Functional Responses In Habitat Selection Research Articles

Uniting statistical and individual-based approaches for animal movement modelling.

The dynamic nature of their internal states and the environment directly shape animals' spatial behaviours and give rise to emergent properties at broader scales in natural systems. However, integrating these dynamic features into habitat selection studies remains challenging, due to practically impossible field work to access internal states and the inability of current statistical models to produce dynamic outputs. To address these issues, we developed a robust method, which combines statistical and individual-based modelling. Using a statistical technique for forward modelling of the IBM has the advantage of being faster for parameterization than a pure inverse modelling technique and allows for robust selection of parameters. Using GPS locations from caribou monitored in Québec, caribou movements were modelled based on generative mechanisms accounting for dynamic variables at a low level of emergence. These variables were accessed by replicating real individuals' movements in parallel sub-models, and movement parameters were then empirically parameterized using Step Selection Functions. The final IBM model was validated using both k-fold cross-validation and emergent patterns validation and was tested for two different scenarios, with varying hardwood encroachment. Our results highlighted a functional response in habitat selection, which suggests that our method was able to capture the complexity of the natural system, and adequately provided projections on future possible states of the system in response to different management plans. This is especially relevant for testing the long-term impact of scenarios corresponding to environmental configurations that have yet to be observed in real systems.

Weather Conditions Drive Dynamic Habitat Selection in a Generalist Predator

Despite the dynamic nature of habitat selection, temporal variation as arising from factors such as weather are rarely quantified in species-habitat relationships. We analysed habitat use and selection (use/availability) of foraging, radio-tagged little owls (Athene noctua), a nocturnal, year-round resident generalist predator, to see how this varied as a function of weather, season and availability. Use of the two most frequently used land cover types, gardens/buildings and cultivated fields varied more than 3-fold as a simple function of season and weather through linear effects of wind and quadratic effects of temperature. Even when controlling for the temporal context, both land cover types were used more evenly than predicted from variation in availability (functional response in habitat selection). Use of two other land cover categories (pastures and moist areas) increased linearly with temperature and was proportional to their availability. The study shows that habitat selection by generalist foragers may be highly dependent on temporal variables such as weather, probably because such foragers switch between weather dependent feeding opportunities offered by different land cover types. An opportunistic foraging strategy in a landscape with erratically appearing feeding opportunities in different land cover types, may possibly also explain decreasing selection of the two most frequently used land cover types with increasing availability.

Habitat Selection and Risk of Predation: Re-colonization by Lynx had Limited Impact on Habitat Selection by Roe Deer

Risk of predation is an evolutionary force that affects behaviors of virtually all animals. In this study, we examined how habitat selection by roe deer was affected by risk of predation by Eurasian lynx – the main predator of roe deer in Scandinavia. Specifically, we compared how habitat selection by roe deer varied (1) before and after lynx re-established in the study area and (2) in relation to habitat-specific risk of predation by lynx. All analyses were conducted at the spatial and temporal scales of home ranges and seasons. We did not find any evidence that roe deer avoided habitats in which the risk of predation by lynx was greatest and information-theoretic model selection showed that re-colonization by lynx had limited impact on habitat selection by roe deer despite lynx predation causing 65% of known mortalities after lynx re-colonized the area. Instead we found that habitat selection decreased when habitat availability increased for 2 of 5 habitat types (a pattern referred to as functional response in habitat selection). Limited impact of re-colonization by lynx on habitat selection by roe deer in this study differs from elk in North America altering both daily and seasonal patterns in habitat selection at the spatial scales of habitat patches and home ranges when wolves were reintroduced to Yellowstone National Park. Our study thus provides further evidence of the complexity by which animals respond to risk of predation and suggest that it may vary between ecosystems and predator-prey constellations.

Linking trade-offs in habitat selection with the occurrence of functional responses for moose living in two nearby study areas

A species may modify its relative habitat use with changing availability, generating functional responses in habitat selection. Functional responses in habitat selection are expected to occur when animals experience trade-offs influencing their habitat selection, but only a few studies to date have explicitly linked functional responses to the underlying trade-offs faced by the animals. We used data from 39 female moose fitted with GPS telemetry collars in two nearby study areas in Canada to investigate if moose (1) were faced with a food/cover trade-off in habitat selection, as typically acknowledged in the literature, and (2) showed a functional response in their use of food/cover-rich habitats. We also examined how habitat selection patterns varied seasonally, and between study areas. The occurrence of functional responses varied strongly between study areas, and could not always be related to a measurable food/cover trade-off. Functional responses were observed more often in the study area where the environmental conditions were more severe (colder temperatures, higher precipitations, and lower food availability). Selection coefficients were also less variable among individuals in that study area, suggesting that severe environmental conditions may constrain individuals to a few selection tactics and promote the development of functional responses. Moose reacted to the availability of different habitat types in different seasons, reflecting the changing trade-offs faced by the animals. We found considerable behavioral differences between individuals from two adjacent study areas, and therefore recommend caution when extrapolating habitat selection results. We advocate for the wider use of functional responses to identify critical habitats for a species from a management or conservation perspective.

