Bram Van Moorter

Habitat–performance relationships: finding the right metric at a given spatial scale

The field of habitat ecology has been muddled by imprecise terminology regarding what constitutes habitat, and how importance is measured through use, selection, avoidance and other bio-statistical terminology. Added to the confusion is the idea that habitat is scale-specific. Despite these conceptual difficulties, ecologists have made advances in understanding ‘how habitats are important to animals’, and data from animal-borne global positioning system (GPS) units have the potential to help this clarification. Here, we propose a new conceptual framework to connect habitats with measures of animal performance itself—towards assessing habitat–performance relationship (HPR). Long-term studies will be needed to estimate consequences of habitat selection for animal performance. GPS data from wildlife can provide new approaches for studying useful correlates of performance that we review. Recent examples include merging traditional resource selection studies with information about resources used at different critical life-history events (e.g. nesting, calving, migration), uncovering habitats that facilitate movement or foraging and, ultimately, comparing resources used through different life-history strategies with those resulting in death. By integrating data from GPS receivers with other animal-borne technologies and combining those data with additional life-history information, we believe understanding the drivers of HPRs will inform animal ecology and improve conservation.

A model-driven approach to quantify migration patterns: individual, regional and yearly differences

1. Animal migration has long intrigued scientists and wildlife managers alike, yet migratory species face increasing challenges because of habitat fragmentation, climate change and over-exploitation. Central to the understanding migratory species is the objective discrimination between migratory and nonmigratory individuals in a given population, quantifying the timing, duration and distance of migration and the ability to predict migratory movements. 2. Here, we propose a uniform statistical framework to (i) separate migration from other movement behaviours, (ii) quantify migration parameters without the need for arbitrary cut-off criteria and (iii) test predictability across individuals, time and space. 3. We first validated our novel approach by simulating data based on established theoretical movement patterns. We then formulated the expected shapes of squared displacement patterns as nonlinear models for a suite of movement behaviours to test the ability of our method to distinguish between migratory movement and other movement types. 4. We then tested our approached empirically using 108 wild Global Positioning System (GPS)-collared moose Alces alces in Scandinavia as a study system because they exhibit a wide range of movement behaviours, including resident, migrating and dispersing individuals, within the same population. Applying our approach showed that 87% and 67% of our Swedish and Norwegian subpopulations, respectively, can be classified as migratory. 5. Using nonlinear mixed effects models for all migratory individuals we showed that the distance, timing and duration of migration differed between the sexes and between years, with additional individual differences accounting for a large part of the variation in the distance of migration but not in the timing or duration. Overall, the model explained most of the variation (92%) and also had high predictive power for the same individuals over time (69%) as well as between study populations (74%). 6. The high predictive ability of the approach suggests that it can help increase our understanding of the drivers of migration and could provide key quantitative information for understanding and managing a broad range of migratory species.

A Migratory Northern Ungulate in the Pursuit of Spring: Jumping or Surfing the Green Wave?

The forage-maturation hypothesis (FMH) states that herbivores migrate along a phenological gradient of plant development in order to maximize energy intake. Despite strong support for the FMH, the actual relationship between plant phenology and ungulate movement has remained enigmatic. We linked plant phenology (MODIS–normalized difference vegetation index [NDVI] data) and space use of 167 migratory and 78 resident red deer (Cervus elaphus), using a space-time-time matrix of “springness,” defined as the instantaneous rate of green-up. Consistent with the FMH, migrants experienced substantially greater access to early plant phenology than did residents. Deer were also more likely to migrate in areas where migration led to greater gains in springness. Rather than “surfing the green wave” during migration, migratory red deer moved rapidly from the winter to the summer range, thereby “jumping the green wave.” However, migrants and, to a lesser degree, residents did track phenological green-up through parts of the growing season by making smaller-scale adjustments in habitat use. Despite pronounced differences in their life histories, we found only marginal differences between male and female red deer in this study. Those differences that we did detect pointed toward additional constraints on female space-use tactics, such as those posed by calving and caring for dependent offspring. We conclude that whereas in some systems migration itself is a way to surf the green wave, in others it may simply be a means to reconnect with phenological spring at the summer range. In the light of ubiquitous anthropogenic environmental change, understanding the relationship between the green wave and ungulate space use has important consequences for the management and conservation of migratory ungulates and the phenomenon of migration itself.

Temperature-mediated habitat use and selection by a heat-sensitive northern ungulate

The behavioural response of animals to unfavourable climatic conditions has received increased attention recently. While many studies have examined the behavioural responses of endotherms to cold temperatures, thermoregulatory behaviour may also occur in response to heat stress. We evaluated whether a heat-sensitive northern ungulate, the moose, Alces alces, showed thermoregulatory behaviour in response to ambient temperature in two populations in southern Norway. We quantified the seasonal habitat use of GPS-collared adult females, as well as fine-scale habitat selection patterns, in relation to time of day and critical temperature thresholds thought to induce heat stress. We also assessed whether temperature-associated changes in spatial behaviour led to a shift in the availability of thermal cover and forage at the chosen location. Frequent exposure to temperatures above critical thresholds occurred in both summer and winter and in both study areas. Moose responded by seeking thermal shelter in mature coniferous forest and avoiding open habitat types, leading to a decrease in local forage availability in summer but not in winter. Differences in habitat choice in response to temperature were most pronounced at twilight. We found that fine-scale habitat selection analyses, using step-selection functions, more effectively revealed thermoregulatory behaviour in both seasons and populations than did habitat use. This is because habitat selection analyses are better able to identify limiting factors operating at different spatiotemporal scales than is habitat use. Our results clearly show that ambient temperature affected fine-scale behavioural decisions of moose with consequences for forage accessibility, especially during summer. As the climate changes, the limiting effect of ambient temperature (cold and high) on animal behaviour is likely to increase, potentially influencing individual fitness and population dynamics. 2012 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

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.

