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1.

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.
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2.

Context

The analysis of individual movement choices can be used to better understand population-level resource selection and inform management.

Objectives

We investigated movements and habitat selection of 13 bobcats in Vermont, USA, under the assumption individuals makes choices based upon their current location. Results were used to identify “movement-defined” corridors.

Methods

We used GPS-collars and GIS to estimate bobcat movement paths, and extracted statistics on land cover proportions, topography, fine-scale vegetation, roads, and streams within “used” and “available” space surrounding each movement path. Compositional analyses were used to determine habitat preferences with respect to landcover and topography; ratio tests were used to determine if used versus available ratios for vegetation, roads, and streams differed from 1. Results were used to create travel cost maps, a primary input for corridor analysis.

Results

Forested and scrub-rock land cover were most preferred for movement, while developed land cover was least preferred. Preference depended on the composition of the “available” landscape: Bobcats moved?>?3 times more quickly through forest and scrub-rock habitat when these habitats were surrounded by agriculture or development than when the available buffer was similarly composed. Overall, forest edge, wetland edge and higher stream densities were selected, while deep forest core and high road densities were not selected. Landscape-scale connectivity maps differed depending on whether habitat suitability, preference, or selection informed the travel cost map.

Conclusions

Both local and landscape scale land cover characteristics affect habitat preferences and travel speed of bobcats, which in turn can inform management and conservation activities.
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3.

Context

Landscape resistance is vital to connectivity modeling and frequently derived from resource selection functions (RSFs). RSFs estimate relative probability of use and tend to focus on understanding habitat preferences during slow, routine animal movements (e.g., foraging). Dispersal and migration, however, can produce rarer, faster movements, in which case models of movement speed rather than resource selection may be more realistic for identifying habitats that facilitate connectivity.

Objective

To compare two connectivity modeling approaches applied to resistance estimated from models of movement rate and resource selection.

Methods

Using movement data from migrating elk, we evaluated continuous time Markov chain (CTMC) and movement-based RSF models (i.e., step selection functions [SSFs]). We applied circuit theory and shortest random path (SRP) algorithms to CTMC, SSF and null (i.e., flat) resistance surfaces to predict corridors between elk seasonal ranges. We evaluated prediction accuracy by comparing model predictions to empirical elk movements.

Results

All connectivity models predicted elk movements well, but models applied to CTMC resistance were more accurate than models applied to SSF and null resistance. Circuit theory models were more accurate on average than SRP models.

Conclusions

CTMC can be more realistic than SSFs for estimating resistance for fast movements, though SSFs may demonstrate some predictive ability when animals also move slowly through corridors (e.g., stopover use during migration). High null model accuracy suggests seasonal range data may also be critical for predicting direct migration routes. For animals that migrate or disperse across large landscapes, we recommend incorporating CTMC into the connectivity modeling toolkit.
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4.

Context

Multispecies and multiscale habitat suitability models (HSM) are important to identify the environmental variables and scales influencing habitat selection and facilitate the comparison of closely related species with different ecological requirements.

Objectives

This study explores the multiscale relationships of habitat suitability for the pine (Martes martes) and stone marten (M. foina) in northern Spain to evaluate differences in habitat selection and scaling, and to determine if there is habitat niche displacement when both species coexist.

Methods

We combined bivariate scaling and maximum entropy modeling to compare the multiscale habitat selection of the two martens. To optimize the HSM, the performance of three sampling bias correction methods at four spatial scales was explored. HSMs were compared to explore niche differentiation between species through a niche identity test.

Results

The comparison among HSMs resulted in the detection of a significant niche divergence between species. The pine marten was positively associated with cooler mountainous areas, low levels of human disturbance, high proportion of natural forests and well-connected forestry plantations, and medium-extent agroforestry mosaics. The stone marten was positively related to the density of urban areas, the proportion and extensiveness of croplands, the existence of some scrub cover and semi-continuous grasslands.

Conclusions

This study outlines the influence of the spatial scale and the importance of the sampling bias corrections in HSM, and to our knowledge, it is the first comparing multiscale habitat selection and niche divergence of two related marten species. This study provides a useful methodological framework for multispecies and multiscale comparatives.
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5.

Context

Although multi-scale approaches are commonly used to assess wildlife-habitat relationships, few studies have examined selection at multiple spatial scales within different hierarchical levels/orders of selection [sensu Johnson’s (1980) orders of selection]. Failure to account for multi-scale relationships within a single level of selection may lead to misleading inferences and predictions.

