Context

In southwestern Alberta, human development, including roads, is encroaching on the landscape and into the range of a partially migratory population of elk (Cervus elaphus).

Objectives

To quantify factors influencing among- and within-home-range selection of winter range in this population.

Methods

We studied individual habitat selection and road avoidance at two biologically relevant spatial scales. We outlined availability extents for 107 individual elk-years based on observed fall migration distance, and based on a minimum convex polygon around winter telemetry relocations. To model the response by elk to road disturbance, we fit a habitat-selection model to each elk-year at each of the two availability extents, and examined population-level and individual variation in space-use. We then evaluated the relationship between inferred selection at the two scales and the functional response in selection.

Results

Roads had a ubiquitous influence on elk across scales. Elk, individually and as a population, avoided roads when migrating to their winter range and within this seasonal home range. Individual elk that avoided roads more strongly relative to the population did so at both scales of analysis.Further, the avoidance of low-use roads decreased with increasing road density. These results support bottom-up habitat-selection patterns (i.e., scale-independent) and functional response in habitat selection.

Conclusions

Overall, using a multi-scale habitat selection analysis, we show that road avoidance is a major determinant of elk space-use behaviour across multiple scales. Consequently, any new road construction or increases in road-use intensity could have detrimental effects on migratory elk populations by restricting space-use.

Context

Landscape and habitat filters are major drivers of biodiversity of small habitat islands by influencing dispersal and extinction events in plant metapopulations.

Objectives

We assessed the effects of landscape and habitat filters on the species richness, abundance and trait composition of grassland specialist and generalist plants in small habitat islands. We studied traits related to functional spatial connectivity (dispersal ability by wind and animals) and temporal connectivity (clonality and seed bank persistence) using model selection.

Methods

We sampled herbaceous plants, landscape (local and regional isolation) and habitat filters (inclination, woody encroachment and disturbance) in 82 grassland islands in Hungary.

Results

Isolation decreased the abundance of good disperser specialist plants due to the lack of directional vectors transferring seeds between suitable habitat patches. Clonality was an effective strategy, but persistent seed bank did not support the survival of specialist plants in isolated habitats. Generalist plants were unaffected by landscape filters due to their wide habitat breadth and high propagule availability. Clonal specialist plants could cope with increasing woody encroachment due to their high resistance against environmental changes; however, they could not cope with intensive disturbance. Steep slopes providing environmental heterogeneity had an overall positive effect on species richness.

Conclusions

Specialist plants were influenced by the interplay of landscape filters influencing their abundance and habitat filters affecting species richness. Landscape filtering by isolation influenced the abundance of specialist plants by regulating seed dispersal. Habitat filters sorted species that could establish and persist at a site by influencing microsite availability and quality.

Context

Scale dependence of bat habitat selection is poorly known with few studies evaluating relationships among landscape metrics such as class versus landscape, or metrics that measure composition or configuration. This knowledge can inform conservation approaches to mitigate habitat loss and fragmentation.

Objectives

We evaluated scale dependence of habitat associations and scaling patterns of landscape metrics in relation to bat occurrence or capture rate in forests of southwestern Nicaragua.

Methods

We captured 1537 bats at 35 locations and measured landscape and class metrics across 10 spatial scales (100–1000 m) surrounding capture locations. We conducted univariate scaling across the 10 scales and identified scales and variables most related to bat occurrence or capture rate.

Results

Edge and patch density, at both landscape and class levels, were the most important variables across species. Feeding guilds varied in their response to metrics. Certain landscape and configuration metrics were most influential at fine (100 m) and/or broad (1000 m) spatial scales while most class and composition metrics were influential at intermediate scales.

Conclusions

These results provide insight into the scale dependence of habitat associations of bat species and the influence of fine and broad scales on habitat associations. The effects of scale, examined in our study and others from fine (100 m) to broad (5 km) indicate habitat relationships for bats may be more informative at larger scales. Our results suggest there could be general differences in scale relationships for different groups of landscape metrics, which deserves further evaluation in other taxonomic groups.

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.

