Wherever I may roam-Human activity alters movements of red deer (Cervus elaphus) and elk (Cervus canadensis) across two continents.

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by resource (Normalized Difference Vegetation Index, NDVI) predictability and topography, but those factors to be superseded by human impact.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.

Settle Down! Ranging Behaviour Responses of Roe Deer to Different Capture and Release Methods.

The fitting of tracking devices to wild animals requires capture and handling which causes stress and can potentially cause injury, behavioural modifications that can affect animal welfare and the output of research. We evaluated post capture and release ranging behaviour responses of roe deer (Capreolus capreolus) for five different capture methods. We analysed the distance from the centre of gravity and between successive locations, using data from 14 different study sites within the EURODEER collaborative project. Independently of the capture method, we observed a shorter distance between successive locations and contextual shift away from the home range centre of gravity after the capture and release event. However, individuals converged towards the average behaviour within a relatively short space of time (between 10 days and one month). If researchers investigate questions based on the distance between successive locations of the home range, we recommend (1) initial investigation to establish when the animals start to behave normally again or (2) not using the first two to three weeks of data for their analysis. We also encourage researchers to continually adapt methods to minimize stress and prioritize animal welfare wherever possible, according to the Refinement of the Three R's.

Fine-scale movement patterns and habitat selection of little owls (Athene noctua) from two declining populations.

Advances in bio-logging technology for wildlife monitoring have expanded our ability to study space use and behavior of many animal species at increasingly detailed scales. However, such data can be challenging to analyze due to autocorrelation of GPS positions. As a case study, we investigated spatiotemporal movements and habitat selection in the little owl (Athene noctua), a bird species that is declining in central Europe and verges on extinction in Denmark. We equipped 6 Danish food-supplemented little owls and 6 non-supplemented owls in the Czech Republic with high-resolution GPS loggers that recorded one position per minute. Nightly space use, measured as 95% kernel density estimates, of Danish male owls were on average 62 ha (± 64 SD, larger than any found in previous studies) compared to 2 ha (± 1) in females, and to 3 ± 1 ha (males) versus 3 ± 5 ha (females) in the Czech Republic. Foraging Danish male owls moved on average 4-fold further from their nest and at almost double the distance per hour than Czech males. To create availability data for the habitat selection analysis, we accounted for high spatiotemporal autocorrelation of the GPS data by simulating correlated random walks with the same autocorrelation structure as the actual little owl movement trajectories. We found that habitat selection was similar between Danish and Czech owls, with individuals selecting for short vegetation and areas with high structural diversity. Our limited sample size did not allow us to infer patterns on a population level, but nevertheless demonstrates how high-resolution GPS data can help to identify critical habitat requirements to better formulate conservation actions on a local scale.

Understanding spatio-temporal patterns of deer-vehicle collisions to improve roadkill mitigation.

Vehicles collide with hundreds of thousands of deer on European roads each year. This leads to animal deaths and suffering, economic damage and risks for human safety, making the reduction of road mortality a major field in conservation biology. In order to successfully reduce roadkill, we need improved knowledge regarding spatio-temporal patterns of deer-vehicle collisions (DVCs) on a landscape scale. Here, we analyzed >85,000 DVCs collected over 17 years in Denmark to investigate changes in the number of DVCs over time and to find spatio-temporal patterns of DVC occurrence. We used a use-availability design - originally developed for habitat selection analyses - to compare DVCs involving roe deer (Capreolus capreolus), red deer (Cervus elaphus) and fallow deer (Dama dama) with random road locations on a landscape scale. This approach enabled us to combine temporal (seasonal and diel variation), spatial (land cover, road density and type) and other relevant variables (deer population density, traffic, and deer activity) within the same analysis. We found that factors related to infrastructure and land cover were most important in explaining patterns of DVCs, but seasonal and diel changes, deer activity, and population density were also important in predicting the occurrence of DVCs. Importantly, patterns of DVCs were largely similar between the three deer species, with more DVCs occurring at intermediate traffic density, increasing forest cover, during dusk and dawn, and with increasing deer activity and population density. The strong and consistent patterns found here will allow the development of flexible mitigation measures. We propose that our findings could be used to develop a spatio-temporally flexible warning system for smartphones and navigation systems that is based on existing map providers, making it a widely available and cheap mitigation measure.

Ungulate management in European national parks: Why a more integrated European policy is needed.

