One size does not fit all: European bison habitat selection across herds and spatial scales

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.

     

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

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

Linking habitat suitability and road mortalities across geographic ranges

Protected areas are established to conserve biodiversity and facilitate resilience to threatening processes. Yet protected areas are not isolated environmental compounds. Many threats breach their borders, including transportation infrastructure. Despite an abundance of roads in many protected areas, the impact of roads on biota within these protected areas is usually unaccounted for in threat mitigation efforts. As landscapes become further developed and the importance of protected areas increases, knowledge of how roads impact on the persistence of species at large scales and whether protected areas provide relief from this process is vital. We took a two-staged approach to analysing landscape-scale habitat use and road-kill impacts of the common wombat (Vombatus ursinus), a large, widely distributed herbivore, within New South Wales (NSW), Australia. Firstly, we modelled their state-wide distribution from atlas records and evaluated the relationship between habitat suitability and wombat road fatalities at that scale. Secondly, we used local-scale fatality data to derive an annual estimate of wombats killed within an optimal habitat area. We then combined these two approaches to derive a measure of total wombats killed on roads within the protected area network. Our results showed that common wombats have a broad distribution (290,981?km²), one quarter (24.9?%) of their distribution lies within protected areas, and the percentage of optimal habitat contained within protected areas is 35.6?%, far greater than the COP10 guidelines of 17?%. Problematically, optimal habitat within protected areas was not a barrier to the effects of road-kill, as we estimated that the total annual count of wombat road-kill in optimal habitat within protected areas could be as high as 13.6?% of the total NSW population. These findings suggest that although protected areas are important spatial refuges for biodiversity, greater effort should be made to evaluate how reserves confer resilience from the impacts of roads across geographic ranges.     

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

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

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

Context

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

Objective

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

Methods

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

Results

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

Conclusions

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

     

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

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

Landscape composition influences roe deer habitat selection at both home range and landscape scales

Understanding how patterns of habitat selection vary in relation to landscape structure is essential to predict ecological responses of species to global change and inform management. We investigated behavioural plasticity in habitat selection of roe deer (Capreolus capreolus) in relation to variable habitat availability across a heterogeneous agricultural landscape at the home range and landscape scales. As expected, woodland was heavily selected, but we found no functional response for this habitat, i.e. no shift in habitat selection with changing habitat availability, possibly due to the presence of hedgerows which were increasingly selected as woodlands were less abundant. Hedgerows may thus function as a substitutable habitat for woodlands by providing roe deer with similar resources. We observed a functional response in the use of hedgerows, implying some degree of landscape complementation between hedgerows and open habitats, which may in part compensate for lower woodland availability. We also expected selection for woodland to be highest at the wider spatial scale, especially when this habitat was limiting. However, our results did not support this hypothesis, but rather indicated a marked influence of habitat composition, as both the availability and distribution of resources conditioned habitat selection. There was no marked between-sex difference in the pattern of habitat selection at either scale or between seasons at the landscape scale, however, within the home range, selection did differ between seasons. We conclude that landscape structure has a marked impact on roe deer habitat selection in agricultural landscapes through processes such as landscape complementation and supplementation.     

Multi-scale habitat selection modeling: a review and outlook

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.

     

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

Context

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

Objectives

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

Methods

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

Results

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

Conclusions

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

     

Predicting across scales: Theory development and testing

Landscape ecologists deal with processes that occur at a variety of temporal and spatial scales. The ability to make predictions at more than one level of resolution requires identification of the processes of interest and parameters that affect this process at different scales, the development of rules to translate information across scales, and the ability to test these predictions at the relevant spatial and temporal scales. This paper synthesizes discussions from a workshop on Predicting Across Scales: Theory Development and Testing that was held to discuss current research on scaling and to identify key research issues.     

Grain-dependent functional responses in habitat selection

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.

     

Adaptive models for large herbivore movements in heterogeneous landscapes

Enzymes are protein catalysts of extraordinary efficiency, capable of bringing about rate enhancements of their biochemical reactions that can approach factors of 10²⁰. Theories of enzyme catalysis, which seek to explain the means by which enzymes effect catalytic transformation of the substrate molecules on which they work, have evolved over the past century from the “lock-and-key” model proposed by Emil Fischer in 1894 to models that explicitly rely on transition state theory to the most recent theories that strive to provide accounts that stress the essential role of protein dynamics. In this paper, I attempt to construct a metaphysical framework within which these new models of enzyme catalysis can be developed. This framework is constructed from key doctrines of process thought, which gives ontologic priority to becoming over being, as well as tenets of a process philosophy of chemistry, which stresses environmentally-responsive molecular transformation. Enzyme catalysis can now be seen not as enzyme acting on its substrate, but rather as enzyme and substrate entering into a relation which allows them to traverse the reaction coordinate as an ontologic unity x̄.     

