Norway spruce Picea abies is an economically important tree species in Europe, actively managed for forestry. Among the most negative biotic factors for growth and hence forest production is damage caused by wildlife, such as damage through bark stripping by red deer Cervus elaphus. We quantified bark stripping damage on Norway spruce trees in across 450 stands (aged 20-72 years) spanning a 400 km latitudinal range along Norway's west coast and analysed the underlying mechanisms driving increased probability of bark stripping by red deer. A total of 74% of tree stands had bark stripping damage. The mean percentage of damaged trees was 16.0%, but 50 stands (11.1% of the stands) had more than 50.0% damaged trees. The most important factor determining probability for bark stripping was broad-scale red deer density, where the probability increased markedly when density reached approximately two harvested red deer per km2. In addition, proximity to agricultural farmland, distance from roads, site productivity, distance between twig whorls and terrain ruggedness index increased the probability of bark stripping. Our study on bark stripping on Norway spruce highlights the importance of red deer population control, but also the importance of evaluating environmental factors as well as site factors and tree characteristics in forestry planning to mitigate damage from red deer.

Reliable estimates of the size and composition of harvested populations over time are key to designing adequate population management plans, regardless of management objectives. In Norway, a national system for collecting and analysing hunter-reported data on red deer (Cervus elaphus) has been operational for about 20 years. The system was expected to provide population metrics that would substantially improve deer population management routines at the municipal level. This has proven to be challenging when using existing state-of-the-art estimation methodology. The main reasons are that the variation in the observation data is generally much larger than population abundance variability, and that one does not have a clear understanding of the stochastic process generating the observation data. Here, using hunter-reported observation data and harvest data from six Norwegian municipalities collected in the period 2007-2023, we show that a straightforward estimation methodology based on population modelling can produce robust abundance estimates despite frequent low quality of the observation data. Its major assets are that it does not involve strong assumptions about the stochastic processes underlying the observation process and that it does not involve assumptions about initial population size and structure in terms of prior statistical distributions. We anticipate that the method can be applied in several other population management contexts, and we think that the results offer fresh perspectives on to what extent noisy citizen-collected time series data can be used to inform management decisions.

Large herbivores at northern latitudes often forage on agricultural farmland. In these populations, the presence of both resident and migrant individuals (termed partial migration) is common, but how migrants and residents differ in their selection of farmland is not well‐understood. Higher access to farmland may provide benefits to residents compensating for not following the ‘green wave’ of emerging vegetation such as migrants. According to sexual segregation theory, males and females differ in body‐size‐related nutritional needs and risk sensitivity associated with farmland. Yet, how the sexes differ in the selection of farmland through an annual cycle remains unclear. We quantified seasonal variation in the selection of farmland by partially migratory red deer ( Cervus elaphus ) at broad, landscape scale and at fine, within‐home range scale using 16 years of data (2005–2020) from 329 females and 115 males in Norway. We tested predictions related to the partial migration and sexual segregation theories using resource selection functions. We predicted higher selection for farmland by residents than by migrants, and higher selection by females than by males due to higher nutritional needs, but that higher perceived predation risk would impact their diurnal selection patterns. The time spent on farmland was higher in winter (14%–18%) than in summer (8%–14%). Residents selected farmland more than migrants mainly at broad, landscape scale, while differences were smaller and less consistent at a fine, within‐home range scale. Females showed higher broad‐scale selection for farmland in winter, whereas males showed higher selection in summer. At a fine, within‐home range scale, females selected farmland more in summer during darkness, whereas sex differences were small otherwise. The fine‐scale selection of farmland was markedly higher during low‐light conditions than in daylight. A high population density index was correlated with high broad‐scale selection of farmland, i.e. high farmland availability in the home ranges, whereas the effect of the density index was weak at a fine, within‐home range scale. Our study emphasises how hypotheses deriving from the theories of partial migration and sexual segregation can improve our understanding of ungulates' selection of farmland. The higher selection by residents during summer highlights the importance of retaining landscape connectivity, allowing for migration and reducing pressure on local resources.