Objectives

We examined habitat selection of the federally threatened eastern indigo snake (Drymarchon couperi) in peninsular Florida at the level of the home range (Level II selection) and individual telemetry location (Level III selection) to identify influential habitat covariates and predict relative probability of selection.

Methods

Within each level, we identified the characteristic scale for each habitat covariate to create multi-scale resource selection functions. We used home range selection functions to model Level II selection and paired logistic regression to model Level III selection.

Results

At both levels, EIS selected undeveloped upland land covers and habitat edges while avoiding urban land covers. Selection was generally strongest at the finest scales with the exception of Level II urban edge which was avoided at a broad scale indicating avoidance of urbanized land covers rather than urban edge per se.

Conclusions

Our study illustrates how characteristic scales may vary within a single level of selection and demonstrates the utility of multi-level, scale-optimized habitat selection analyses. We emphasize the importance of maintaining large mosaics of natural habitats for eastern indigo snake conservation.
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6.

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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7.

Context

Increased edge density is among the main negative effects of habitat loss and fragmentation. Roads are linear infrastructures that may promote barrier effects due to disturbance and mortality effects. We hypothesized that edges of habitat patches bordered by roads are less permeable than roadless edges.

Objectives

We tested whether edge permeability and avoidance are influenced by the presence of paved and dirt roads bordering habitat patches, relatively to roadless edges.

Methods

We translocated 55 montane akodonts (Akodon montensis) from the interior of vegetation remnants to their edges, and tracked fine-scale movements using spool-and-line devices. Edges were bordered by dirt roads (n = 12 mice), paved roads (n = 21) or were not bordered by roads (n = 22). We assessed edge permeability by comparing the number of tracks with crossings, and by comparing the empirical data to simulated correlated random walks. We also assessed edge avoidance by comparing the net direction travelled and net displacement from edge.

Results

No edge crossings were recorded in roaded edges, whereas 36% of tracks in roadless edges crossed the edge at least once. Simulations indicated a significantly lower permeability of roaded edges, while the observed number of crossings in roadless edges was within the expected range. We found no evidence of higher avoidance of roaded edges, as both net direction travelled and displacement were similar across edge types.

Conclusions

Roads decreased edge permeability for the montane akodont. This is likely to increase population isolation among vegetation remnants by reducing the structural connectivity in the already fragmented landscape.
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8.
9.

Context

Connectivity assessments typically rely on resistance surfaces derived from habitat models, assuming that higher-quality habitat facilitates movement. This assumption remains largely untested though, and it is unlikely that the same environmental factors determine both animal movements and habitat selection, potentially biasing connectivity assessments.

Objectives

We evaluated how much connectivity assessments differ when based on resistance surfaces from habitat versus movement models. In addition, we tested how sensitive connectivity assessments are with respect to the parameterization of the movement models.

Methods

We parameterized maximum entropy models to predict habitat suitability, and step selection functions to derive movement models for brown bear (Ursus arctos) in the northeastern Carpathians. We compared spatial patterns and distributions of resistance values derived from those models, and locations and characteristics of potential movement corridors.

Results

Brown bears preferred areas with high forest cover, close to forest edges, high topographic complexity, and with low human pressure in both habitat and movement models. However, resistance surfaces derived from the habitat models based on predictors measured at broad and medium scales tended to underestimate connectivity, as they predicted substantially higher resistance values for most of the study area, including corridors.

Conclusions

Our findings highlighted that connectivity assessments should be based on movement information if available, rather than generic habitat models. However, the parameterization of movement models is important, because the type of movement events considered, and the sampling method of environmental covariates can greatly affect connectivity assessments, and hence the predicted corridors.
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10.

Context

Transportation and wildlife agencies may consider the need for barrier structures and safe wildlife road-crossings to maintain the long-term viability of wildlife populations. In order to prioritize these efforts, it is important to identify species that are most at risk of extirpation from road-related impacts.

Purpose

Our goal was to identify reptiles and amphibians in California most susceptible to road mortality and fragmentation. With over 160 species and a lack of species-specific research data, we developed an objective risk assessment method based upon road ecology science.

Methods

Risk scoring was based upon a suite of life history and space-use characteristics associated with negative road effects applied in a hierarchical manner from individuals to species. We evaluated risk to both aquatic and terrestrial connectivity and calculated buffer distances to encompass 95% of population-level movements. We ranked species into five relative categories of road-related risk (very-high to very-low) based upon 20% increments of all species scores.

