Floris M. van Beest

What determines variation in home range size across spatiotemporal scales in a large browsing herbivore

1. Most studies of intraspecific variation in home range size have investigated only a single or a few factors and often at one specific scale. However, considering multiple spatial and temporal scales when defining a home range is important as mechanisms that affect variation in home range size may differ depending on the scale under investigation. 2. We aim to quantify the relative effect of various individual, forage and climatic determinants of variation in home range size across multiple spatiotemporal scales in a large browsing herbivore, the moose (Alces alces), living at the southern limit of its distribution in Norway. 3. Total home range size and core home range areas were estimated for daily to monthly scales in summer and winter using both local convex hull (LoCoH) and fixed kernel home range methods. Variance in home range size was analysed using linear mixed-effects models for repeated measurements. 4. Reproductive status was the most influential individual-level factor explaining variance in moose home range size, with females accompanied by a calf having smaller summer ranges across all scales. Variation in home range size was strongly correlated with spatiotemporal changes in quantity and quality of natural food resources. Home range size decreased with increasing browse density at daily scales, but the relationship changed to positive at longer temporal scales. In contrast, browse quality was consistently negatively correlated with home range size except at the monthly scale during winter when depletion of high-quality forage occurs. Local climate affected total home range size more than core areas. Temperature, precipitation and snow depth influenced home range size directly at short temporal scales. 5. The relative effects of intrinsic and extrinsic determinants of variation in home range size differed with spatiotemporal scale, providing clear evidence that home range size is scale dependent in this large browser. Insight into the behavioural responses of populations to climatic stochasticity and forage variability is essential in view of current and future climate change, especially for populations with thermoregulatory restrictions living at the southern limit of their distribution.

Forage quantity, quality and depletion as scale-dependent mechanisms driving habitat selection of a large browsing herbivore.

1. Mechanisms that affect the spatial distribution of animals are typically scale-dependent and may involve forage distribution. Forage quality and quantity are often inversely correlated and a much discussed trade-off is whether or not to select for high-quality forage at the expense of forage abundance. This discussion has rarely involved scale-dependence or been applied to Northern browsing herbivores. At small spatial scales, browsers are assumed to select for the best quality forage. But, as high-quality forage resources are often scarce and may become depleted, coarse-scale habitat selection is assumed to be driven by forage availability. 2. To evaluate if moose selection for forage quantity and quality is scale-dependent we modelled summer and winter habitat selection of 32 GPS-marked female moose (Alces alces) at two spatial scales (landscape-scale vs. within-home range-scale). We used mixed-effects resource selection functions (RSFs) and landscape-scale forage availability models of six tree species of varying quality for moose. We considered silver birch (Betula pendula), downy birch (Betula pubescens.), Scots pine (Pinus sylvestris) as low quality browse species and rowan (Sorbus aucuparia), aspen (Populus tremula), willow (Salix spp.) as high-quality species. 3. As expected, the overall selection patterns for available browse biomass and quality varied across spatiotemporal scales. At the landscape-scale, moose selected for habitat with high available browse biomass of low quality species while at the within-home range-scale moose selected for sites with the highest quality browse species available. Furthermore, selection patterns during summer remained fairly stable, while during winter, selection at the within-home range-scale switched from sites with high quality to sites with lower quality browse species which suggests depletion of high-quality species. Consistent with expectations from seasonal resource depletion, site fidelity (bimonthly home range overlap) was much lower in winter than in summer. 4. Coarse-scale habitat selection by moose as a function of forage variability revealed a scale-dependent trade-off between available browse quantity and browse quality. Moreover, resource depletion changed the winter selection criteria of free-ranging moose and we demonstrate how the behavioural response to such a dynamic process can be inferred from RSFs.

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.

Impacts of wildlife baiting and supplemental feeding on infectious disease transmission risk: A synthesis of knowledge

Baiting and supplemental feeding of wildlife are widespread, yet highly controversial management practices, with important implications for ecosystems, livestock production, and potentially human health. An often underappreciated threat of such feeding practices is the potential to facilitate intra- and inter-specific disease transmission. We provide a comprehensive review of the scientific evidence of baiting and supplemental feeding on disease transmission risk in wildlife, with an emphasis on large herbivores in North America. While the objectives of supplemental feeding and baiting typically differ, the effects on disease transmission of these practices are largely the same. Both feeding and baiting provide wildlife with natural or non-natural food at specific locations in the environment, which can result in large congregations of individuals and species in a small area and increased local densities. Feeding can lead to increased potential for disease transmission either directly (via direct animal contact) or indirectly (via feed functioning as a fomite, spreading disease into the adjacent environment and to other animals). We identified numerous diseases that currently pose a significant concern to the health of individuals and species of large wild mammals across North America, the spread of which are either clearly facilitated or most likely facilitated by the application of supplemental feeding or baiting. Wildlife diseases also have important threats to human and livestock health. Although the risk of intra- and inter-species disease transmission likely increases when animals concentrate at feeding stations, only in a few cases was disease prevalence and transmission measured and compared between populations. Mostly these were experimental situations under controlled conditions, limiting direct scientific evidence that feeding practices exacerbates disease occurrence, exposure, transmission, and spread in the environment. Vaccination programs utilizing baits have received variable levels of success. Although important gaps in the scientific literature exist, current information is sufficient to conclude that providing food to wildlife through supplemental feeding or baiting has great potential to negatively impact species health and represents a non-natural arena for disease transmission and preservation. Ultimately, this undermines the initial purpose of feeding practices and represents a serious risk to the maintenance of biodiversity, ecosystem functioning, human health, and livestock production. Managers should consider disease transmission as a real and serious concern in their decision to implement or eliminate feeding programs. Disease surveillance should be a crucial element within the long-term monitoring of any feeding program in combination with other available preventive measures to limit disease transmission and spread.