Our understanding of how migratory wildlife populations affect incidence of infectious diseases spilling over into humans is limited. Ticks are expanding their distribution towards northern latitudes, causing emergence of tick-borne diseases. Deer serve as reproduction hosts for adult ticks, supporting the tick populations. However, in northern areas, deer populations are partially migratory, and migrants occupy high elevation summer ranges that are unsuitable for ticks. Migration can thus theoretically lead to escape from exposure to ticks and to deer feeding fewer ticks, which lower disease hazard to humans. Combining data on Ixodes ricinus tick abundance with GPS-tracking data of red deer (Cervus elaphus), we quantified the distributional overlap of ticks and red deer along elevational gradients in Norway. Furthermore, we correlated both deer density and the proportion of migratory deer with the incidence of Lyme disease in humans. We found that migratory deer summer ranges had colder climate and overlapped less with tick distribution than those of resident deer. Deer density consistently increased Lyme disease incidence in humans. However, we found only weak evidence that deer migration negatively affected Lyme disease incidence. Our study provides a rare quantification of how host availability, in terms of both host density and migratory movement, affects risk of a zoonotic disease.

The timing of migration is fundamental for species exploiting seasonally variable environments. For ungulates, earlier spring migration is expected with earlier vegetation green-up. However, other drivers, such as access to agricultural farmland and variation in local conditions, are also known to affect migration. We investigated the timing of spring migration for 96 male and 201 female red deer ( Cervus elaphus ) using a long-term dataset (2005–2020). Overall, the timing of migration was mainly characterized by large individual variability between and within years (95% range 6 April to 18 June). The spring migration timing was, as expected, later with colder winter and spring conditions (North Atlantic Oscillation (NAO) winter and April indices) and later peak vegetation green-up (NDVI), with a five-day delay in green-up causing a migration delay of 1.2 days. Timing was also influenced by local conditions in summer and winter home ranges. Red deer with greater access to farmland and a more variable topography (hence variable plant phenology) in winter delayed migration. Similarly, individuals with higher-elevation summer ranges (with delayed onset of plant growth) also delayed migration. Our analyses highlight that the timing of red deer migration is determined by multiple drivers affecting foraging conditions in the landscape, indicative of considerable phenotypic plasticity.

Grazing by wildlife on agricultural land is widespread across geographical regions, and can cause human–wildlife conflicts due to reduced crop yield when the grazing pressure is high. Growing red deer Cervus elaphus populations in Europe call for an increased understanding of their grazing patterns to mitigate damages. We quantified how red deer grazing pressure (grazing presence and grazing level) on agricultural grass meadows (n = 60) in Norway varied across multiple spatial scales. We used a nested, hierarchical study design transcending from a broad scale (meadows across the landscape) to intermediate (between nearby meadows) and local (within‐meadow) scales, allowing us to identify at which scale the variation in grazing pressure was strongest. We estimated how grazing was determined by broad‐scale factors influencing forage availability and quality through population density, distance to coastline, and differences between the first versus second harvest, by intermediate‐scale factors in terms of meadow management causing differences in botanical composition and quality, and by local‐scale factors in terms of perceived predation risk and disturbance. At a broad scale, higher population densities were associated with higher grazing pressure, and more grazing occurred before the first compared to the second harvest. Intermediate‐scale factors explained the most variation of grazing pressure from red deer, with higher grazing pressure on newly renewed meadows compared to other nearby meadows. On a local scale, more grazing occurred closer to the forest edge, providing cover, and further away from infrastructure, with increased risk and disturbance. Overall, our study highlights how drivers of grazing pressure on agricultural land vary across spatial scales. Population reductions on a broader scale may have some effect in reducing the grazing pressure, but renewed meadows will nevertheless attract red deer, causing higher grazing pressure compared to neighbouring meadows. This insight is crucial for determining effective mitigation strategies facing rising red deer populations across Europe.