Winter resource selection by female mule deer Odocoileus hemionus: functional response to spatio‐temporal changes in habitat

Populations of mule deer Odocoileus hemionus have declined throughout most of their historic range in recent decades, and habitat alteration has been hypothesized as one potential cause of those declines. Consequently, understanding how patterns of behaviour change as landscapes are altered through time may provide important insights into mechanisms underlying observed demographic trends in populations of mule deer. We examined resource selection in relation to habitat change by mule deer on the Tex Creek winter range in southeastern Idaho, USA. We created a GIS‐based map of habitats available to mule deer during two time periods, past (1985‐1986) and current (2007‐2009), to document changes in habitat over time. We then modeled past and current patterns of resource selection by mule deer based on locations obtained from visual observation, radio‐telemetry and GPS collars. Abundance and distribution of juniper Juniperus spp., aspen Populus tremuloides, sagebrush steppe Artemisia spp. and riparian habitat did not change significantly between past and current time periods. In contrast, we documented an increase in grassland from 3.5 to 30.7% of our study area, and a corresponding decrease in agricultural land, which provides high‐quality forage for mule deer, from 37.8 to 12.5% of our study area. Patterns of resource selection largely were similar between the two time periods. Nevertheless, mule deer significantly increased selection of agricultural fields and areas far from roads between the 1980s and 2007‐2009. In addition, juniper stands were strongly selected in all years, and importance values (use × availability rescaled to 100%) for grassland and sagebrush steppe increased between past and current time periods. Our results indicated that mule deer responded behaviourally to declining availability of high‐quality forage (i.e. agricultural land) by increasing selection of agricultural fields. Such functional responses in habitat selection may have important consequences for dynamics of mule deer populations, but additional research is necessary to link those responses to demography and population performance. Although juniper encroachment often is thought to have affected mule deer negatively on many summer ranges, strong selection for juniper in our study highlights the potential importance of that species on winter range, likely because it provides both thermal and hiding cover.

Functional responses in the habitat selection of a generalist mega‐herbivore, the African savannah elephant

Resource selection function (RSF) models are commonly used to quantify species/habitat associations and predict species occurrence on the landscape. However, these models are sensitive to changes in resource availability and can result in a functional response to resource abundance, where preferences change as a function of availability. For generalist species, which utilize a wide range of habitats and resources, quantifying habitat selection is particularly challenging. Spatial and temporal changes in resource abundance can result in changes in selection preference affecting the robustness of habitat selection models. We examined selection preference across a wide range of ecological conditions for a generalist mega‐herbivore, the African savanna elephant Loxodonta africana, to quantify general patterns in selection and to illustrate the importance of functional responses in elephant habitat selection. We found a functional response in habitat selection across both space and time for tree cover, with tree cover being unimportant to habitat selection in the mesic, eastern populations during the wet season. A temporal functional response for water was also evident, with greater variability in selection during the wet season. Selection for low slopes, high tree cover, and far distance from people was consistent across populations; however, variability in selection coefficients changed as a function of the abundance of a given resource within the home range. This variability of selection coefficients could be used to improve confidence estimations for inferences drawn from habitat selection models. Quantifying functional responses in habitat selection is one way to better predict how wildlife will respond to an ever‐changing environment, and they provide promising insights into the habitat selection of generalist species.

Seasonal shifts in habitat selection of a large herbivore and the influence of human activity

Failure to recognize factors contributing to variation in habitat models like resource selection functions (RSFs) can affect their application for projecting probabilities of occurrence, and thereby limit their relevance for conservation and management. We compared seasonal RSFs (2006–2008) for 16 adult female moose ( Alces alces) with home ranges located in western Algonquin Provincial Park (APP), Ontario, Canada, to those of 14 adult females located in provincial Wildlife Management Unit (WMU) 49, 40 km west of the protected area. Wildlife and habitat management practices differed between regions: hunting was higher in WMU 49 compared to APP, and APP preserved large tracts of old growth forest rarely found in WMU 49. Seasonal RSFs projected expected similarities in moose resource use between regions (e.g., responses to wetlands and stands of eastern hemlock, Tsuga canadensis [in winter]); however, we also observed differences consistent with the hypothesis that animals, through effects of hunting, would shift habitat use seasonally and in response to roads. We further observed evidence of functional responses in habitat selection due to underlying differences in forestry practices (e.g., responses to stands of old-growth hemlock forest). Given the close proximity and shared biogeographic region between study areas, we believe that observed spatial dynamics in RSFs were ultimately reflective of divergent management strategies between areas and ensuing differences in predation and hunting mortality risk, and functional habitat.

