many temperate zone bats are well documented at microhabitat scales, but fewer studies have included multi-scale assessments of landscape patterns in bat roost site selection.

Objectives

To identify and assess at the landscape-scale the location of spring and early season maternity roosts of female northern long-eared bats (Myotis septentrionalis) from 2015 to 2016 at Mammoth Cave National Park (MACA), Kentucky, USA.

Methods

We used mist-nets and radiotelemetry to catch and track bats to roost trees across the landscape of MACA. Data on roosting sites were evaluated using spatial point pattern analysis to examine distributional trends of roosts. A variety of spatial covariates were used to model the effect of landscape pattern, including: forest type, elevation, and proximity to hibernacula, water, and road corridors.

Results

Data indicate that roost locations of female northern long-eared bats in MACA were typically situated within 2000 m of known winter hibernacula, occurring more often at higher elevations in mesic upland deciduous forests, and in close proximity to water sources and roads. We present hypotheses to account for the patterns observed in relation to landscape features and habitat resources in the Park.

Conclusions

Our data indicate that a more comprehensive understanding of habitat requirements which includes empirically-based, landscape-scale patterns, and not solely considerations at stand or local levels, could lead to better informed management policies targeting conservation of maternity habitat of forest-dwelling bats, including the northern long-eared bat, a species in decline throughout much of its distribution in North America.
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4.

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.
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5.

Context

Urban sprawl and the expanding transportation infrastructure drive land consumption and landscape fragmentation, causing environmental deterioration and loss of species. Current understanding of how these drivers interact to shape landscape fragmentation is still poor. However, a strong correlation between urban sprawl and landscape fragmentation patterns is commonly assumed.

Objectives

Our main objective was to test the strength, non-stationarity, and scale-dependency of the relationship between urban sprawl and landscape fragmentation patterns (‘sprawl-fragmentation relationship’). Subsequently, we propose an extended framework for the links between urban sprawl, expansion of transport infrastructure, and landscape fragmentation.

Methods

We quantified spatial patterns of urban sprawl and landscape fragmentation for mainland Spain at multiple scales. We then fitted global regression models and geographically weighted regression models with metrics of landscape fragmentation and urban sprawl.

Results

Most variation in landscape fragmentation values (almost 80 % on average) is not explained by urban sprawl metrics through global modeling. Local models show substantial improvements in model performance, with an average of 37 % of the variance remaining unexplained. The contribution of urban sprawl to landscape fragmentation patterns varies locally and depends on scale, with higher contributions at coarser scales and at higher organizational levels.

Conclusions

Our investigation revealed three critical characteristics of the sprawl-fragmentation relationship: it does not prevail, is non-stationary, and scale-dependent. We propose four mechanisms that may have resulted in this mismatch: scale, time-lagged development, spatial arrangement of development, and other external variables including teleconnections. These spatial mismatches provide windows of opportunity for conservation through better development strategies.
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6.

Context

An increasing number of studies have investigated the impact of environmental heterogeneity on faunal assemblages when measured at multiple spatial scales. Few studies, however, have considered how the effects of heterogeneity on fauna vary with the spatial scale at which the response variable is characterised.

Objectives

We investigated the relationship between landscape properties in a region characterised by diverse fire mosaics, and the structure and composition of avian assemblages measured at both the site- (1 ha) and landscape-scale (100 ha).

Methods

We surveyed birds and calculated spatial landscape properties in sub-tropical woodlands of central Queensland, Australia.

Results

Environmental heterogeneity, as measured by topographic complexity, was consistently important for bird species richness and composition. However, the explanatory power of topographic complexity varied depending on the spatial scale and the component of diversity under investigation. We found different correlates of richness within particular foraging guilds depending on the scale at which richness was measured. Extent of long-unburnt habitat (>10 years since fire) was the most important variable for the landscape-scale richness of frugivores, insectivores and canopy feeders, whereas environmental heterogeneity in the surrounding landscape was more important for site-scale richness of these foraging guilds.

Conclusions

The response of species richness to landscape characteristics varies among scales, and among components of diversity. Thus, depending on the scale at which a biodiversity conservation goal is conceptualised—maximising richness at a site, or across a landscape—different landscape management approaches may be preferred.
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7.