Caring for newborn offspring hampers resource acquisition of mammalian females, curbing their ability to meet the high energy expenditure of early lactation. Newborns are particularly vulnerable, and, among the large herbivores, ungulates have evolved a continuum of neonatal antipredator tactics, ranging from immobile hider (such as roe deer fawns or impala calves) to highly mobile follower offspring (such as reindeer calves or chamois kids). How these tactics constrain female movements around parturition is unknown, particularly within the current context of increasing habitat fragmentation and earlier plant phenology caused by global warming. Here, using a comparative analysis across 54 populations of 23 species of large herbivores from 5 ungulate families (Bovidae, Cervidae, Equidae, Antilocapridae and Giraffidae), we show that mothers adjust their movements to variation in resource productivity and heterogeneity according to their offspring’s neonatal tactic. Mothers with hider offspring are unable to exploit environments where the variability of resources occurs at a broad scale, which might alter resource allocation compared with mothers with follower offspring. Our findings reveal that the overlooked neonatal tactic plays a key role for predicting how species are coping with environmental variation. Combining a large-scale dataset of 23 ungulate species (in which newborns follow contrasting tactics of predator avoidance) with continuous-time stochastic movement models, the authors reveal that there are multiple dimensions of maternal movement behaviour and space use.

Parasitic nematodes are ubiquitous and can negatively impact their host by reducing fecundity or increasing mortality, yet the driver of variation in the parasite community across a wildlife host's geographic distribution remains elusive for most species. Based on an extensive collection of fecal samples (n = 264) from GPS marked moose (Alces alces), we used DNA metabarcoding to characterize the individual (sex, age class) and seasonal parasitic nematode community in relation to habitat use and migration behavior in five populations distributed across a wide latitudinal gradient (59.6°N to 70.5°N) in Norway. We detected 21 distinct nematode taxa with the six most common being Ostertagia spp., Nematodirella spp., Trichostongylus spp., T. axei, Elaphostrongylus alces, and an unclassified Strongylida. There was higher prevalence of livestock parasites in areas with larger sheep populations indicating a higher risk of spillover events. The individual level nematode richness was mostly consistent across study areas, while the number and type of nematode taxa detected at each study area varied considerably but did not follow a latitudinal gradient. While migration distance affected nematode beta-diversity across all sites, it had a positive effect on richness at only two of the five study areas suggesting population specific effects. Unexpectedly, nematode richness was higher in winter than summer when very few nematodes were detected. Here we provide the first extensive description of the parasitic nematode community of moose across a wide latitudinal range. Overall, the population-specific impact of migration on parasitism across the distribution range and variation in sympatry with other ruminants suggest local characteristics affect host-parasite relationships.

In Europe, over a third of the agricultural area is grass meadows used for livestock grazing and fodder production. Grass meadows provide a food source for wild ungulates causing human-wildlife conflicts due to forage removal. Few experimental studies have quantified biomass loss with enough replicates to determine how surrounding environments influences level of biomass removal. Using an exclosure experiment on 57 grassland meadows over five years at the northwest coast of Norway covering 10 650 km2, we quantified biomass removal by red deer (Cervus elaphus L.) and how environmental factors impacted biomass loss (Study 1). Furthermore, we examined development of biomass loss and crude protein concentration in five fields throughout the growing season (Study 2). The average predicted biomass loss to red deer grazing was 16

The management of infectious wildlife diseases often involves tackling pathogens that infect multiple host species. Chronic wasting disease (CWD) is a prion disease that can infect most cervid species. CWD was detected in reindeer (Rangifer tarandus) in Norway in 2016. Sympatric populations of red deer (Cervus elaphus) and moose (Alces alces) are at immediate risk. However, the estimation of spillover risk across species and implementation of multispecies management policies are rarely addressed for wildlife. Here, we estimated the broad risk of CWD spillover from reindeer to red deer and moose by quantifying the probability of co-occurrence based on both (1) population density and (2) habitat niche overlap from GPS data of all three species in Nordfjella, Norway. We describe the practical challenges faced when aiming to reduce the risk of spillover through a marked reduction in the population densities of moose and red deer using recreational hunters. This involves setting the population and harvest aims with uncertain information and how to achieve them. The niche overlap between reindeer and both moose and red deer was low overall but occurred seasonally. Migratory red deer had a moderate niche overlap with the CWD-infected reindeer population during the calving period, whereas moose had a moderate niche overlap during both calving and winter. Incorporating both habitat overlap and the population densities of the respective species into the quantification of co-occurrence allowed for more spatially targeted risk maps. An initial aim of a 50% reduction in abundance for the Nordfjella region was set, but only a moderate population decrease of less than 20% from 2016 to 2021 was achieved. Proactive management in the form of marked population reduction is invasive and unpopular when involving species of high societal value, and targeting efforts to zones with a high risk of spillover to limit adverse impacts and achieve wider societal acceptance is important.