1. Primary objectives of national parks usually include both, the protection of natural processes and species conservation. When these objectives conflict, as occurs because of the cascading effects of large mammals (i.e., ungulates and large carnivores) on lower trophic levels, park managers have to decide upon the appropriate management while considering various local circumstances. 2. To analyse if ungulate management strategies are in accordance with the objectives defined for protected areas, we assessed the current status of ungulate management across European national parks using the naturalness concept and identified the variables that influence the management. 3. We collected data on ungulate management from 209 European national parks in 29 countries by means of a large-scale questionnaire survey. Ungulate management in the parks was compared by creating two naturalness scores. The first score reflects ungulate and large carnivore species compositions, and the second evaluates human intervention on ungulate populations. We then tested whether the two naturalness score categories are influenced by the management objectives, park size, years since establishment, percentage of government-owned land, and human impact on the environment (human influence index) using two generalized additive mixed models. 4. In 67.9% of the national parks, wildlife is regulated by culling (40.2%) or hunting (10.5%) or both (17.2%). Artificial feeding occurred in 81.3% of the national parks and only 28.5% of the national parks had a non-intervention zone covering at least 75% of the area. Furthermore, ungulate management differed greatly among the different countries, likely because of differences in hunting traditions and cultural and political backgrounds. Ungulate management was also influenced by park size, human impact on the landscape, and national park objectives, but after removing these variables from the full model the reduced models only showed a small change in the deviance explained. In areas with higher anthropogenic pressure, wildlife diversity tended to be lower and a higher number of domesticated species tended to be present. Human intervention (culling and artificial feeding) was lower in smaller national parks and when park objectives followed those set by the International Union for the Conservation of Nature (IUCN). 5. Our study shows that many European national parks do not fulfil the aims of protected area management as set by IUCN guidelines. In contrast to the USA and Canada, Europe currently has no common ungulate management policy within national parks. This lack of a common policy together with differences in species composition, hunting traditions, and cultural or political context has led to differences in ungulate management among European countries. To fulfil the aims and objectives of national parks and to develop ungulate management strategies further, we highlight the importance of creating a more integrated European ungulate management policy to meet the aims of national parks.

European hares do not avoid newly pesticide-sprayed fields: Overspray as unnoticed pathway of pesticide exposure.

Although risk assessments for pesticides involve quantifying exposure routes for animals, little empirical evidence exists on how individuals use areas where pesticides were used. Further, the European Food Safety Authority guidelines for the risk assessment of birds and mammals currently only include direct dietary uptake from exposed plants as important pathway for pesticides, arguing that dermal exposure is generally negligible. Here, we use the European hare (Lepus europaeus) as a model of a farmland specialist to investigate if (1) hares adjust habitat use in response to pesticide spraying, using GPS data, and (2) calculate the pesticide exposure and uptake over different pathways, i.e. foraging uptake versus overspray and subsequent oral grooming, using an agent-based modelling approach. Apart from avoidance of sprayed fields on the spraying day by inactive hares, which was likely caused by the mechanical disturbance rather than the pesticide itself, we found no evidence that hares reduced the use of pesticide-sprayed fields compared to control observations where no pesticides were applied. Our simulation showed that both the proportion of exposed individuals and the pesticide uptake were related to the pathway of exposure (foraging versus overspray/oral grooming), and depended on the age class and the vegetation type. We found that pesticide uptake via overspray/oral grooming might be 7-fold higher compared to foraging and might thus be an important pathway of pesticide exposure in hares. Our findings emphasize that policy makers, specifically the European Food Safety Authority, should incorporate alternative pathways of pesticide exposure for birds and mammals, such as overspray and oral grooming, when conducting environmental risk assessments and take variation in vegetation structure and age-related animal behavior into account, because these factors might affect pesticide exposure. Interactions between pesticide application and vegetation structure may both increase or decrease exposure but can be predicted using a simulation approach.

Discussion: Illegal kills of protected wolves call for public reasoning.

Illegal wolf kills happens around in Europe despite the European wolf is protected under the EU Habitats Directive. The reason for this is conflicts with farmers and local hunters and in some instances also direct fear. In April 2018, a wolf was killed in Denmark after 1st recolonization since the 18th century. This caused a heated debate and calls for better communication and management of the Danish and entire European wolf population. Here we discuss the challenges of illegal wolf kills and call for European governments to take action. We specifically encourage European governments to create facilitated spaces for public deliberation on wildlife management by integrating facts and values, not separating them.

Habitat selection by the European hare in arable landscapes: The importance of small-scale habitat structure for conservation.

