Animals exhibit a diversity of movement tactics [1]. Tracking resources that change across space and time is predicted to be a fundamental driver of animal movement [2]. For example, some migratory ungulates (i.e., hooved mammals) closely track the progression of highly nutritious plant green-up, a phenomenon called "green-wave surfing" [3-5]. Yet general principles describing how the dynamic nature of resources determine movement tactics are lacking [6]. We tested an emerging theory that predicts surfing and the existence of migratory behavior will be favored in environments where green-up is fleeting and moves sequentially across large landscapes (i.e., wave-like green-up) [7]. Landscapes exhibiting wave-like patterns of green-up facilitated surfing and explained the existence of migratory behavior across 61 populations of four ungulate species on two continents (n = 1,696 individuals). At the species level, foraging benefits were equivalent between tactics, suggesting that each movement tactic is fine-tuned to local patterns of plant phenology. For decades, ecologists have sought to understand how animals move to select habitat, commonly defining habitat as a set of static patches [8, 9]. Our findings indicate that animal movement tactics emerge as a function of the flux of resources across space and time, underscoring the need to redefine habitat to include its dynamic attributes. As global habitats continue to be modified by anthropogenic disturbance and climate change [10], our synthesis provides a generalizable framework to understand how animal movement will be influenced by altered patterns of resource phenology.© 2020 Elsevier Inc.

Faced with rapid environmental changes, individuals may express different magnitude and plasticity in their response to a given stressor. However, little is known about the causes of variation in phenotypic plasticity of the stress response in wild populations. In the present study, we repeatedly captured individual roe deer (Capreolus capreolus) from two wild populations in Sweden exposed to differing levels of predation pressure and measured plasma concentrations of stress-induced cortisol and behavioral docility. While controlling for the marked effects of habituation, we found clear between-population differences in the stress-induced cortisol response. Roe deer living in the area that was recently recolonized by lynx (Lynx lynx) and wolves (Canis lupus) expressed cortisol levels that were around 30% higher than roe deer in the human-dominated landscape free of large carnivores. In addition, for the first time to our knowledge, we investigated the stress-induced cortisol response in free-ranging newborn fawns and found no evidence for hypo-responsiveness during early life in this species. Indeed, stress-induced cortisol levels were of similar magnitude and differed between populations to a similar extent in both neonates and adults. Finally, at an individual level, we found that both cortisol and docility levels were strongly repeatable, and weakly negatively inter-correlated, suggesting that individuals differed consistently in how they respond to a stressor, and supporting the existence of a stress-management syndrome in roe deer. © 2018, The Author(s).

Context Intraspecific differences in behaviour can affect censuses and bias population estimates, suggesting that choice and implementation of census methods are fundamental, and need to be adapted to behavioural variations. Aims We investigated temporal variations in activity patterns during the rut among red deer (Cervus elaphus) categories and the implications for two different census methods. Methods We used a long-term dataset collected during 17 consecutive red deer rutting seasons in southernmost Sweden. The two census methods were: (1) a collection of observation ratios; and (2) a count of individuals including identification of males. Both methods are commonly used in ungulate management. Key results There was a difference in activity among age and sex categories, with a temporal variation in activity and/or presence at rutting grounds of adult (≥6 years) and subadult (2-5 years) males. Observation ratios of adult and subadult males increased from low at the start of the rut to a top level during peak rut, with subadults lagging behind adults. Before and during peak rut, the proportion of adult males was higher than that of subadults. After peak rut, the proportion of adult males decreased, whereas subadult males remained high, resulting in a higher number of subadults than of adults. The comparison of the two census methods revealed a strong correlation regarding the trends of population size and for the age and sex categories. There was also a strong consistency concerning the calf/female ratio. The male proportion was, however, consistently lower in the collected observations than in the counts. Conclusions The lower proportion of males in observations compared with counts may be explained by behavioural differences among male age classes, i.e. by temporal variations in presence and activity. That females, calves and yearling males are stationary during the rut, but adult and subadult males arrive and depart the rutting grounds at varying points of time, can lead to an underestimated male proportion in continuously collected observation data. Implications The results suggest that census should be conducted during peak rut, and that incorporating identification of individual males in the monitoring may be beneficiary. © CSIRO 2017.