Supplemental feeding of cervids during winter is a widespread management practice, but feeding may increase the risk of disease transmission. Therefore, legal regulations to limit supplemental feeding are often implemented when dealing with severe infectious diseases, such as chronic wasting disease (CWD) in cervids. However, it is currently unclear whether these regulations result in decreased spatial clustering and aggregation as intended. Supplemental feeding is expected to restrict the movement of cervids. Therefore, a ban on feeding may also result in wider space use and a risk of geographic spread of disease. The space use of 63 GPS-marked red deer ( Cervus elaphus ) was investigated before ( n = 34) and after ( n = 29) the implementation of a legal regulation aimed at limiting the supplemental feeding of cervids during winter in a CWD-affected region of Nordfjella, Norway. Snow depth was the main determinant of the space use for red deer. A moderate reduction in the number of GPS positions in spatial clusters was evident during periods of deep snow once the ban was in place. Sizes of core areas (Kernel 50%), home ranges (Kernel 95%), and dispersion (MCP 100%, number of 1 km 2 pixels visited per deer) declined from January to March and with increasing snow depth. Dispersion (number of 1 km 2 pixels visited per deer) did not depend on snow depth after the ban, and red deer used larger areas when snow depth was high after the ban compared to before. The ban on supplementary feeding had no effect on size of core areas or home ranges. Several potential factors can explain the overall weak effect of the ban on space use, including the use of agricultural fields by red deer, other anthropogenic feeding, and landscape topography. This study highlights that snow depth is the main factor determining space use during winter, and it remains to be determined whether the moderate reduction in spatial clustering during deep snow after the ban was sufficient to lower the risk of disease transmission.

Coprophagy, the eating of feces, has been documented in a wide range of species but appears to be rare or difficult to detect in deer (Cervidae). Here, we report the first observation of coprophagy in moose Alces alces, which was recorded using camera collars on free-ranging moose in Norway. The footage shows an instance of allocoprophagy by an adult female moose in spring (May). We summarize the current knowledge about coprophagy in deer and briefly discuss potential drivers and possible implications for disease transmission. Further research is needed to determine whether coprophagy occurs frequently in moose and whether this behavior is positive (e.g., increased intake of nutrients) or negative (increased infection by parasites or pathogens).

COVID-19 lockdowns in early 2020 reduced human mobility, providing an opportunity to disentangle its effects on animals from those of landscape modifications. Using GPS data, we compared movements and road avoidance of 2300 terrestrial mammals (43 species) during the lockdowns to the same period in 2019. Individual responses were variable with no change in average movements or road avoidance behavior, likely due to variable lockdown conditions. However, under strict lockdowns 10-day 95th percentile displacements increased by 73%, suggesting increased landscape permeability. Animals' 1-hour 95th percentile displacements declined by 12% and animals were 36% closer to roads in areas of high human footprint, indicating reduced avoidance during lockdowns. Overall, lockdowns rapidly altered some spatial behaviors, highlighting variable but substantial impacts of human mobility on wildlife worldwide.

Data for Agricultural grasslands buffer density effects in red deer populations Initially accepted in Journal of Wildlife Management

Abstract Many wild animals perceive humans as predators, and human disturbance, especially in the form of hunting, triggers antipredatory behavior among prey. Yet, knowledge of how game species react to different types of human disturbance and adapt to repeated disturbances is limited. We investigated how disturbance in the form of a solitary human approacher (stalker) impacted behavior (flight response and short‐term habitat use) of 28 GPS‐collared red deer (Cervus elaphus) in two populations with contrasting population densities in Norway. We studied how the behavioral response differed: (1) with season (pre‐hunting vs. hunting); (2) by consecutive approaches within a day; (3) among replicated experiments within the same season; and (4) between two regions with contrasting densities of red deer. The average flight initiation distance (FID) increased by 15% during the hunting season, and consecutive approaches within the same day caused the red deer to move 49% longer distances. Flight initiation distance was longer in the high‐density population, while escape distance was longer in the low‐density population. Red deer moved out of their weekly home range after 52% approaches, and after the onset of hunting season, time spent outside the home range increased by 89%. Red deer preferred denser resting sites after the disturbance and animal sites had shorter sighting distance and higher canopy cover than control plots. Tree density and canopy cover at animal sites increased at the onset of hunting season, from first to second approach within day, and after replicated experiments within season. Our results suggest that red deer preferred dense resting sites, especially in the hunting season. However, these animal sites had the same amount of the favorable forage plant bilberry (Vaccinium myrtillus), indicating no clear food–cover trade‐off in selection of habitat. Our study showed that onset of hunting initiates stronger fear responses in red deer, which may in turn affect red deer distribution and harvesting efficiency.