Agricultural land-use practices have intensified over the last decades, leading to population declines of various farmland species, including the European hare (Lepus europaeus). In many European countries, arable fields dominate agricultural landscapes. Compared to pastures, arable land is highly variable, resulting in a large spatial variation of food and cover for wildlife over the course of the year, which potentially affects habitat selection by hares. Here, we investigated within-home-range habitat selection by hares in arable areas in Denmark and Germany to identify habitat requirements for their conservation. We hypothesized that hare habitat selection would depend on local habitat structure, that is, vegetation height, but also on agricultural field size, vegetation type, and proximity to field edges. Active hares generally selected for short vegetation (1-25 cm) and avoided higher vegetation and bare ground, especially when fields were comparatively larger. Vegetation >50 cm potentially restricts hares from entering parts of their home range and does not provide good forage, the latter also being the case on bare ground. The vegetation type was important for habitat selection by inactive hares, with fabaceae, fallow, and maize being selected for, potentially providing both cover and forage. Our results indicate that patches of shorter vegetation could improve the forage quality and habitat accessibility for hares, especially in areas with large monocultures. Thus, policymakers should aim to increase areas with short vegetation throughout the year. Further, permanent set-asides, like fallow and wildflower areas, would provide year-round cover for inactive hares. Finally, the reduction in field sizes would increase the density of field margins, and farming different crop types within small areas could improve the habitat for hares and other farmland species.

Common Starlings (Sturnus vulgaris) increasingly select for grazed areas with increasing distance-to-nest.

The abundant and widespread Common Starling (Sturnus vulgaris) is currently declining across much of Europe due to landscape changes caused by agricultural intensification. The proximate mechanisms causing adverse effects to breeding Starlings are unclear, hampering our ability to implement cost-efficient agri-environmental schemes to restore populations to former levels. This study aimed to show how this central foraging farmland bird uses and selects land cover types in general and how use of foraging habitat changes in relation to distance from the nest. We attached GPS-loggers to 17 breeding Starlings at a Danish dairy cattle farm in 2015 and 2016 and analysed their use of different land cover types as a function of distance intervals from the nest and their relative availability. As expected for a central place forager, Starlings increasingly avoided potential foraging areas with greater distance-to-nest: areas ≥ 500 m were selected > 100 times less frequently than areas within 100 m. On average, Starlings selected the land cover category Grazed most frequently, followed by Short Grass, Bare Ground, Meadow and Winter Crops. Starlings compensated for elevated travel costs by showing increasing habitat selection the further they foraged from the nest. Our results highlight the importance of Grazed foraging habitats close to the nest site of breeding Starlings. The ecological capacity of intensively managed farmlands for insectivorous birds like the Starling is decreasing through conversion of the most strongly selected land cover type (Grazed) to the least selected (Winter Crops) which may be further exacerbated through spatial segregation of foraging and breeding habitats.

Using population viability analysis, genomics, and habitat suitability to forecast future population patterns of Little Owl Athene noctua across Europe.

The agricultural scene has changed over the past decades, resulting in a declining population trend in many species. It is therefore important to determine the factors that the individual species depend on in order to understand their decline. The landscape changes have also resulted in habitat fragmentation, turning once continuous populations into metapopulations. It is thus increasingly important to estimate both the number of individuals it takes to create a genetically viable population and the population trend. Here, population viability analysis and habitat suitability modeling were used to estimate population viability and future prospects across Europe of the Little Owl Athene noctua, a widespread species associated with agricultural landscapes. The results show a high risk of population declines over the coming 100 years, especially toward the north of Europe, whereas populations toward the southeastern part of Europe have a greater probability of persistence. In order to be considered genetically viable, individual populations must count 1,000-30,000 individuals. As Little Owl populations of several countries count <30,000, and many isolated populations in northern Europe count <1,000 individuals, management actions resulting in exchange of individuals between populations or even countries are probably necessary to prevent losing <1% genetic diversity over a 100-year period. At a continental scale, a habitat suitability analysis suggested Little Owl to be affected positively by increasing temperatures and urban areas, whereas an increased tree cover, an increasing annual rainfall, grassland, and sparsely vegetated areas affect the presence of the owl negatively. However, the low predictive power of the habitat suitability model suggests that habitat suitability might be better explained at a smaller scale.

Weather conditions drive dynamic habitat selection in a generalist predator.

Despite the dynamic nature of habitat selection, temporal variation as arising from factors such as weather are rarely quantified in species-habitat relationships. We analysed habitat use and selection (use/availability) of foraging, radio-tagged little owls (Athene noctua), a nocturnal, year-round resident generalist predator, to see how this varied as a function of weather, season and availability. Use of the two most frequently used land cover types, gardens/buildings and cultivated fields varied more than 3-fold as a simple function of season and weather through linear effects of wind and quadratic effects of temperature. Even when controlling for the temporal context, both land cover types were used more evenly than predicted from variation in availability (functional response in habitat selection). Use of two other land cover categories (pastures and moist areas) increased linearly with temperature and was proportional to their availability. The study shows that habitat selection by generalist foragers may be highly dependent on temporal variables such as weather, probably because such foragers switch between weather dependent feeding opportunities offered by different land cover types. An opportunistic foraging strategy in a landscape with erratically appearing feeding opportunities in different land cover types, may possibly also explain decreasing selection of the two most frequently used land cover types with increasing availability.