Factors affecting intraspecific variation in home range size of a large African herbivore

Factors affecting intraspecific variation in home range size have rarely been examined using modern statistical and remote sensing methods. This is especially true for animals in seasonal savanna environments in Africa, despite this biome??s importance for both conservation and development goals. We studied the impacts of spatial and temporal variability in environmental conditions, along with individual and social factors, on home range sizes in African buffalo (Syncerus caffer) in northeastern Namibia. Our data set spans 4?years, is derived from 32 satellite tracking collars, and contains over 35,000 GPS locations. We used the local convex hull method to estimate home range size from 31 buffalo captured at 6 sites. We used a variety of remotely sensed data to characterize potential anthropogenic and natural boundaries, as well as seasonal and temporal heterogeneity in environmental conditions. Using an information-theoretic, mixed effects approach, our analyses showed that home ranges varied over two orders of magnitude and are among the largest recorded for this species. Variables relating to vegetation and habitat boundaries were more important than abiotic environmental conditions and individual or social factors in explaining variation in home range size. The relative contributions of environmental, individual, social, and linear boundary variables to intraspecific home range size have rarely been examined and prior to this had not been assessed for any species in seasonal savannas of Africa. Understanding the factors that condition space-use patterns of wildlife in this area will lead to better-informed conservation and sustainable development decisions.     

Integrating landscape resistance and multi-scale predictor of habitat selection for amphibian distribution modelling at large scale

Context

Species distribution modelling is a common tool in conservation biology but two main criticisms remain: (1) the use of simplistic variables that do not account for species movements and/or connectivity and (2) poor consideration of multi-scale processes driving species distributions.

Objectives

We aimed to determine if including multi-scale and fine-scale movement processes in SDM predictors would improve accuracy of SDM for low-mobility amphibian species compared with species-level analysis.

Methods

We tested and compared different SDMs for nine amphibian species with four different sets of predictors: (1) simple distance-based predictors; (2) single-scale compositional predictors; (3) multi-scale compositional predictors with a priori selection of scale based on knowledge of species mobility and scale-of-effect; and (4) multi-scale compositional predictors calculated using a friction-based functional grain to account for resource accessibility with landscape resistance to movement.

Results

Using friction-based functional grain predictors produced slight to moderate improvements of SDM performance at large scale. The multi-scale approach, with a priori scale selection, led to ambiguous results depending on the species studied, in particular for generalist species.

Conclusion

We underline the potential of using a friction-based functional grain to improve SDM predictions for species-level analysis.

     

Surface metrics for landscape ecology: a comparison of landscape models across ecoregions and scales

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.

     

Resource depletion versus landscape complementation: habitat selection by a multiple central place forager

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.

     

Human settlement ecology and chimpanzee habitat selection in Mali

Customary land-use practices create distinctive cultural landscapes, including landscapes where abandoned settlements host vegetation that attracts wild animals. Understanding how landscape patterns relate to land-use history can help clarify the ecological effects of particular land uses. This study examines relationships between chimpanzee habitat selection and Maninka settlement practice, to determine how settlement history has affected chimpanzee habitat in Mali’s Bafing Biosphere Reserve, where conservation practitioners assume that the characteristic settlement pattern reflects a process of settlement expansion into undisturbed habitat. Three types of data are reported: (1) ethnographic interviews on settlement history and practice; (2) systematic sampling of chimpanzee habitat use; and (3) ground-based mapping of settlement sites, surface water, and fruit-tree patches. These data show that the Maninka have a shifting settlement system, meaning that most sites are occupied for only relatively brief periods; and that some abandoned settlement sites host fruit-tree patches that are seasonally important chimpanzee habitat. Two main conclusions are: (1) settlement expansion has not occurred; instead, historic settlement has created habitat that is both attractive and available to chimpanzees. Anthropogenic habitat does not appear to be vital for chimpanzee survival, but it spatially and temporally supplements natural habitats. (2) Conservation policies meant to reduce the presumed threat of settlement expansion may have initiated an unintended, long-term threat of habitat loss for chimpanzees. While settlement practices may be a component of short-term threats to chimpanzees, such as hunting, when addressing these threats conservation practitioners should consider long-term settlement processes to avoid creating new threats.     

Multi-level,multi-scale habitat selection by a wide-ranging,federally threatened snake

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.