Deer (Cervidae) cause considerable damage to forest plantations, crops, and protected habitats. The most common response to this damage is to implement strategies to lower population densities. However, lowering deer density may not always be desirable from hunting, recreational, or conservation perspectives. Therefore, knowledge is needed about additional factors beyond deer density that affect damage levels, and management actions that consider competing management goals. We studied the relationships between levels of bark-stripping by red deer (Cervus elaphus) on Norway spruce (Picea abies) and (1) relative deer density indices (pellet group count and deer harvest data), (2) availability of alternative natural forage (cover of forage species) and (3) proportion forest in the landscape, both at a forest stand scale and at a landscape scale. Extensive variation in damage level was evident between the six study areas. On a stand scale, the proportion of spruce damaged was positively related to pellet group density, indicating the importance of local deer usage of stands. In addition, available alternative forage in the field layer within spruce stands and proportion forest surrounding stands was negatively related to damage level. On the landscape scale, damage level was negatively related to availability of forage in the field and shrub layers and proportion forest, but was not related to any of the relative deer density indices. Increasing alternative forage may thus decrease damage and thereby reduce conflicts. Additionally, the proportion of forest in the landscape affects damage levels and should thus be considered in landscape planning and when forecasting damage risk. The relationship between local deer usage of stands and damage level suggests that future studies should try to separate the effects of local deer usage and deer density. © 2014 Jarnemo et al.

Knowledge about deer spatial use is essential for damage mitigation, conservation, and harvest management. We assess annual and seasonal home range sizes in relation to habitat composition for red deer (Cervus elaphus) in Sweden, using GPS-data from two regions with different management systems. We compare our findings with reviewed data on red deer home range sizes in Europe. Annual and seasonal home ranges during calving, hunt, and winter-spring, decreased with increasing proportion forest. Female annual home ranges in a mixed agricultural-forest landscape were three times larger than in a forest-dominated landscape. Core areas (50% Kernels) were approximately 1/5 of the full annual and seasonal home ranges (95% Kernels) regardless of habitat composition. Home range size in the forest-dominated landscape showed little inter-seasonal variation. In the agricultural-forest landscape, home ranges were larger during calving, hunt, and winter-spring compared to summer and rut. In the forest-dominated landscape, management areas are large enough to cover female spatial use. In the agricultural-forest landscape, female spatial use covers several license units. Here, the coordinated license system is needed to reach trade-offs between goals of conservation, game management, and damage mitigation. Males had in general larger home ranges than females, and the majority of the males also made a seasonal migration to and from the rutting areas. The license system area in the agricultural-forest landscape is large enough to manage migrating males. In the forest landscape, a coordination of several management areas is needed to encompass male migrations. We conclude that management needs to adapt to deer spatial use in different types of landscapes to reach set goals. © 2023, The Author(s).

Deer can show transitional use between agricultural fields and forests for foraging and shelter. Such transitional use may affect forest damage as nutrient balancing theory suggests that if deer ingest large amounts of nutrient-rich food, complementary browse, such as bark, may be required to balance the diet. We investigated the relationship between the level of red deer Cervus elaphus bark-stripping damage in 68 Norway spruce Picea abies stands and the presence of rapeseed Brassica napus fields – an energy-rich crop preferred by red deer – in the surroundings, hypothesizing that damage increases with decreasing distance to rapeseed fields. We also considered other potentially influencing factors, such as supplemental feeding, alternative forage availability, and deer use of spruce stands as indexed by a pellet group count. Spruce stands closer to rapeseed had a significantly higher proportion of damaged stems. The increased level of bark-stripping damage was not explained by a higher stand use of deer closer to rapeseed fields, indicating that deer increase their consumption of bark in order to balance their diet. Similarly, spruce stands closer to supplemental feeding stations had significantly higher damage levels. In line with earlier findings, damage levels were negatively related to the amount of available browse in the forest. This emphasizes the importance of alternative forage for reducing the damage risk in forest plantations. Our study shows that the availability of fields with nutrient-rich food, such as rapeseed, as well as supplemental feeding needs to be considered when predicting the level of forest damage. With a high availability of nutrient-rich food in the vicinity of forest stands, a higher damage level can be expected and counteractions could be taken such as increased disturbance, harvest or changed choice of both crop and supplemental feed types. These actions may also be combined with a push–pull strategy where the deer are steered to disturbance-free zones insensitive to damage and with alternative forage. The importance of alternative forage availability on damage levels highlights the necessity for managers to actively promote tree and shrub forage within and around their production stands. © 2022 The Authors. Ecological Solutions and Evidence published by John Wiley & Sons Ltd on behalf of British Ecological Society.

