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.

Summary1. 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.

Summary1. There is a rapidly growing literature on how climate affects populations of vertebrates. For large herbivorous mammals, most attention has been paid to demographic responses to climate variation. Much less information is available regarding how climate affects animal behaviour, i.e. the climate mechanisms. Further, the appropriate measurement scale of climate variables remains debated. Here, we investigate how local climate variables determine home range sizes at four temporal scales using the Bo¨rger-method on GPS telemetry data from 47 female red deer Cervus elaphus L. in Norway. 2. If local climate operates directly on the immediate activity level of the animal, we predict home range sizes to show season-specific variation on short temporal scale (weekly-daily) related to temperature and precipitation. If local climate operate indirectly through plant growth, we rather predict variation in home range sizes to be apparent on longer time scales (biweekly-monthly), and during summer only. 3. At all time scales home range size was positively correlated with temperature during winter and negatively during summer, while the effect of precipitation was season-and scale-specific, except when accumulating as snow. Extensive snow cover decreased home range size, indicating direct effects of climate. 4. The effects of local climate was weaker at the shortest time scales (weekly-daily) compared to the longest time scales (monthly-biweekly), while the effects of day length on home range size was only apparent on the monthly and daily scale. At the longest time scales variation in local climate had a large effect on home range size. This is consistent with climatic variables operating indirectly through plant growth, but we cannot exclude a certain direct effect even at longer time scales. 5. We show how local climate-home range size correlations measured over different temporal scales can be used to infer direct and indirect climate mechanisms. Insight on the behavioural basis of responses to climate enables more accurate predictions of possible nonlinear relationships to future global warming.

Highlights d Ungulates moved to track forage in landscapes with wavelike spring green-up d Patterns of green-up explained where migratory behavior occurred in many ecosystems d At the species level, migrants and residents received equivalent foraging benefits d Movement tactics represent behavioral adaptations to specific landscapes

Autumn has to a large extent been neglected in the climate effect literature, yet autumn events, e.g., plant senescence and animal migration, affect fitness of animals differently than spring events. Understanding how variables including plant phenology influence timing of autumn migrations is important to gain a comprehensive understanding of the full annual cycle of migratory species. Here we use 13 yr of data from 60 male and 168 female red deer (Cervus elaphus) to identify triggers of autumn migration. We relate the timing of autumn migration to environmental variables like snow fall, temperature, and plant phenology (NDVI), and to onset of hunting, sex, and migration distance. Severe weather has been suggested as the main trigger of autumn migration, but we found that the majority of the individuals had left the summer range well before snow fall (80.3%) and frost (70.5%), and also before the peak deterioration in forage quality (71.9%). Declining temperatures were associated with a higher daily migration potential. Onset of hunting showed the largest effect on migration potential, with a marked increase during the first days of hunting. Individuals still present in the summer range when snow fall, frost, or peak forage deterioration occurred showed a significantly higher migration potential around these events. Males were less responsive to environmental cues, suggesting rutting activity, starting earlier in males, initiate movement prior to such conditions. Also, individuals with longer migration distances had a higher migration potential late in the season than individuals with shorter migration distances. Our study shows that factors beyond weather and plant phenology, such as onset of hunting, may be important triggers of autumn migration. Severe weather and forage deterioration were important triggers for the individuals experiencing this, which suggests a hierarchical response to environmental cues. The trade-off between staying longer in the summer range and increased energy expenditures if surprised by severe weather is asymmetric, and leaving well in advance can be seen as a risk-averse tactic.

The forage maturation hypothesis (FMH) states that herbivores should follow the onset of growth in spring to obtain access to forage of higher quality and quantity, the so-called "green wave surfing." Several studies have found correlative evidence in support of this by associating animal movement with plant phenology. However, experimental manipulation of vast natural systems determining causes of large herbivore movement is usually beyond reach. The unique management system involving winter enclosures for wild red deer (Cervus elaphus) in Germany facilitated an opportunity for an experimental approach. We manipulated release dates of red deer into free-ranging conditions in spring, predicting increased overall access to high quality forage if released early (1 April), and more rapid initial movement speed towards higher elevation if released late (15 May). The latter had lower access to high quality forage than individuals released early, as they missed parts of the green wave. In strong support of the FMH, individuals released late moved at faster initial speed than early released individuals which tracked the green wave more closely, both settling when reaching similar elevations. This shows that red deer were flexible in their movements, and they can adapt to new patterns of phenology by phenotypic plasticity.

. 2016. Behavioral buffering of extreme weather events in a high-Arctic herbivore. Ecosphere 7(6):e01374. 10.1002/ecs2.1374Abstract. As global warming advances, there is a growing concern about the impact of extreme weather events on ecosystems. In the Arctic, more frequent unseasonal warm spells and rain-on-snow events in winter cause changes in snow-pack properties, including ground icing. Such extreme weather events are known to have severe effects across trophic levels, for instance, causing die-offs of large herbivores. However, the extent to which individuals and populations are able to buffer such events through behavioral plasticity is poorly understood. Here, we analyze responses in space use to rain-on-snow and icing events, and their fitness correlates, in wild reindeer in high-Arctic Svalbard. Range displacement among GPS-collared females occurred mainly in icy winters to areas with less ice, lower over-winter body mass loss, lower mortality rate, and higher subsequent fecundity, than the departure area. Our study provides rare empirical evidence that mammals may buffer negative effects of climate change and extreme weather events by adjusting behavior in highly stochastic environments. Under global warming, behavioral buffering may be important for the long-term population persistence in mobile species with long generation time and therefore limited ability for rapid evolutionary adaptation.

Summary1. Human harvesting has a large impact on natural populations and may cause undesirable life-history changes. In wild ungulate populations, unrestricted trophy hunting may cause strong selection pressures resulting in evolutionary change towards smaller trophies. It has rarely been tested how harvesting selection varies in space and time, and whether directional hunter selection is sufficiently strong to induce long-term decreases in trophy size in centuryscale data. 2. We analysed two unique data sets of harvesting records spanning decade and century scales to identify changes in trophy size and how harvesting selection varies in space and time in red deer Cervus elaphus. We contrasted predictions from the trophy-hunting depletion, the restricted trophy hunting and the hunting pressure hypotheses. 3. Foreign hunters selected older and larger males than local hunters, but selection patterns for age-specific trophy size between counties and over time were dynamic. Patterns of red deer trophy size development from exhibitions (representing the 'upper tail' of antler sizes) were remarkably similar across Hungary from 1881 to 2008. A weak decline in trophy size between 1881 and 1958 was followed by a strong increase in trophy size between 1958 and 1974, culminating in a period of stable antler tine numbers and a weak decline in beam length until 2008. 4. We rejected the trophy hunting depletion hypothesis due to the increase in trophy size after a period of decline; patterns were most consistent with the hunting pressure hypothesis. Large increases in trophy size during 1958-1974 were likely due to a relief in hunting pressure due to implementation of strict management regulations allowing stags to grow old after the massive overharvesting during World War II, but we cannot exclude impacts from environmental factors, and that data from trophy exhibitions may underestimate trends. 5. Synthesis and applications. Trophy hunting does not necessarily lead to a non-reversible decline in trophy size, even over century-long time-scales. To ensure sustainable trophy hunting management, we need to consider factors such as spatial and temporal refuges, compensatory culling, saving stags until prime-age culmination and higher prices for larger trophies.