Context

Habitat loss and fragmentation are among the major drivers of population declines and extinction, particularly in large carnivores. Connectivity models provide practical tools for assessing fragmentation effects and developing mitigation or conservation responses. To be useful to conservation practitioners, connectivity models need to incorporate multiple scales and include realistic scenarios based on potential changes to habitat and anthropogenic pressures. This will help to prioritize conservation efforts in a changing landscape.

Objectives

The goal of our paper was to evaluate differences in population connectivity for lions (Panthera leo) across the Kavango-Zambezi Trans-frontier Conservation Area (KAZA) under different landscape change scenarios and a range of dispersal distances.

Methods

We used an empirically optimized resistance surface, based on analysis of movement pathways of dispersing lions in southern Africa to calculate resistant kernel connectivity. We assessed changes in connectivity across nine landscape change scenarios, under each of which we explored the behavior of lions with eight different dispersal abilities.

Results

Our results demonstrate that reductions in the extent of the protected area network and/or fencing protected areas will result in large declines in the extent of population connectivity, across all modeled dispersal abilities. Creation of corridors or erection of fences strategically placed to funnel dispersers between protected areas increased overall connectivity of the population.

Conclusions

Our results strongly suggest that the most effective means of maintaining long-term population connectivity of lions in the KAZA region involves retaining the current protected area network, augmented with protected corridors or strategic fencing to direct dispersing individuals towards suitable habitat and away from potential conflict areas.

Context

Native vegetation is often used as a proxy for habitat to estimate habitat availability in landscapes. This approach may lead to incorrect estimates of the impacts of habitat loss and fragmentation on species, which have not been thoroughly quantified so far.

Objectives

We quantified to what extent the loss of native vegetation reflect actual habitat loss by native species in landscapes. We tested the hypothesis that habitat availability declines at greater rates than native vegetation and thus is overestimated when it is quantified on the basis of native vegetation.

Methods

Using simulations, we quantified how the loss of native vegetation in artificial and real landscapes affects habitat availability for species with different habitat requirements. We contrasted a generalist species, which uses all native vegetation, with 10 habitat-specialist species classified into three categories (interior, patchy and riparian species).

Results

Habitat availability generally declined at greater rates than native vegetation for all specialist species. This pattern was apparent for different specialist species in a broad range of landscape types. Interior species always lost habitat availability more rapidly than the generalist species. Most riparian species lost habitat availability more rapidly than the generalist species. Responses of patchy species were more complex, depending on their dispersal abilities and landscape structure.

Conclusions

Habitat availability is likely to be overestimated when native vegetation is used as proxy for habitat, because habitat availability will generally decline at greater rates than native vegetation. Therefore, a species-centered approach should be adopted when estimating habitat availability in landscapes.

Context

Management of wintering waterfowl in North America requires adaptability because constant landscape and environmental change challenges existing management strategies regarding waterfowl habitat use at large spatial scales. Migratory waterfowl including mallards (Anas platyrhynchos) use the lower Mississippi Alluvial Valley (MAV) for wintering habitat, making this an important area of emphasis for improving wetland conservation strategies, while enhancing the understanding of landscape-use patterns.

Objectives

We used aerial survey data collected in the Arkansas portion of the MAV (ARMAV) to explain the abundance and distribution of mallards in relation to variable landscape conditions.

Methods

We used two-stage, hierarchical spatio-temporal models with a random spatial effect to identify covariates related to changes in mallard abundance and distribution within and among years.

Results

We found distinct spatio-temporal patterns existed for mallard distributions across the ARMAV and these distributions are dependent on the surrounding landscape structure and changing environmental conditions. Models performing best indicated seasonal surface water extent, rice field, wetland and fallow (uncultivated) fields positively influenced mallard presence. Rice fields, surface water and weather were found to influence mallard abundance. Additionally, the results suggest weather and changing surface water affects mallard presence and abundance throughout the winter.

Conclusions

Using novel datasets to identify which environmental factors drive changes in regional wildlife distribution and abundance can improve management by providing managers additional information to manage land over landscapes spanning private and public lands. We suggest our analytical approach may be informative in other areas and for other wildlife species.

Context

Allometric scaling laws are foundational to structuring processes from cellular to ecosystem levels. The idea that allometric relationships underlie species characteristic selection scales, the spatial scales at which species respond to landscape features, has recently been investigated, however, supporting empirical evidence is scarce.