Common understory vegetation species such as the ericaceous shrubs bilberry (Vaccinium myrtillus), cowberry (V. vitis-idaea) and heather (Calluna vulgaris), are key forage plant species for moose and other large herbivores, as well as fulfilling many additional ecosystem functions and services. Here we developed models to predict above-ground biomass of these ericaceous species in coniferous forests, using data on their percentage cover, height, and different stand characteristics. We also built models to understand how the aforementioned variables affect the proportion of the shrubs commonly utilized as forage by large herbivores. We found that the percentage cover of shrubs was the most important explanatory variable when predicting above-ground biomass, explaining 51%, 47% and 71% of the variation (marginal R²) in bilberry, cowberry and heather biomass, respectively. By adding ramet height to the model with percentage cover, the variation explained increased to 77% for bilberry, 75% for cowberry and 87% for heather. The best outcome for candidate models was obtained by adding stand site index and spruce basal area to the model, improving the variation explained in bilberry to 83%, to 81% for cowberry, and 91% for heather. When modelling the proportion of the shrubs commonly utilized as forage by large herbivores, stand site index and spruce basal area often played important roles. Some of the best fitting models for forage biomass explained 51% of the variation in bilberry, 59% in cowberry and 30% in heather. Site location did not have a major role in improving the variability explained in either type of model, which indicated the applicability of the models regardless of study location. Our models therefore have a high potential to be implemented in forestry decision support systems. Their inclusion should provide better large-scale estimations of forage resources, aiding forest management, and thereby taking an important step forward to determine the ecosystem carrying capacity of large herbivores. © 2023 The Author(s)

Human activity and associated landscape modifications alter the movements of animals with consequences for populations and ecosystems worldwide. Species performing long-distance movements are thought to be particularly sensitive to human impact. Despite the increasing anthropogenic pressure, it remains challenging to understand and predict animals' responses to human activity. Here we address this knowledge gap using 1206 Global Positioning System movement trajectories of 815 individuals from 14 red deer (Cervus elaphus) and 14 elk (Cervus canadensis) populations spanning wide environmental gradients, namely the latitudinal range from the Alps to Scandinavia in Europe, and the Greater Yellowstone Ecosystem in North America. We measured individual-level movements relative to the environmental context, or movement expression, using the standardized metric Intensity of Use, reflecting both the directionality and extent of movements. We expected movement expression to be affected by resource (Normalized Difference Vegetation Index, NDVI) predictability and topography, but those factors to be superseded by human impact. Red deer and elk movement expression varied along a continuum, from highly segmented trajectories over relatively small areas (high intensity of use), to directed transitions through restricted corridors (low intensity of use). Human activity (Human Footprint Index, HFI) was the strongest driver of movement expression, with a steep increase in Intensity of Use as HFI increased, but only until a threshold was reached. After exceeding this level of impact, the Intensity of Use remained unchanged. These results indicate the overall sensitivity of Cervus movement expression to human activity and suggest a limitation of plastic responses under high human pressure, despite the species also occurring in human-dominated landscapes. Our work represents the first comparison of metric-based movement expression across widely distributed populations of a deer genus, contributing to the understanding and prediction of animals' responses to human activity. Global Change Biology© 2023 The Authors. Global Change Biology published by John Wiley & Sons Ltd.