Context

A recent hypothesis, the habitat amount hypothesis, predicts that the total amount of habitat in the landscape can replace habitat patch size and isolation in studies of species richness in fragmented landscapes.

Objectives

To test the habitat amount hypothesis by first evaluating at which spatial scale the relationship between species richness in equal-sized sample quadrats and habitat amount was the strongest, and then test the importance of spatial configuration of habitat—measured as local patch size and isolation—when habitat amount was taken into account.

Methods

A quasi-experimental setup with 20 habitat patches of dry calcareous grasslands varying in patch size, patch isolation and habitat amount at the landscape scale was established in the inner Oslo fjord, Southern Norway. We recorded species richness of habitat specialists of vascular plants in equal-sized sample quadrats and analysed the relationship between species richness, habitat amount in the landscape and patch size and isolation.

Results

Although the total amount of habitat in a 3 km-radius around the local patch was positively related to species richness in the sample quadrats, local patch size had an additional positive effect, and the effect of patch size was higher when the amount of habitat within the 3 km-radius was high than when it was low.

Conclusions

In our study system of specialist vascular plants in dry calcareous grasslands, we do not find support for the habitat amount hypothesis.
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8.

Context

Spatial scale is an important consideration for understanding how animals select habitat, and multi-scalar designs in resource selection studies have become increasingly common. Despite this, examination of functional responses in habitat selection at multiple scales is rare. The perceptual range of an animal changes as a function of vegetation association, suggesting that use, selection and functional responses may all be habitat- and scale-dependent.

Objectives

Our objective was to determine how varying grain size affects our interpretation of functional response in habitat selection and to elucidate scalar and landscape effects on habitat selection.

Methods

We quantified the functional response of GPS-collared, female white-tailed deer (Odocoileus virginianus, n = 18) in Riding Mountain National Park, Canada, to different habitat types. Functional responses were quantified at multiple spatial scales by regressing proportion of habitat used against proportion of habitat available at different buffer radii (ranging from 75–1000 m radius) surrounding used (telemetry) locations and available points within the individual’s seasonal home range. We examined how functional responses changed as a function of grain by plotting grain size against the slope of the functional response.

Results

We detected functional responses in most habitat types. As expected, functional responses tended to converge towards 1 (use proportional to availability) at large buffer sizes; however, the relationship between scale and functional response was typically non-linear and depended on habitat type.

Conclusions

We conclude that a multi-scalar approach to modelling animal functional responses in habitat selection is important for understanding patterns in animal behaviour and resource use.
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9.

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, Nterritories = 66, Ncontrols = 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, Nterritories = 78, Ncontrols = 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.
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10.

Context

Landscape metrics represent powerful tools for quantifying landscape structure, but uncertainties persist around their interpretation. Urban settings add unique considerations, containing habitat structures driven by the surrounding built-up environment. Understanding urban ecosystems, however, should focus on the habitats rather than the matrix.

Objectives

We coupled a multivariate approach with landscape metric analysis to overcome existing shortcomings in interpretation. We then explored relationships between landscape characteristics and modelled ecosystem service provision.

Methods

We used principal component analysis and cluster analysis to isolate the most effective measures of landscape variability and then grouped habitat patches according to their attributes, independent of the surrounding urban form. We compared results to the modelled provision of three ecosystem services. Seven classes resulting from cluster analysis were separated primarily on patch area, and secondarily by measures of shape complexity and inter-patch distance.

Results

When compared to modelled ecosystem services, larger patches up to 10 ha in size consistently stored more carbon per area and supported more pollinators, while exhibiting a greater risk of soil erosion. Smaller, isolated patches showed the opposite, and patches larger than 10 ha exhibited no additional areal benefit.

Conclusions

Multivariate landscape metric analysis offers greater confidence and consistency than analysing landscape metrics individually. Independent classification avoids the influence of the urban matrix surrounding habitats of interest, and allows patches to be grouped according to their own attributes. Such a grouping is useful as it may correlate more strongly with the characteristics of landscape structure that directly affect ecosystem function.
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11.