Habitat quality influences population distribution, individual space use and functional responses in habitat selection by a large herbivore

Identifying factors shaping variation in resource selection is central for our understanding of the behaviour and distribution of animals. We examined summer habitat selection and space use by 108 Global Positioning System (GPS)-collared moose in Norway in relation to sex, reproductive status, habitat quality, and availability. Moose selected habitat types based on a combination of forage quality and availability of suitable habitat types. Selection of protective cover was strongest for reproducing females, likely reflecting the need to protect young. Males showed strong selection for habitat types with high quality forage, possibly due to higher energy requirements. Selection for preferred habitat types providing food and cover was a positive function of their availability within home ranges (i.e. not proportional use) indicating functional response in habitat selection. This relationship was not found for unproductive habitat types. Moreover, home ranges with high cover of unproductive habitat types were larger, and smaller home ranges contained higher proportions of the most preferred habitat type. The distribution of moose within the study area was partly related to the distribution of different habitat types. Our study shows how distribution and availability of habitat types providing cover and high-quality food shape ungulate habitat selection and space use.Electronic supplementary materialThe online version of this article (doi:10.1007/s00442-011-2072-3) contains supplementary material, which is available to authorised users.

Functional responses in habitat selection by tropical birds moving through fragmented forest

Summary 1. The ability of animals to move through a landscape is a fundamental determinant of population persistence in fragmented habitats. This movement can be affected by both the composition and configuration of the remaining habitat. To date, few studies have examined the habitat selection of animals moving in novel landscapes or addressed whether animals exhibit a functional response in selection as available habitat changes. 2. To assess habitat selection during movement, we translocated 60 individuals of two species of birds with differing forest dependency in three configuration treatments in a highly fragmented, tropical dry forest landscape: along a forested riparian corridor, along a fencerow (row of live trees) or across pasture. We closely followed the return routes of translocated birds to determine their choice of habitat and proximity to the forest edge. We then tested whether habitat composition or configuration (treatment) best explained individual variation in habitat selection. 3. Both species preferred habitat closer to the forest edge, but this preference was weaker in the forest specialist, the barred antshrike Thamnophilus doliatus. This species selected routes in forest habitat, which included riparian corridors, over fencerow and stepping stone habitat, which were all preferred over pasture habitat. 4. By contrast, the forest generalist, the rufous‐naped wren Campylorhynchus rufinucha preferred forest equally to fencerow and stepping stone habitat over pasture habitat. For it, fencerows were selected more strongly than stepping stones. 5. Analysis of the individual variation in selection for forest habitat revealed that both species exhibited a functional response to habitat configuration, selecting forest more strongly in riparian corridor treatments where it was also more abundant. The forest specialist also reduced its preference for edge habitat in riparian corridor treatments. 6. Synthesis and applications. The unprecedented precision of our route information demonstrates the extent to which our forest specialist preferred to travel in forest relative to fencerow and stepping stone habitat. Functional responses to habitat configuration indicated that both species make more use of other habitats when forested corridors are not present. Stepping stones in particular may be important features to the conservation of forest birds in highly fragmented habitats.

Scale dependency and functional response in moose habitat selection

Habitat selection can be influenced by the distribution of the habitat types in the landscape as well as net gain in visiting patches of resources, causing individual variation in habitat selection. Moreover, the hypothesis of functional response in habitat selection predicts that the degree of selection of a resource depends on its relative availability. We used radio‐telemetry data from individual moose on an island off the coast of northern Norway to evaluate whether the selection of habitat types at the landscape scale differed from the choice of habitat types within the home range, and investigated the functional response in habitat selection by relating individual habitat selection to home range characteristics. At the landscape scale, moose selected for habitat types that provided both good forage and cover, with small differences between sex and age groups. At the home range scale, all individuals selected habitat types that were associated with cover and low human impact. Habitat selection was not modified by local moose density, but was related to home range size at both spatial scales. Larger home ranges contained larger proportions of non‐preferred habitat types compared to smaller home ranges. At the home range scale, the selection for a habitat type decreased with its relative availability, indicating a functional response in habitat selection. This suggests that habitat selection is modified by home range size, which influences the availability of habitat types and shapes individual habitat selection patterns. Our results support previous suggestions that analyses of habitat or resource selection should follow a multi‐scale approach. Both the relative availability of habitat types as well as individual variation in home range size should be accounted for in order to disentangle the complex mechanisms that contribute to shape patterns of resource selection in animals.