Limited mapping of migrations hampers conservation

Herbivores like cervids usually graze on widely scattered forage, but anthropogenic food sources may cause spatial revisitation and aggregation, posing a risk for transmission of infectious diseases. In 2016, chronic wasting disease (CWD) was first detected in Norway. A legal regulation to ban supplemental feeding of cervids and to fence stored hay bales was implemented to lower aggregation of cervids. Knowledge of further patterns and causes of spatial revisitation can inform disease management. We used a recently developed revisitation analysis on GPS-positions from 13 red deer (Cervus elaphus) to identify the pattern of spatial clustering, and we visited 185 spatial clusters during winter to identify the causes of clustering. Anthropogenic food sources were found in 11.9% of spatial clusters, which represented 31.0% of the clusters in agricultural fields. Dumping of silage and hay bales were the main anthropogenic food sources (apart from agricultural fields), and unfenced hay bales were available despite the regulation. The probability of the clusters being in agricultural fields was high during winter. It may be necessary to find other ways of disposing of silage and enforcing the requirement of fencing around hay bales to ensure compliance, in particular during winters with deep snow.

Individual age is an important element in models of population demographics, but the limitations of the methods used for age determination are not always clear. We used known-age data from moose ( Alces alces ), red deer ( Cervus elaphus ), semi-domestic reindeer ( Rangifer tarandus tarandus ) and Svalbard reindeer ( Rangifer tarandus platyrhynchus ) to evaluate the accuracy and repeatability of age estimated by cementum annuli analysis of longitudinally sectioned permanent incisors. Four observers with varying experience performed blind duplicate age estimation of 37 specimens from each cervid. The relationship between known age and estimated age was linear, except for Svalbard reindeer where a quadratic model gave a slightly better fit. After correcting for observer ID and animal ID, there was a slightly declining probability to assess the correct age with increasing age for moose, red deer and Svalbard reindeer. Across cervids and observers, estimated age equalled known age in 69% of all readings, while 95% age ± 1 year. Predicted probability of correct age assessment for experienced observers was 93% for red deer, 89% for Svalbard reindeer, 84% for moose and 73% for semi-domestic reindeer. Regardless of observer experience and cervid, there was a high agreement between repeated assessments of a given animal’s tooth sections. The accuracy varied between cervids but was generally higher for observers with former ageing experience with a given cervid. We conclude that the accuracy of estimated age using longitudinally sectioned incisors is generally high, and even more so if performed by observers with former ageing experience of a given species. To ensure consistency over time, a reference material from known-age individuals for each species analysed should be available for calibration and training of observers.

With climate change, the effect of global warming on snow cover is expected to cause range expansion and enhance habitat suitability for species at their northern distribution limits. However, how this depends on landscape topography and sex in size-dimorphic species remains uncertain, and is further complicated for migratory animals following climate-driven seasonal resource fluctuations across vast landscapes. Using 11 years of data from a partially migratory ungulate at their northern distribution ranges, the red deer (Cervus elaphus), we predicted sex-specific summer and winter habitat suitability in diverse landscapes under medium and severe global warming. We found large increases in future winter habitat suitability, resulting in expansion of winter ranges as currently unsuitable habitat became suitable. Even moderate warming decreased snow cover substantially, with no suitability difference between warming scenarios. Winter ranges will hence not expand linearly with warming, even for species at their northern distribution limits. Although less pronounced than in winter, summer ranges also expanded and more so under severe warming. Summer habitat suitability was positively correlated with landscape topography and ranges expanded more for females than males. Our study highlights the complexity of predicting future habitat suitability for conservation and management of size-dimorphic, migratory species under global warming.