Comparative space use and habitat selection of moose around feeding stations.

Abstract The practice of feeding cervids in winter, either as a supplement to enhance nutritional status or to divert animals away from roads, railways, or vulnerable habitats, is rising noticeably. Moose (Alces alces) densities in Scandinavia are currently at historically high levels, resulting in amplified damage to economically important young Scots pine (Pinus sylvestris) forest stands. Nevertheless, there is limited information as to how diversionary feeding affects herbivore space use and habitat selection. We followed 32 female moose marked with Global Positioning System collars to evaluate 1) if feeding stations serve as attraction points to the extent that habitat-selection patterns resemble those of central-place foragers (i.e., high usage and more uniform selection close to the attraction point), and 2) if moose using feeding sites select young pine stands less than those not using feeding sites. Moose that used diversionary forage concentrated their space use around feeding stations and selected habitats as predicted for a central-place forager with a decreasing probability of using areas away from feeding sites and a low degree of habitat selectivity close to feeding sites. However, moose that used feeding sites continued to select young pine stands to the same extent as moose that did not use feeding sites. Feeding sites were, therefore, not successful in diverting moose away from valuable natural browse, so we recommend wildlife managers establish feeding sites in sacrifice areas where moose browsing is permissible and, if possible, >1 km from young pine plantations.

Increasing density leads to generalization in both coarse‐grained habitat selection and fine‐grained resource selection in a large mammal

Density is a fundamental driver of many ecological processes including habitat selection. Theory on density-dependent habitat selection predicts that animals should be distributed relative to profitability of habitat, resulting in reduced specialization in selection (i.e. generalization) as density increases and competition intensifies. Despite mounting empirical support for density-dependent habitat selection using isodars to describe coarse-grained (interhabitat) animal movements, we know little of how density affects fine-grained resource selection of animals within habitats [e.g. using resource selection functions (RSFs)]. Using isodars and RSFs, we tested whether density simultaneously modified habitat selection and within-habitat resource selection in a rapidly growing population of feral horses (Equus ferus caballus Linnaeus; Sable Island, Nova Scotia, Canada; 42% increase in population size from 2008 to 2012). Among three heterogeneous habitat zones on Sable Island describing population clusters distributed along a west-east resource gradient (west-central-east), isodars revealed that horses used available habitat in a density-dependent manner. Intercepts and slopes of isodars demonstrated a pattern of habitat selection that first favoured the west, which generalized to include central and east habitats with increasing population size consistent with our understanding of habitat quality on Sable Island. Resource selection functions revealed that horses selected for vegetation associations similarly at two scales of extent (total island and within-habitat zone). When densities were locally low, horses were able to select for sites of the most productive forage (grasslands) relative to those of poorer quality. However, as local carrying capacity was approached, selection for the best of available forage types weakened while selection for lower-quality vegetation increased (and eventually exceeded that of grasslands). Isodars can effectively describe coarse-grained habitat selection in large mammals. Our study also shows that the main predictions of density-dependent habitat selection are highly relevant to our interpretation of RSFs in space and time. At low but not necessarily high population size, density will be a leading indicator of habitat quality. Fitness maximization from specialist vs. generalist strategies of habitat and resource selection may well be apparent at multiple spatial extents and grains of resolution.

Behavioural responses to thermal conditions affect seasonal mass change in a heat-sensitive northern ungulate.