Results

All chelonids, 72% of snakes, 50% of anurans, 18% of lizards and 17% of salamander species in California were ranked at high or very-high risk from negative road impacts. Results were largely consistent with local and global scientific literature in identifying high risk species and groups.

Conclusions

This comparative risk assessment method provides a science-based framework to identify species most susceptible to negative road impacts. The results can inform regional-scale road mitigation planning and prioritization efforts and threat assessments for special-status species. We believe this approach is applicable to numerous landscapes and taxonomic groups.
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11.

Context

Species distributions are a function of an individual’s ability to disperse to and colonize habitat patches. These processes depend upon landscape configuration and composition.

Objectives

Using Blanchard’s cricket frogs (Acris blanchardi), we assessed which land cover types were predictive of (1) presence at three spatial scales (pond-shed, 500 and 2500 m) and (2) genetic structure. We predicted that forested, urban, and road land covers would negatively affect cricket frogs. We also predicted that agricultural, field, and aquatic land covers would positively affect cricket frogs.

Methods

We surveyed for cricket frogs at 28 sites in southwestern Ohio, USA to determine presence across different habitats and analyze genetic structure among populations. For our first objective, we examined if land use (crop, field, forest, and urban habitat) and landscape features (ponds, streams, and roads) explained presence; for our second objective, we assessed whether these land cover types explained genetic distance between populations.

Results

Land cover did not have a strong influence on cricket frog presence. However, multiple competing models suggested effects of roads, streams, and land use. We found genetic structuring: populations were grouped into five major clusters and nine finer-scale clusters. Highways were predictive of increased genetic distance.

Conclusions

By combining a focal-patch study with landscape genetics, our study suggests that major roads and waterways are key features affecting species distributions in agricultural landscapes. We demonstrate that cricket frogs may respond to landscape features at larger spatial scales, and that presence and movement may be affected by different environmental factors.
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12.

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.
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13.

Context

Conservation corridors must facilitate long-distance dispersal movements to promote gene flow, prevent inbreeding, and allow animals to shift ranges with climate change. Least-cost models are used to identify areas that support long-distance movement. These models rely on estimates of landscape resistance, which are typically derived from habitat suitability.

Objectives

We examine two key steps in estimating resistance from habitat suitability: choosing a procedure to estimate habitat suitability, and choosing a transformation function to translate habitat suitability into resistance.

Methods

We used linear and nonlinear functions to convert three types of habitat suitability estimates (from expert opinion, resource selection functions, and step selection functions) into resistances for elk (Cervus canadensis) and desert bighorn sheep (Ovis canadensis nelsoni). We evaluated the resulting resistance maps on an independent set of observed long-distance, prospecting movements.

Results

A negative exponential function best described the relationship between resistance values and habitat suitability for desert bighorn sheep indicating long-distance movers readily travel through moderately-suitable areas and avoid only the least suitable habitat. For desert bighorn sheep, all three suitability estimates performed better than chance, and resource and step selection functions outperformed expert opinion. For elk, all three suitability estimates performed the same as chance.

Conclusions

When designing corridors to facilitate long-distance movements of mobile animals, we recommend transforming habitat suitability into resistance with a negative exponential function. Use of an exponential transformation means that larger fractions of the landscape offer low resistance, allowing greater flexibility in where a corridor is located.
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14.

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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15.

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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16.

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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17.

Context

In heterogeneous landscapes, local patterns of community structure are a product of the habitat size and condition within a patch interacting with adjacent habitat patches of varying composition and quantity. While evidence for local versus landscape factors have been found in terrestrial biomes, support for such multi-scale effects shaping marine ecological communities is equivocal.

Objectives

We investigated whether within-patch habitat condition can override seascape context to explain the community structure of macroalgae-associated reef fishes across a tropical seascape.

Methods

We mapped the distribution and abundance of a diverse family of reef fishes (Labridae) occupying macroalgae meadows within a tropical reef ecosystem, and using best-subsets model selection, investigated the potential for habitat structural connectivity and/or local habitat quality for predicting variations in fish community structure across the seascape.

Results

Local habitat quality (canopy structure, hard habitat complexity) and area of coral-dominated habitat within 500 m of a macroalgal meadow provided the best predictors of fish community structure. However, the specific importance of a given predictor varied with fish life history stage and functional trophic group. Interestingly, macroalgae meadow area was among the least important predictors.