Context

Jack pine (Pinus banksiana)-dominated ecosystems of northern Lower Michigan are the primary breeding habitat for the federally endangered Kirtland’s warbler (Setophaga kirtlandii, KW). Historically, young stands used by KW were produced by stand-replacing wildfires, but fire suppression has necessitated the management of jack pine plantations for KW habitat since the 1970s. Effects of this long-term management on landscape age heterogeneity have previously not been quantified.

Objectives

We hypothesized that forest management has altered the spatial and temporal distribution of jack pine-dominated ecosystems beyond their historic range of variability.

Methods

By developing a diameter-age relationship for jack pine, we estimated ages of pre-European settlement trees found in General Land Office survey notes. We compared pre-European and current landscapes using geostatistical modeling of survey notes, and landscape metrics to quantify changes in pattern.

Results

Three KW management-based age classes (<20, 21–50, >50 years) are now more evenly distributed (31, 39, and 30 %, respectively) compared to the pre-European distribution (5, 19, 76 %) with little variability over time. Landscape metrics suggest the current landscape is younger and more fragmented than the pre-European landscape. These changes indicate restriction of the historic range of age variability, largely due to conversion of older jack pine stands to young KW habitat plantations.

Conclusions

Management has met KW population objectives, but has altered the temporal variability of the landscape’s age structure. Pre-European settlement patterns of stand-ages may provide a foundation for an ecosystem-based management plan for the region that supports both KW and the ecosystems upon which they depend.
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12.

Context

Scale is the lens that focuses ecological relationships. Organisms select habitat at multiple hierarchical levels and at different spatial and/or temporal scales within each level. Failure to properly address scale dependence can result in incorrect inferences in multi-scale habitat selection modeling studies.

Objectives

Our goals in this review are to describe the conceptual origins of multi-scale habitat selection modeling, evaluate the current state-of-the-science, and suggest ways forward to improve analysis of scale-dependent habitat selection.

Methods

We reviewed more than 800 papers on habitat selection from 23 major ecological journals published between 2009 and 2014 and recorded a number of characteristics, such as whether they addressed habitat selection at multiple scales, what attributes of scale were evaluated, and what analytical methods were utilized.

Results

Our results show that despite widespread recognition of the importance of multi-scale analyses of habitat relationships, a large majority of published habitat ecology papers do not address multiple spatial or temporal scales. We also found that scale optimization, which is critical to assess scale dependence, is done in less than 5 % of all habitat selection modeling papers and less than 25 % of papers that address “multi-scale” habitat analysis broadly defined.

Conclusions

Our review confirms the existence of a powerful conceptual foundation for multi-scale habitat selection modeling, but that the majority of studies on wildlife habitat are still not adopting multi-scale frameworks. Most importantly, our review points to the need for wider adoption of a formal scale optimization of organism response to environmental variables.
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13.

Context

The anthropocene is characterised by global landscape modification, and the structure of remnant habitats can explain different patterns of species richness. The most pervasive processes of degradation include habitat loss and fragmentation. However, a recovery of modified landscape is occurring in some areas.

Objectives

The main goal is to know how lichen and bryophyte epiphytic richness growing on Mediterranean forests is influenced not only by fragments characteristics but also by the structure of the landscape. We introduce a temporal dimension in order to evaluate if the historical landscape structure is relevant for current epiphytic communities.

Methods

40 well-preserved forest fragments were selected in a landscape with a large habitat loss over decades, but with a recovery of forest surface in the last 55 years. The most relevant fragment and landscape-scale attributes were considered. Some of the variables were measured in three different years to incorporate a temporal framework.

Results

The results showed that variables at fragment scale had a higher influence, whereas variables at the landscape scale were irrelevant. Among all the historical variables analyzed, only the shift in forest fragment size had influence on species richness.

Conclusions

Mediterranean forests had suffered fragmentation along centuries. Their epiphytic communities also suffer the hard conditions of Mediterranean climate. Our results indicate that Mediterranean epiphytic communities may be in a threshold since it they will never be similar to those communities existing previous fragmentation process even a recovery habitat occur or, they may require more time to response to this habitat recovery.
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14.