Crop raiding ungulates can cause costly yield losses and conflicts between agriculture, game management, and conservation. It is therefore crucial to know how, where, and when ungulates use different habitats and crops. In this study we used resource selection functions and GPS-tagged red deer Cervus elaphus (n = 38) in two different study areas (North – mainly forested land and South – mainly arable land), to investigate how red deer use arable land and different crop types in relation to availability and distance to forest cover in Sweden. Our study shows a transitional use of arable land and forests by red deer. Red deer spend on average 45% and 21% of their time in arable land in South and North respectively. In the North, arable land was selected while forest and wetlands were selected in the South. The selection of different crops also differed between the two study areas, for example rapeseed was highly selected during both seasons in South while it was used to a lower degree in relation to its availability in the North. In both study areas the probability of red deer presence on the agricultural fields decreased with distance to forest. The significant use of arable land and unharvested crops by the increasing red deer population in Sweden highlights a risk for crop damage that may need further consideration for farming practices as well as for damage and wildlife management. In our case damage mitigation should focus on rapeseed in the South, whereas there is a less clear pattern of selection among growing crops in the North. The differences in habitat and crop selection between the two areas also highlights the need of knowledge about red deer habitat selection at a regional level to be able to adapt damage mitigation and wildlife management strategies accordingly. Moreover, the transitional use of arable land and forests by red deer in mixed landscapes may imply that consumption of certain crops can affect browsing patterns and damage levels within forests and vice versa, that may have implication for both agriculture and forestry and calls for future studies. © 2021

The most common framework under which ungulate migration is studied predicts that it is driven by spatio–temporal variation in plant phenology, yet other hypotheses may explain differences within and between species. To disentangle more complex patterns than those based on single species/ single populations, we quantified migration variability using two sympatric ungulate species differing in their foraging strategy, mating system and physiological constraints due to body size. We related observed variation to a set of hypotheses. We used GPS-collar data from 537 individuals in 10 roe Capreolus capreolus and 12 red deer Cervus elaphus populations spanning environmental gradients across Europe to assess variation in migration propensity, distance and timing. Using time-to-event models, we explored how the probability of migration varied in relation to sex, landscape (e.g. topography, forest cover) and temporally-varying environmental factors (e.g. plant green-up, snow cover). Migration propensity varied across study areas. Red deer were, on average, three times more migratory than roe deer (56% versus 18%). This relationship was mainly driven by red deer males which were twice as migratory as females (82% versus 38%). The probability of roe deer migration was similar between sexes. Roe deer (both sexes) migrated earliest in spring. While territorial male roe deer migrated last in autumn, male and female red deer migrated around the same time in autumn, likely due to their polygynous mating system. Plant productivity determined the onset of spring migration in both species, but if plant productivity on winter ranges was sufficiently high, roe deer were less likely to leave. In autumn, migration coincided with reduced plant productivity for both species. This relationship was stronger for red deer. Our results confirm that ungulate migration is influenced by plant phenology, but in a novel way, that these effects appear to be modulated by species-specific traits, especially mating strategies. © 2018 The Authors. Oikos © 2018 Nordic Society Oikos

Wildlife tagging provides critical insights into animal movement ecology, physiology, and behavior amid global ecosystem changes. However, the stress induced by capture, handling, and tagging can impact post-release locomotion and activity and, consequently, the interpretation of study results. Here, we analyze post-tagging effects on 1585 individuals of 42 terrestrial mammal species using collar-collected GPS and accelerometer data. Species-specific displacements and overall dynamic body acceleration, as a proxy for activity, were assessed over 20 days post-release to quantify disturbance intensity, recovery duration, and speed. Differences were evaluated, considering species-specific traits and the human footprint of the study region. Over 70% of the analyzed species exhibited significant behavioral changes following collaring events. Herbivores traveled farther with variable activity reactions, while omnivores and carnivores were initially less active and mobile. Recovery duration proved brief, with alterations diminishing within 4-7 tracking days for most species. Herbivores, particularly males, showed quicker displacement recovery (4 days) but slower activity recovery (7 days). Individuals in high human footprint areas displayed faster recovery, indicating adaptation to human disturbance. Our findings emphasize the necessity of extending tracking periods beyond 1 week and particular caution in remote study areas or herbivore-focused research, specifically in smaller mammals. © 2024. The Author(s).