Conclusions

Given the complex interplay between local habitat quality and spatial context effects on fish biodiversity, our study reveals the multi-scale predictors that should be used in spatial conservation and management approaches for tropical fish diversity. Moreover, our findings question the ubiquity of habitat area effects in patchy landscapes, and cautions against a sole reliance on habitat quantity in spatial management.
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18.

Context

Habitat loss and habitat fragmentation negatively affect amphibian populations. Roads impact amphibian species through barrier effects and traffic mortality. The landscape variable ‘accessible habitat’ considers the combined effects of habitat loss and roads on populations.

Objectives

The aim was to test whether accessible habitat was a better predictor of amphibian species richness than separate measures of road effects and habitat loss. I assessed how accessible habitat and local habitat variables determine species richness and community composition.

Methods

Frog and tadpole surveys were conducted at 52 wetlands in a peri-urban area of eastern Australia. Accessible habitat was delineated using a highway. Regressions were used to examine relationships between species richness and eleven landscape and local habitat variables. Redundancy analysis was used to examine relationships between community composition and accessible habitat and local habitat variables.

Results

Best-ranked models of species richness included both landscape and local habitat variables. There were positive relationships between species richness and accessible habitat and distance to the highway, and uncertain relationships with proportion cover of native vegetation and road density. There were negative relationships between species richness and concreted wetlands and wetland electrical conductivity. Four species were positively associated with accessible habitat, whereas all species were negatively associated with wetland type.

Conclusions

Barrier effects caused by the highway and habitat loss have negatively affected the amphibian community. Local habitat variables had strong relationships with species richness and community composition, highlighting the importance of both availability and quality of habitat for amphibian conservation near major roads.
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19.

Context

Although animal movement behaviors are influenced by spatial heterogeneity, such behaviors can also generate spatial heterogeneity via interactions with the emergent spatial structure and other individuals (i.e., the social landscape).

Objective

Elucidate the behavioral and ecological mechanisms of pattern formation in a homogeneous resource landscape.

Methods

We analyzed the movement pathways and space-use patterns of the lesser grain borer (Rhyzopertha dominica) within homogeneous resource landscapes (wheat kernels). Experimental trials consisted of individual beetles foraging alone or paired with a member of the same or different sex.

Results

We identified two sources of pattern formation: (1) beetles were attracted to areas where they or another beetle had previously fed, leading to increased patchiness via positive reinforcement; and (2) the presence of conspecifics affected whether and at what scales patchiness occurred. Solitary males had lower rates of movement and less tortuous pathways than solitary females, but both sexes generated fine-scale patchiness in the resource distribution. Patchy resource landscapes were also generated by male–female pairs, but not by same-sex pairings. Paired females in particular exhibited significantly greater daily net displacements and more random space use than solitary females.

Conclusions

Pattern formation is a complex process, even in a relatively simple, homogeneous resource landscape. In particular, patterns created by individuals when foraging alone versus in pairs underscores how social interactions can fundamentally alter the resultant pattern of heterogeneity that emerges in resource landscapes.
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20.

Context

Routine movements of large herbivores, often considered as ecosystem engineers, impact key ecological processes. Functional landscape connectivity for such species influences the spatial distribution of associated ecological services and disservices.

Objectives

We studied how spatio-temporal variation in the risk-resource trade-off, generated by fluctuations in human activities and environmental conditions, influences the routine movements of roe deer across a heterogeneous landscape, generating shifts in functional connectivity at daily and seasonal time scales.

Methods

We used GPS locations of 172 adult roe deer and step selection functions to infer landscape connectivity. In particular, we assessed the influence of six habitat features on fine scale movements across four biological seasons and three daily periods, based on variations in the risk-resource trade-off.

Results

The influence of habitat features on roe deer movements was strongly dependent on proximity to refuge habitat, i.e. woodlands. Roe deer confined their movements to safe habitats during daytime and during the hunting season, when human activity is high. However, they exploited exposed open habitats more freely during night-time. Consequently, we observed marked temporal shifts in landscape connectivity, which was highest at night in summer and lowest during daytime in autumn. In particular, the onset of the autumn hunting season induced an abrupt decrease in landscape connectivity.

Conclusions

Human disturbance had a strong impact on roe deer movements, generating pronounced spatio-temporal variation in landscape connectivity. However, high connectivity at night across all seasons implies that Europe’s most abundant and widespread large herbivore potentially plays a key role in transporting ticks, seeds and nutrients among habitats.
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