Moose Alces alces habitat use at multiple temporal scales in a human-altered landscape

Abstract Habitat alteration by humans may change the supply of food and cover for wild ungulates, but few studies have examined how these resources are utilised over time by individuals of different sex and reproductive status. We examined circadian and seasonal variation in habitat utilisation within a moose Alces alces population in central Norway. Our study area covers forests and open habitats, both influenced by human alterations (e.g. forestry and agriculture). We expected moose to select habitats with good forage and cover in all seasons, but to select open foraging habitats mainly during night-time. Moose selected good foraging habitats, such as young forest stands and cultivated land during night, whereas the utilisation of older forest stands providing cover increased during daytime. This circadian pattern changed throughout the year, seemingly related to variation in hours of daylight and provision of forage. Young forest stands provided higher density of preferred food plants compared to older stands and were highly selected from spring until autumn. Relative to young forest, the selection for older forest stands increased towards winter, likely due to provision of higher plant quality late in the growing season, and to reduced accumulation of movement-impeding snow during winter. Selection of cultivated land varied among seasons, being highest when crop biomass was high. We also found some indications of state-dependent habitat selection as reproducing females avoided open, food rich areas in the first months after their calves were born, whereas males and females without young selected these areas in spring and summer. Our results clearly show that moose exploit the variations in cover and food caused by forestry and agriculture. This is particularly relevant for moose in Norway as current changes in forestry practice lead to a reduction in young, food-rich forest stands, possibly aggravating the already declining body conditions and recruitment rates of moose.

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.

Maternal and individual effects in selection of bed sites and their consequences for fawn survival at different spatial scales

We examined the relationship between survival of roe deer (Capreolus capreolus) fawns at Trois Fontaines, Champagne-Ardennes, France, and factors related to bed-site selection (predator avoidance and thermoregulation) and maternal food resources (forage availability in the maternal home range). Previous studies have demonstrated that at small scales, the young of large herbivores select bed sites independently from their mothers, although this selection takes place within the limits of their mother’s home range. Fawn survival was influenced largely by the availability of good bed sites within the maternal home range, not by the fawn’s selection of bed sites; however, selection for thermal cover when selecting bed sites positively influenced survival of young fawns. Typical features of a good home range included close proximity to habitat edges, which is related to forage accessibility for roe deer. The availability of bed sites changed as fawns aged, probably due to an increased mobility of the fawn or a different use of the home range by the mother; sites offering high concealment and thermal protection became less available in favor of areas with higher forage accessibility. Despite the minor influence of bed-site selection on survival, roe deer fawns strongly selected their bed sites according to several environmental factors linked to predator avoidance and thermoregulation. Fawns selected for sites providing concealment, light penetration, and avoided signs of wild boar (Sus scrofa) activity. Avoidance of sites with high light penetration by young fawns positively affected their survival, confirming a negative effect on thermoregulation due to reduced thermal cover. Selection for light penetration by older fawns was less clear. We discuss these results in the context of cross-generational effects in habitat selection across multiple scales, and the potential influence of the ‘ghost of predation past’.

Understanding scales of movement: animals ride waves and ripples of environmental change

Animal movements are the primary behavioural adaptation to spatiotemporal heterogeneity in resource availability. Depending on their spatiotemporal scale, movements have been categorized into distinct functional groups (e.g. foraging movements, dispersal, migration), and have been studied using different methodologies. We suggest striving towards the development of a coherent framework based on the ultimate function of all movement types, which is to increase individual fitness through an optimal exploitation of resources varying in space and time. We developed a novel approach to simultaneously study movements at different spatiotemporal scales based on the following proposed theory: the length and frequency of animal movements are determined by the interaction between temporal autocorrelation in resource availability and spatial autocorrelation in changes in resource availability. We hypothesized that for each time interval the spatiotemporal scales of moose Alces alces movements correspond to the spatiotemporal scales of variation in the gains derived from resource exploitation when taking into account the costs of movements (represented by their proxies, forage availability NDVI and snow depth respectively). The scales of change in NDVI and snow were quantified using wave theory, and were related to the scale of moose movement using linear mixed models. In support of the proposed theory we found that frequent, smaller scale movements were triggered by fast, small-scale ripples of changes, whereas infrequent, larger scale movements matched slow, large-scale waves of change in resource availability. Similarly, moose inhabiting ranges characterized by larger scale waves of change in the onset of spring migrated longer distances. We showed that the scales of movements are driven by the scales of changes in the net profitability of trophic resources. Our approach can be extended to include drivers of movements other than trophic resources (e.g. population density, density of related individuals, predation risk) and may facilitate the assessment of the impact of environmental changes on community dynamics and conservation.

Identifying Movement States From Location Data Using Cluster Analysis

Abstract Animal movement studies regularly use movement states (e.g., slow and fast) derived from remotely sensed locations to make inferences about strategies of resource use. However, the number of movement state categories used is often arbitrary and rarely inferred from the data. Identifying groups with similar movement characteristics is a statistical problem. We present a framework based on k-means clustering and gap statistic for evaluating the number of movement states without making a priori assumptions about the number of clusters. This allowed us to distinguish 4 movement states using turning angle and step length derived from Global Positioning System locations and head movements derived from tip switches in a neck collar of free-ranging elk (Cervus elaphus) in west central Alberta, Canada. Based on movement characteristics and on the linkage between each state and landscape features, we were able to identify inter-patch movements, intra-patch foraging, rest, and inter-patch foraging movements. Linking behavior to environment (e.g., state-dependent habitat use) can inform decisions on landscape management for wildlife.