Functional response in habitat selection and the tradeoffs between foraging niche components in a large herbivore

How herbivore behaviour is influenced by changes in resource levels is central for understanding trophic interactions. We examined whether foraging tradeoffs change with food levels by comparing habitat selection and space use within and between two neighbouring, predator‐free Svalbard reindeer populations. The populations faced different food levels due to contrasting grazing history. Summer resource selection in radiocollared females was assessed by a multi‐dimensional niche approach based on habitat variables obtained from a satellite image (e.g. the normalised difference vegetation index, NDVI) and a digital terrain model. The population at the overgrazed Brøggerhalvøya faced overall lower plant cover, biomass and primary productivity (i.e. lower NDVI) than the population at Sarsøyra. At Brøggerhalvøya, most reindeer selected for productive habitat when choosing home range and patches within the home range. In contrast, habitat selection at Sarsøyra was more affected by abiotic conditions such as moisture, which may influence plant quality. Here, reindeer used patches with even less biomass than the average reindeer at the poorer Brøggerhalvøya. Such a difference in habitat preference with different habitat availability (a functional response in habitat selection) probably reflected increased selection for high‐quality forage at the expense of high forage quantity at Sarsøyra. Accordingly, a negative relationship between habitat productivity and home range size was only present across individuals within Brøggerhalvøya, where forage quantity was the important foraging niche component. Individuals having poor (and large) home ranges apparently could not compensate for this by higher patch selectivity compared to individuals with richer home ranges. The results indicate changes in foraging tradeoffs at contrasting resource levels and that strong interactions occur between habitat selection, space use and the foraging niche structure in the absence of predation.

Temporal scales, trade‐offs, and functional responses in red deer habitat selection

Animals selecting habitats often have to consider many factors, e.g., food and cover for safety. However, each habitat type often lacks an adequate mixture of these factors. Analyses of habitat selection using resource selection functions (RSFs) for animal radiotelemetry data typically ignore trade-offs, and the fact that these may change during an animal's daily foraging and resting rhythm on a short-term basis. This may lead to changes in the relative use of habitat types if availability differs among individual home ranges, called functional responses in habitat selection. Here, we identify such functional responses and their underlying behavioral mechanisms by estimating RSFs through mixed-effects logistic regression of telemetry data on 62 female red deer (Cervus elaphus) in Norway. Habitat selection changed with time of day and activity, suggesting a trade-off in habitat selection related to forage quantity or quality vs. shelter. Red deer frequently used pastures offering abundant forage and little canopy cover during nighttime when actively foraging, while spending much of their time in forested habitats with less forage but more cover during daytime when they are more often inactive. Selection for pastures was higher when availability was low and decreased with increasing availability. Moreover, we show for the first time that in the real world with forest habitats also containing some forage, there was both increasing selection of pastures (i.e., not proportional use) and reduced time spent in pastures (i.e., not constant time use) with lowered availability of pastures within the home range. Our study demonstrates that landscape-level habitat composition modifies the trade-off between food and cover for large herbivorous mammals. Consequently, landscapes are likely to differ in their vulnerability to crop damage and threat to biodiversity from grazing.

Combining information from range use and habitat selection: sex‐specific spatial responses to habitat fragmentation in tawny owls Strix aluco

How individuals respond to habitat heterogeneity is usually measured as variation in range size and by ranking the relative importance of habitat types (habitat selection). The combined effect of how individuals incorporate different habitat types in their home ranges and allocate their time budget between them is rarely derived. Additionally, when home range size varies between individuals, habitat selection analyses might be flawed if foraging decisions are based on variation in absolute rather than proportional availability. We investigated the suitability of standard analytical approaches by measuring the spatial responses of tawny owls to habitat fragmentation. These owls inhabited woodland of various sizes, representing a fragmentation gradient from open farmland with small, isolated woodland patches, to continuous woodland within their home ranges. In 17 territories within open farmland, the available area covered by woodland increased with the square root of the area of open land embraced in the home range. The owls did not display functional response in habitat selection, but females selected woodland more strongly than males. Females utilised woodland 10 times more intensively in farmland than in continuous woods, whereas males utilised farmland woods 3.2 times more intensively. Moreover, females in farmland exploited woodland 3.2 times as intensively as males, apparently because of higher travel costs in open areas. Since the extensive variation in intensity of use as a function of total availability was not indicated from the analysis of habitat selection, we suggest that information about intensity of use be more widely used as a supplementary measure of habitat use patterns than appears to be the practice at present.