Objectives

Lack of pattern can be explained by inaccurate estimation, low power, confounding factors, or absence of a relationship. In this paper, we evaluate the relationship between body size and species characteristic selection scales after overcoming limitations of previous study designs.

Methods

We conducted 1328 avian point counts across the state of Nebraska using the robust sampling design to account for imperfect detection. We used Bayesian latent indicator scale selection with N-mixture models to estimate species’ characteristic selection scales of six habitat features for 86 species. We propagated the uncertainty associated with assigning characteristic scales to a model of the relationship between body size and characteristic spatial scales.

Results

Species characteristic scales varied across habitat predictors, and varied in the uncertainty associated with selecting single characteristic scales. After propagating uncertainty our results do not support a relationship between species’ body size and the spatial scales at which they respond to landscape features.

Conclusions

As species abundance integrates birth, death, immigration, and emigration processes, each of which are influenced by ecological processes manifesting at various scales, we question whether a general allometric relationship should be expected. Our results suggest that selection may act on responses to specific environmental features, rather than responses to spatial scale per se.

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.

Context

Ungulate browsers often alter plant composition and reduce diversity in forests worldwide, yet our ability to predict browse impact on vegetation remains equivocal. Theory suggests, however, that ungulate distribution and foraging impacts are shaped by scale-dependent decisions based on variation in habitat composition and structure encountered within their home range.

Objective

Examine how variation in habitat composition at landscape (259 ha) scales modulates browse impact on vegetation at local scales.

Methods

We measured vegetation richness and abundance in plots with and without white-tailed deer (Odocoileus virginianus) at 23 northern hardwood forest sites distributed across a 6500 km² area in Pennsylvania, USA. Experimental sites were embedded within landscapes with varying levels of habitat composition and deer densities.

Results

Browsing reduced vegetation richness and cover by as much as 53 and 70%, respectively; however, we found browse impact was modulated by variation in the relative abundance of managed habitats that alter forage availability. Specifically, relative to fenced areas, browse impact weakened and ultimately disappeared as the proportion of forage-rich habitats (e.g., recent harvests) increased to ≥20%. Conversely, vegetation grew increasingly depauperate as landscapes contained greater proportions of forage-poor habitats (i.e., older harvests), particularly when browsed.

Conclusions

Our results underscore how management actions that alter forage availability to ungulates throughout the landscape (i.e. the foodscape) can shape forest-ungulate interactions and suggest a new paradigm whereby managers evaluate and undertake actions at the appropriate spatio-temporal scales to proactively limit the deleterious impact of browsing on plant biodiversity.

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.

Context

Beyond the recognized importance of protecting large areas of contiguous habitat, conservation efforts for many species are complicated by the fact that patch suitability may also be affected by characteristics of the landscape within which the patch is located. Currently, little is known about the spatial scales at which species respond to different aspects of the landscape surrounding an occupied patch.

Objectives

Using grassland bird point count data, we describe an approach to evaluating scale-specific effects of landscape composition on patch occupancy.

Methods

We used data from 793 point count surveys conducted in idle and grazed grasslands across Wisconsin, USA from 2012 to 2014 to evaluate scale-dependencies in the response of grassland birds to landscape composition. Patch occupancy models were used to evaluate the relationship between occupancy and landscape composition at scales from 100 to 3000 m.

Results

Bobolink (Dolichonyx oryzivorus) exhibited a pattern indicating selection for grassland habitats in the surrounding landscape at all spatial scales while selecting against other habitats. Eastern Meadowlark (Sturnella magna) displayed evidence of scale sensitivity for all habitat types. Grasshopper Sparrow (Ammodramus savannarum) showed a strong positive response to pasture and idle grass at all scales and negatively to cropland at large scales. Unlike other species, patch occupancy by Henslow’s Sparrow (A. henslowii) was primarily influenced by patch area.

Conclusions

Our results suggest that both working grasslands (pasture) and idle conservation grasslands can play an important role in grassland bird conservation but also highlight the importance of considering species-specific patch and landscape characteristics for effective conservation.

