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Aim The non‐stationarity in habitat selection of expanding populations poses a significant challenge for spatial forecasting. Focusing on the grey wolf (Canis lupus) natural recolonization of Germany, we compared the performance of different distribution modelling approaches for predicting habitat suitability in unoccupied areas. Furthermore, we analysed whether grey wolf showed non‐stationarity in habitat selection in newly colonized areas, which will impact the predictions for potential habitat. Location Germany. Methods Using telemetry data as presence points, we compared the predictive performance of five modelling approaches based on combinations of distribution modelling algorithms—GLMM, MaxEnt and ensemble modelling—and two background point selection strategies. We used a homogeneous Poisson point process to draw background points from either the minimum convex polygons derived from telemetry or the whole area known to be occupied by wolves. Models were fit to the data of the first years and validated against independent data representing the expansion of the species. The best‐performing approach was then used to further investigate non‐stationarity in the species' response in spatiotemporal restricted datasets that represented different colonization steps. Results While all approaches performed similarly when evaluated against a subset of the data used to fit the models, the ensemble model based on integrated data performed best when predicting range expansion. Models for subsequent colonization steps differed substantially from the global model, highlighting the non‐stationarity of wolf habitat selection towards human disturbance during the colonization process. Main Conclusions While telemetry‐only data overfitted the models, using all available datasets increased the reliability of the range expansion forecasts. The non‐stationarity in habitat selection pointed to wolves settling in the best areas first, and filling in nearby lower‐quality habitat as the population increases. Our results caution against spatial extrapolation and space‐for‐time substitutions in habitat models, at least with expanding species.

Aim Several large‐mammal species in Europe have recovered and recolonized parts of their historical ranges. Knowing where suitable habitat exists, and thus where range expansions are possible, is important for proactively promoting coexistence between people and large mammals in shared landscapes. We aimed to assess the opportunities and limitations for range expansions of Europe's two largest herbivores, the European bison (Bison bonasus) and moose (Alces alces). Location Central Europe. Methods We used large occurrence datasets from multiple populations and species distribution models to map environmentally suitable habitats for European bison and moose across Central Europe, and to assess human pressure inside the potential habitat. We then used circuit theory modeling to identify potential recolonization corridors. Results We found widespread suitable habitats for both European bison (>120,000 km2) and moose (>244,000 km2), suggesting substantial potential for range expansions. However, much habitat was associated with high human pressure (37% and 43% for European bison and moose, respectively), particularly in the west of Central Europe. We identified a strong east–west gradient of decreasing connectivity, with major barriers likely limiting natural recolonization in many areas. Main conclusions We identify major potential for restoring large herbivores and their functional roles in Europe's landscapes. However, we also highlight considerable challenges for conservation planning and wildlife management, including areas where recolonization likely leads to human–wildlife conflict and where barriers to movement prevent natural range expansion. Conservation measures restoring broad‐scale connectivity are needed in order to allow European bison and moose to recolonize their historical ranges. Finally, our analyses and maps indicate suitable but isolated habitat patches that are unlikely to be colonized but are candidate locations for reintroductions to establish reservoir populations. More generally, our work emphasizes that transboundary cooperation is needed for restoring large herbivores and their ecological roles, and to foster coexistence with people in Europe's landscapes.

Large carnivores are currently recolonizing Europe following legal protection, but increased mortality in landscapes highly impacted by humans may limit further population expansion. We analyzed mortality and disappearance rates of 35 wolves (of which three emigrated, nine died and 14 disappeared by 1 January 2020) by genetic monitoring in the heavily cultivated and densely populated Jutland peninsula (Denmark and Schleswig‐Holstein, Germany). Annual traffic kill rate estimates ranged from 0.37 (95% CI: 0.11–0.85) to 0.78 (0.51–0.96) in the German part, equivalent to 0.08 (0.02–0.29)–0.25 (0.13–0.46) for the entire region, in the absence of any registered Danish roadkills. In Denmark, annual mortality rate estimates ranged from 0.46 (0.29–0.67) to 0.52 (0.35–0.71), predominantly from cryptic mortality. Despite successful reproductions, we conclude the region is a wolf population sink, primarily driven by cryptic mortality, most likely illegal killing. We hypothesize that frequent encounters between wolves and wolf‐averse persecutors in cultivated landscapes may cause unsustainably high mortality rates despite the majority of hunters respecting protection laws.

Monitoring is a prerequisite for evidence‐based wildlife management and conservation planning, yet conventional monitoring approaches are often ineffective for species occurring at low densities. However, some species such as large mammals are often observed by lay people and this information can be leveraged through citizen science monitoring schemes. To ensure that such wildlife monitoring efforts provide robust inferences, assessing the quantity, quality, and potential biases of citizen science data is crucial. For Eurasian moose (Alces alces), a species currently recolonizing north‐eastern Germany and occurring in very low numbers, we applied three citizen science tools: a mail/email report system, a smartphone application, and a webpage. Among these monitoring tools, the mail/email report system yielded the greatest number of moose reports in absolute and in standardized (corrected for time effort) terms. The reported moose were predominantly identified as single, adult, male individuals, and reports occurred mostly during late summer. Overlaying citizen science data with independently generated habitat suitability and connectivity maps showed that members of the public detected moose in suitable habitats but not necessarily in movement corridors. Also, moose detections were often recorded near roads, suggestive of spatial bias in the sampling effort. Our results suggest that citizen science‐based data collection can be facilitated by brief, intuitive digital reporting systems. However, inference from the resulting data can be limited due to unquantified and possibly biased sampling effort. To overcome these challenges, we offer specific recommendations such as more structured monitoring efforts involving the public in areas likely to be roamed by moose for improving quantity, quality, and analysis of citizen science‐based data for making robust inferences. Citizen science‐based monitoring approaches for moose (Alces alces) in northeast Germany have been analyzed and their characteristics and limitations have been assessed. We provide concrete technical and structural changes to overcome data collection biases and improve the quantity and quality of citizen science data.