Context

How do young birds achieve spatial knowledge about the environment during the initial stages of their life? They may follow adults, so gaining social information and learning; alternatively, young birds may acquire knowledge of the environment themselves by experiencing habitat and landscape features. If learning is at least partially independent of adults then young birds should respond to landscape composition at finer spatial scale than adults, who possess knowledge over a larger area.

Objectives

We studied the responses of juvenile, immature and adult Caspian Gull Larus cachinnans to the same habitat and landscape variables, but at several spatial scales (ranging from 2.5 to 15 km), during post-breeding period.

Methods

We surveyed 61 fish ponds (foraging patches) in southern Poland and counted Caspian gulls.

Results

Juvenile birds responded at finer spatial scales to the factors than did adults. Immature birds showed complicated, intermediate responses to spatial scale. The abundance of juvenile birds was mostly correlated with the landscape composition (positively with the cover of corridors and negatively with barriers). Adult abundance was positively related to foraging patch quality (fish stock), which clearly required previous spatial experience of the environment. The abundance of all age classes were moderately correlated with each other indicating that social behaviour may also contribute to the learning of the environment.

Conclusions

This study shows that as birds mature, they respond differently to components of their environment at different spatial scales. This has considerable ecological consequences for their distribution across environments.
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15.

Context

Understanding the factors contributing to maintaining biodiversity is crucial to mitigate the impact of anthropogenic disturbances. Representing large proportions of green area in highly modified landscapes, residential gardens are often seen as local habitats that can contribute to larger networks of suitable environments at the landscape scale.

Objectives

We investigated the impact of the landscape context on butterfly communities observed in residential gardens, taking into account garden characteristics, land-use types and presence of linear features in the surrounding landscape. We examined how species traits affected butterflies’ response to landscape context and habitat quality.

Methods

We performed a cross-scale study, based on citizen science data documenting butterfly species composition and abundance in 920 gardens across France. We examined the effect of garden quality, the area of different land-use types and the length of linear elements measured at three scales within the surrounding landscape. Species were grouped according to their habitat preference and mobility.

Results

Urbanization negatively affected total species richness and the abundance of butterfly in each group. This was related to declining habitat quality and reduced area of suitable habitat in the surrounding landscape. The magnitude of this effect, however, was negatively correlated with mobility, a trait related to habitat preference. The spatial scale at which landscape context best explained variation in butterfly abundance changed with species’ habitat preference.

Conclusions

This study highlights the importance of preserving high quality habitats in altered landscapes and considering species’ mobility and habitat preference when assessing the impact of landscapes on butterfly communities.
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16.

Context

Emissions of greenhouse gases in urban areas play an important role in climate change. Increasing attention has been given to urban landscape structure–emission relationships (SERs). However, it remains unknown if and to what extent SERs are dependent on observational scale.

Objective

To assess how changing observational scales (in terms of spatial and thematic resolutions) of urban landscape structure affect SERs.

Methods

We examined correlations between 16 landscape metrics and greenhouse gas emissions across 52 European cities, through (1) systematic manipulation of spatial and thematic resolutions of the urban land use/cover (ULUC) dataset, and (2) comparison between available standard ULUC datasets with different spatial resolutions.

Results

Our analyses showed that the observed SERs significantly depend on both thematic and spatial resolutions of the ULUC data. For the 16 landscape metrics, we found diverse spatial/thematic scaling relations exhibiting monotonic, hump-shaped or scale-invariant trends. For different landscape metrics, the SERs were strongest at different spatial scales, suggesting that there is no consistent scaling relation over those observational scales.

Conclusions

SERs are highly sensitive to spatial and thematic resolutions of landscape data. To avoid the problem of ‘ecological fallacy,’ important caveats should be taken for interpretations based on single landscape metrics. Particular consideration should be paid to metrics that are easily interpretable, predictable in scaling behaviors, and important for shaping SERs, such as PLAND, ED, and LPI. Systematic investigations on scaling behaviors of SERs over well-defined scale domains are encouraged in future studies linking greenhouse gas emissions and urban landscape structure.
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17.