Background Empirical tests that link temperature-mediated changes in behaviour (activity and resource selection) to individual fitness or condition are currently lacking for endotherms yet may be critical to understanding the effect of climate change on population dynamics. Moose (Alces alces) are thought to suffer from heat stress in all seasons so provide a good biological model to test whether exposure to non-optimal ambient temperatures influence seasonal changes in body mass. Seasonal mass change is an important fitness correlate of large herbivores and affects reproductive success of female moose. Methodology/Principal Findings Using GPS-collared adult female moose from two populations in southern Norway we quantified individual differences in seasonal activity budget and resource selection patterns as a function of seasonal temperatures thought to induce heat stress in moose. Individual body mass was recorded in early and late winter, and autumn to calculate seasonal mass changes (n = 52 over winter, n = 47 over summer). We found large individual differences in temperature-dependent resource selection patterns as well as within and between season variability in thermoregulatory strategies. As expected, individuals using an optimal strategy, selecting young successional forest (foraging habitat) at low ambient temperatures and mature coniferous forest (thermal shelter) during thermally stressful conditions, lost less mass in winter and gained more mass in summer. Conclusions/Significance This study provides evidence that behavioural responses to temperature have important consequences for seasonal mass change in moose living in the south of their distribution in Norway, and may be a contributing factor to recently observed declines in moose demographic performance. Although the mechanisms that underlie the observed temperature mediated habitat-fitness relationship remain to be tested, physiological state and individual variation in thermal tolerance are likely contributory factors. Climate-related effects on animal behaviour, and subsequently fitness, are expected to intensify as global warming continues.

Reproductive success and failure: the role of winter body mass in reproductive allocation in Norwegian moose

A life history strategy that favours somatic growth over reproduction is well known for long-lived iteroparous species, especially in unpredictable environments. Risk-sensitive female reproductive allocation can be achieved by a reduced reproductive effort at conception, or the subsequent adjustment of investment during gestation or lactation in response to unexpected environmental conditions or resource availability. We investigated the relative importance of reduced investment at conception compared with later in the reproductive cycle (i.e. prenatal, perinatal or neonatal mortality) in explaining reproductive failure in two high-density moose (Alces alces) populations in southern Norway. We followed 65 multiparous, global positioning system (GPS)-collared females throughout the reproductive cycle and focused on the role of maternal nutrition during gestation in determining reproductive success using a quasi-experimental approach to manipulate winter forage availability. Pregnancy rates in early winter were normal (≥0.8) in all years while spring calving rates ranged from 0.4 to 0.83, with prenatal mortality accounting for most of the difference. Further losses over summer reduced autumn recruitment rates to 0.23–0.69, despite negligible predation. Over-winter mass loss explained variation in both spring calving and autumn recruitment success better than absolute body mass in early or late winter. Although pregnancy was related to body mass in early winter, overall reproductive success was unrelated to pre-winter body condition. We therefore concluded that reproductive success was limited by winter nutritional conditions. However, we could not determine whether the observed reproductive allocation adjustment was a bet-hedging strategy to maximise reproduction without compromising survival or whether females were simply unable to invest more resources in their offspring.

Factors driving variation in movement rate and seasonality of sympatric ungulates

Abstract Defining biologically relevant seasons is a critical issue in the interpretation of animal space-use studies. Moreover, understanding the effects of extrinsic (e.g., predation risk) and intrinsic (e.g., age and sex) factors on individual differences in seasonal transition dates will deepen our understanding of the mechanisms driving animal movement and potentially population dynamics. We used nonlinear modeling of movement rate over time using global positioning system–collared nonmigratory elk (Cervus elaphus manitobensis) and white-tailed deer (Odocoileus virginianus) in southern Manitoba, Canada, to derive species- and sex-specific seasonal transition dates. In addition, we used variables known to influence timing of migration in migratory populations to explain individual differences in seasonal transition dates. We found ecologically important differences in start and length of seasons between male and female elk and white-tailed deer. Individual differences in seasonal transition dates were large, and could be explained by a combination of intrinsic and extrinsic factors. Age-class of the individual animal and elevation influenced timing of winter, spring, and date of parturition, whereas predation risk from wolves (Canis lupus) influenced onset of spring, summer, and autumn. Our findings suggest that similar extrinsic and intrinsic factors can influence both large- (i.e., migratory) and small-scale movement patterns and can be used effectively to empirically define biologically relevant seasons for sympatric large herbivores.

Density-Dependent Effects on Group Size Are Sex-Specific in a Gregarious Ungulate

Density dependence can have marked effects on social behaviors such as group size. We tested whether changes in population density of a large herbivore (elk, Cervus canadensis) affected sex-specific group size and whether the response was density- or frequency-dependent. We quantified the probability and strength of changes in group sizes and dispersion as population density changed for each sex. We used group size data from a population of elk in Manitoba, Canada, that was experimentally reduced from 1.20 to 0.67 elk/km2 between 2002 and 2009. Our results indicated that functional responses of group size to population density are sex-specific. Females showed a positive density-dependent response in group size at population densities ≥0.70 elk/km2 and we found evidence for a minimum group size at population density ≤0.70 elk/km2. Changes in male group size were also density-dependent; however, the strength of the relationship was lower than for females. Density dependence in male group size was predominantly a result of fusion of solitary males into larger groups, rather than fusion among existing groups. Our study revealed that density affects group size of a large herbivore differently between males and females, which has important implications for the benefits e.g., alleviating predation risk, and costs of social behaviors e.g., competition for resources and mates, and intra-specific pathogen transmission.