Context

A challenge devising revegetation strategies in fragmented landscapes is conserving for the widest spectrum of biodiversity. Habitat network reconstruction should improve landscape capacity to maintain species populations. However, the location of revegetation often fails to account for species occurrence and dispersal processes operating across spatial scales.

Objectives

Our objective was to integrate metapopulation theory with estimates of landscape capacity and dispersal pathways to highlight connectivity gaps. Maintenance of populations could thereby be facilitated through reconnecting habitat networks across regional and broader scales, with assumed benefit for the dispersal needs of less sensitive species.

Methods

Predicted occupancy and metapopulation capacity were calculated for a generic focal species derived from fragmentation-sensitive woodland birds, mammals and reptiles. A metapopulation connectivity analysis predicted regional dispersal links to identify likely routes through which individuals may move to contribute to the viability of the population. We used the revegetation programmes of the Brigalow–Nandewar Biolinks project, eastern New South Wales, Australia, to demonstrate our approach.

Results

Landscape capacity of the current landscape varied across the region. Low-value links between populations provided greatest opportunities for revegetation and improved landscape capacity. Where regional connectivity did not indicate a pathway between populations, broader scale connectivity provided guidance for revegetation.

Conclusions

The metapopulation-based model, coupled with a habitat dispersal network analysis, provided a platform to inform revegetation locations and better support biodiversity. Our approach has application for directing on-ground action to support viable populations, assess the impact of revegetation schemes or monitor the progress of staged implementations.

Context

Multi-scale approaches to habitat modeling have been shown to provide more accurate understanding and predictions of species-habitat associations. It remains however unexplored how spatial and temporal variations in habitat use may affect multi-scale habitat modeling.

Objectives

We aimed at assessing how seasonal and temporal differences in species habitat use and distribution impact operational scales, variable influence, habitat suitability spatial patterns, and performance of multi-scale models.

Methods

We evaluated the environmental factors driving brown bear habitat relationships in the Cantabrian Range (Spain) based on species presence records (ground observations) for the period 2000–2010, LiDAR data on forest structure, and seasonal estimates of foraging resources. We separately developed multi-scale habitat models for (i) each season (spring, summer, fall and winter) (ii) two sub-periods with different population status: 2000–2004 (with brown bear distribution restricted to the main population nuclei) and 2005–2010 (with expanding bear population and range); and (iii) the entire 2000–2010 period.

Results

Scales of effect remained considerably stable across seasonal and temporal variations, but not the influence of certain environmental variables. The predictive ability of multi-scale models was lower in the seasons or periods in which populations used larger areas and a broader variety of environmental conditions. Seasonal estimates of foraging resources, together with LiDAR data, appeared to improve the performance of multi-scale habitat models.

Conclusions

We highlight that the understanding of multi-scale behavioral responses of species to spatial patterns that continually shift over time may be essential to unravel habitat relationships and produce reliable estimates of species distributions.

Context

Understanding habitat selection can be challenging for species surviving in small populations, but is needed for landscape-scale conservation planning.

Objectives

We assessed how European bison (Bison bonasus) habitat selection, and particularly forest use, varies across subpopulations and spatial scales.

Methods

We gathered the most comprehensive European bison occurrence dataset to date, from five free-ranging herds in Poland. We compared these data to a high-resolution forest map and modelled the influence of environmental and human-pressure variables on habitat selection.

Results

Around 65% of European bison occurrences were in forests, with cows showing a slightly higher forest association than bulls. Forest association did not change markedly across spatial scales, yet differed strongly among herds. Modelling European bison habitat suitability confirmed forest preference, but also showed strong differences in habitat selection among herds. Some herds used open areas heavily and actively selected for them. Similarly, human-pressure variables were important in all herds, but some herds avoided human-dominated areas more than others.

Conclusions

Assessing European bison habitat across multiple herds revealed a more generalist habitat use pattern than when studying individual herds only. Our results highlight that conflicts with land use and people could be substantial if bison are released in human-dominated landscapes. Future restoration efforts should target areas with low road and human population density, regardless of the degree of forest cover. More broadly, our study highlights the importance of considering multiple subpopulations and spatial scales in conservation planning.