Context

Light pollution is a global change affecting a major proportion of global land surface. Although the impacts of Artificial Light At Night (ALAN) have been documented locally for many taxa, the extent of effect of ALAN at a landscape scale on biodiversity is unknown.

Objectives

We characterized the landscape-scale impacts of ALAN on 4 insectivorous bat species Pipistrellus pipistrellus, Pipistrellus kuhlii, Eptesicus serotinus, Nyctalus leisleri, and compared the extent of their effects to other major land-use pressures.

Methods

We used a French national-scale monitoring program recording bat activity among 2-km car transect surveys, and extracted landscape characteristics around transects with satellite and land cover layers. For each species, we performed multi-model averaging at 4 landscape scales (from 200 to 1000 m buffers around transects) to compare the relative effects of the average radiance, the proportion of impervious surface and the proportion of intensive agriculture.

Results

For all species, ALAN had a stronger negative effect than impervious surface at the 4 landscape scales tested. This effect was weaker than the effect of intensive agriculture. The negative effect of ALAN was significant for P. pipistrellus, P. kuhlii and E. serotinus, but not for N. leisleri. The effect of impervious surface varied among species while intensive agriculture had a significant negative effect on the 4 species.

Conclusion

Our results highlight the need to consider the impacts of ALAN on biodiversity in land-use planning and suggest that using only impervious surface as a proxy for urbanization may lead to underestimated impacts on biodiversity.
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18.

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.
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19.

Context

Connectivity models for animal movement frequently use resistance surfaces, but rarely incorporate actual movement data and multiple scale drivers of landscape resistance.

Objectives

Using GPS data, we developed a multi-scale model of landscape resistance for tiger (Panthera tigris) dispersal in central India and evaluated the performance, interpretation and predictions against single scale models.

Methods

Six dispersing tiger paths were subjected to a path level analysis with conditional logistic regression to parameterize a resistance surface. We evaluated for 21 scales of available habitat and selected the best scale for each variable. We derived a scale-optimized multivariate path selection function and predicted landscape resistance across the landscape.

Results

The tigers preferred to move along areas with forest cover at relatively high elevations along the ridges with rugged topography at broad scale, while avoiding areas with agriculture-village matrix at fine scale. We found that the scale that was most supported by Akaike’s information criterion was not always the scale that maximized the magnitude (effect size) of the relationship. Further, the multi-scale optimized model differed substantially from the single scale models in terms of variable importance, magnitude of coefficients and predictions of connectivity.

Conclusions

Our results demonstrate that the variables in landscape resistance models produce markedly different predictions of population connectivity depending on the scales of analyses and interpretation. Thus, scale optimization in parameterization is critical for appropriate inferences and sound management strategies.
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20.

Context

Landscape heterogeneity (the composition and configuration of different landcover types) plays a key role in shaping woodland bird assemblages in wooded-agricultural mosaics. Understanding how species respond to landscape factors could contribute to preventing further decline of woodland bird populations.

Objective

To investigate how woodland birds with different species traits respond to landscape heterogeneity, and to identify whether specific landcover types are important for maintaining diverse populations in wooded-agricultural environments.

Methods

Birds were sampled from woodlands in 58 2 × 2 km tetrads across southern Britain. Landscape heterogeneity was quantified for each tetrad. Bird assemblage response was determined using redundancy analysis combined with variation partitioning and response trait analyses.

Results

For woodland bird assemblages, the independent explanatory importance of landscape composition and landscape configuration variables were closely interrelated. When considered simultaneously during variation partitioning, the community response was better represented by compositional variables. Different species responded to different landscape features and this could be explained by traits relating to woodland association, foraging strata and nest location. Ubiquitous, generalist species, many of which were hole-nesters or ground foragers, correlated positively with urban landcover while specialists of broadleaved woodland avoided landscapes containing urban areas. Species typical of coniferous woodland correlated with large conifer plantations.

Conclusions

At the 2 × 2 km scale, there was evidence that the availability of resources provided by proximate landcover types was highly important for shaping woodland bird assemblages. Further research to disentangle the effects of composition and configuration at different spatial scales is advocated.
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