The distribution and population density of red deer (Cervus elaphus) are increasing in several regions of Europe. The deer may cause severe damage in commercial forestry and agriculture. Bark stripping is the main problem in forests, especially on Norway spruce (Picea abies), and is thought to mostly occur during winter when other forage is scarce. It has been suggested that an imbalance in the nutrient intake, and especially a diet including high amounts of easily-digestible macronutrients, such as agricultural crops, can lead to an increased urge to consume bark. Feeding on brassicas, for example rapeseed (Brassica napus) might have this effect. The aim with this study was to investigate the relationship between intake of rapeseed and bark stripping on Norway spruce by red deer during early spring. We did this by a controlled feeding experiment with four groups of captive red deer in southern Sweden. All groups were given spruce logs every week, while only two groups had access to freshly harvested rapeseed plants. In addition, influence of air temperature and forage nutritional composition was taken into account. Our results show that red deer bark stripping can be considerable not only during winter but also during spring green-up. We found no significant influence of rapeseed on bark stripping performed by the deer. However, at a threshold temperature, deer suddenly started to ingest large amounts of bark biomass, coinciding with a significant change in the bark's concentration of starch. We suggest that the lack of effect of rapeseed feeding can partly be explained by overshadowing effects caused by such seasonal changes of bark characteristics, and partly by the fact that the rapeseed plants in our study contained lower than expected concentrations of easily-digestible macronutrients (apart from protein). We conclude that the risk of damage on spruce can be especially high during certain periods, something that is important to consider when mitigating bark stripping. However, several interactive effects are involved and must be considered in order to more efficiently mitigate damage. © 2022 The Author(s)

Apex predators may affect mesopredators through intraguild predation and/or supply of carrion from their prey, causing a trade-off between avoidance and attractiveness. We used wildlife triangle snow-tracking data to investigate the abundance of red fox (Vulpes vulpes) in relation to lynx (Lynx lynx) and wolf (Canis lupus) occurrence as well as land composition and vole (Microtus spp.) density. Data from the Swedish wolf-monitoring system and VHF/GPS-collared wolves were used to study the effect of wolf pack size and time since wolf territory establishment on fox abundance. Bottom-up processes were more influential than top-down effects as the proportion of arable land was the key indicator of fox abundance at the landscape level. At this spatial scale, there was no effect of wolf abundance on fox abundance, whereas lynx abundance had a positive effect. In contrast, at the wolf territory level there was a negative effect of wolves on fox abundance when including detailed information of pack size and time since territory establishment, whereas there was no effect of lynx abundance. This study shows that different apex predator species may affect mesopredator abundance in different ways and that the results may be dependent on the spatiotemporal scale and resolution of the data. © 2017 The Author(s).

Olfactory signals constitute an important mechanism in interspecific interactions, but little is known regarding their role in communication between predator species. We analyzed the behavioral responses of a mesopredator, the red fox (Vulpes vulpes), to an olfactory cue (scat) of an apex predator, the lynx (Lynx lynx) in BiałowieÅŒa Primeval Forest, Poland, using video camera traps. Red fox visited sites with scats more often than expected and the duration of their visits was longer at scat sites than at control sites (no scat added). Vigilant behavior, sniffing and scent marking (including over-marking) occurred more often at scat sites compared to control sites, where foxes mainly passed by. Vigilance was most pronounced during the first days of the recordings. Red fox behavior was also influenced by foxes previously visiting scat sites. They sniffed and scent marked (multiple over-marking) more frequently when the lynx scat had been over-marked previously by red fox. Fox visits to lynx scats may be seen as a trade-off between obtaining information on a potential food source (prey killed by lynx) and the potential risk of predation by an apex predator. © 2017, The Author(s).