Context

Theory predicts that habitat loss and fragmentation may have drastic consequences on species’ interactions. To date, however, little empirical evidence exists on the strength of interspecific competition in shaping animal communities in fragmented landscapes.

Objectives

Our aim was to measure the degree of ongoing competitive interference between species in fragmented landscapes. Our model system was the community of ground-dwelling rodents in deciduous woodlands in central Italy, composed of a habitat generalist species (Apodemus sylvaticus) and two forest specialists (Apodemus flavicollis and Myodes glareolus). Our objectives were to test whether species were segregated among forest patches and whether spatial segregation was determined by interspecific competition or habitat and resource availability.

Methods

We surveyed the populations inhabiting 29 woodland patches in a highly fragmented landscape using a capture-mark-recapture protocol, capturing >4500 individuals. First we modelled species’ distribution as a function of habitat, resource availability and landscape variables. The second stage of our analyses involved measuring the response of vital rate parameters (body mass, reproduction, survival, recruitment, population density) to competitor density.

Results

The relative distribution of species reflected a spatial segregation of habitat generalists and specialists according to habitat quality, cover and connectivity. Interspecific competition mainly affected individual level vital rates, whereas we found no substantial effects at the population level.

Conclusions

Competitive exclusion of specialist species by generalist species was occurring. However, when compared to other factors such as habitat connectivity and resource availability, interspecific competition played a relatively minor role in shaping the studied community.

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.

Context

Animal movements are inherently linked to landscape structure. Understanding this relationship for highly-mobile species requires documenting their responses to spatiotemporal variability of resources. To that end, characterizing movement behaviors and resource distributions using the principles of habitat connectivity facilitates coordinated landscape planning efforts within highly modified landscapes.

Objectives and methods

We tracked locations and movements for 156 dunlin (Calidris alpina) and 109 long-billed dowitchers (Limnodromus scolopaceus) overwintering in two regions with distinct water distributions in California’s Central Valley. We then compared residency rates, functional connectivity to other regions, and associations between movement distances and average habitat availability and structural connectivity of habitat at multiple temporal and spatial scales.

Results

A widespread yet highly variable regional water distribution was associated with lower residency rates and substantially higher functional connectivity to nearby regions when compared to a stable regional water distribution characterized by a large, contiguous wetland complex. Longer movements were associated with decreasing average availability and spatial aggregation of surface water. Movement models suggested shorebirds primarily responded to habitat availability at smaller scales (<?10 km) and structural connectivity at larger scales (≥?10 km).

Conclusions

Differences in movement behaviors suggested that wintering shorebirds will avoid long distance movements and remain resident within a wetland region when possible. Conservation and management efforts should reliably flood individual wetlands and agricultural lands from November to April and prioritize locations that maximize structural wetland connectivity and limit spatiotemporal variability of surface water throughout the Central Valley.

Context

The conversion of natural environments into agricultural land has profound effects on the composition of the landscape, often resulting in a mosaic of human-altered and natural habitats. The response to these changes may however vary among organisms. Bats are highly vagile, and their requirements often imply the use of distinct habitats, which they select responding to both landscape and local features.

Objectives

We aimed to identify which features influence bat richness and activity within Baixo Vouga Lagunar, a heterogeneous landscape located on the Central-North Portuguese coast, and to investigate if that influence varies across a gradient of focal scales.

Methods

We sampled bats acoustically, while simultaneously sampling insects with light traps. We assessed the relationships between species richness, bat activity, and activity of eco-morphological guilds with landscape and local features, across four scales.

Results

Our results revealed both scale- and guild-dependent responses of bats to landscape and local features. At broader scales we found positive associations between open-space foraging bats and habitat heterogeneity and between edge-space foraging bats and greater edge lengths. Woodland cover and water availability at an intermediate scale and weather conditions and insect abundance at a local scale were the factors that mostly influenced the response variables.

Conclusions

Globally, our results suggest that bats are sensitive to local resource availability and distribution, while simultaneously reacting to landscape features acting at coarser scales. Finally, our results suggest that the responses given by bats are guild-dependent, and some habitats act as keystone structures for bats within this mosaic.