Biological invasions provide excellent study systems to understand evolutionary, genetic and ecological processes during range expansions. There is strong evidence for positive effects of high propagule pressure and the associated higher genetic diversity on invasion success, but some species have become invasive despite small founder numbers. The raccoon (Procyon lotor) is often considered as a typical example for such a successful invasion resulting from a small number of founders. The species' largest non-native population in Germany is commonly assumed to stem from a small number of founders and two separate founding events in the 1930s and 1940s. In the present study we analyzed 407 raccoons at 20 microsatellite loci sampled from the invasive range in Western Europe to test if these assumptions are correct. Contrary to the expectations, different genetic clustering methods detected evidence for at least four independent introduction events that gave rise to genetically differentiated subpopulations. Further smaller clusters were either artifacts or resulted from founder events at the range margin and recent release of captive individuals. We also found genetic evidence for on-going introductions of individuals. Furthermore a novel randomization process was used to determine the potential range of founder population size that would suffice to capture all the alleles present in a cluster. Our results falsify the assumption that this species has become widespread and abundant despite being genetically depauperate and show that historical records of species introductions may be misleading.

As the second largest cause of biodiversity loss worldwide, there is an urgent need to study the dynamics of biological invasions and identify factors limiting the distribution of invasive alien species. In the present study we analyze national-scale hunting bag data from Germany to predict the dispersal of raccoons in the largest non-native population of the species. Our focus is (1) to document changes in the distribution and abundance of raccoons, (2) to identify the species–environment relationship and predict which areas will be suitable for future colonization and (3) to apply a dispersal model to predict how fast the raccoon will spread to these areas. The increase from about 9000 harvested raccoons in 2000/01 to about 71,000 in 2011/12 reflects the extensive amount of suitable habitat for this omnivorous species in Central Europe. The best model for explaining range expansion in Germany identified coverage of agriculture and fragmentation and coverage of forests as the most important explanatory variables. The range of raccoons (area with harvest index >0.1 per 100 ha) increased from 26,515 km² in 2001 to 111,630 km² in 2011, and is predicted to expand to 252,940 km² by 2061, 71 % of the area of Germany. This vast area encompasses strategically important areas for conservation biology, such as wetlands with endangered native terrapins. The combination of merging of separated introduced populations and accelerating population growth highlights the potential for future impacts of raccoons on native communities, ecosystems and economic life in Germany and Central Europe.

The products of the genes of the major histocompatibility complex (MHC) are known to be drivers of pathogen resistance and sexual selection enhancing offspring genetic diversity. The MHC further influences individual odour types and social communication. However, little is known about the receptors and their volatile ligands that are involved in this type of chemical communication. Here, we have investigated chemosensory receptor genes that ultimately enable females to assess male genes through odour cues. As a model, we used an invasive population of North American raccoons ( Procyon lotor ) in Germany. We investigated the effect of two groups of chemosensory receptor genes—trace amine-associated receptors (TAARs) and olfactory receptors (ORs)—on MHC-dependent mate choice. Females with more alleles of the TAAR or OR loci were more likely to choose a male with a diverse MHC. We additionally found that MHC class I genes have a stronger effect on mate choice than the recently reported effect for MHC class II genes, probably because of their immunological relevance for viral resistance. Our study is among the first to show a genetic link between behaviour and chemosensory receptor genes. These results contribute to understanding the link between genetics, olfaction and associated life-history decisions.

Livestock farming, which produces milk and meat, is practised by nearly all cultures. But in addition to this, grazing livestock fulfils other fundamentally important roles, one of which is the maintenance of open landscapes, which in turn provides habitat for many rare plant and animal species. Whether wild large herbivores, such as red deer (Cervus elaphus), benefit from the management of the open landscapes is unknown, and there are conflicting views on the nature of the interaction between livestock and wildlife. To investigate this question, we studied interactions in space and time between red deer and livestock in an 787 ha fenced pasture in eastern Germany. First, we used faecal pellet counts to determine whether red deer were using the fenced pasture and whether livestock were using the entire available pasture. Second, we fitted ten red deer, as well as eight Konik horses (Equus ferus caballus) and eight Heck cattle (Bos taurus) with GPS-collars. These data allowed us to perform analyses of space–time interactions. The faecal pellet data showed that red deer did indeed use the fenced pasture and that livestock used the entire extent of the pasture. Further, we verified a neutral space–time interaction between red deer and livestock within the pasture. Thus, there was no indication of a negative effect of the presence of livestock on red deer inside the pasture. For these reasons, we recommend that low-intensity grazing by cattle and horses can maintain open landscapes without detrimental effects on the use of the area by red deer.

Conservation grazing uses semi-feral or domesticated herbivores to limit encroachment in open areas and to promote biodiversity. However, we are still unaware of its effects on wild herbivores. This study investigates the influence of herded sheep and goats on red deer (Cervus elaphus) spatial behavior by testing three a-priori hypotheses: (i) red deer are expected to avoid areas used by livestock, as well as adjacent areas, when livestock are present, albeit (ii) red deer increase the use of these areas when sheep and goats are temporarily absent and (iii) there is a time-lagged disruption in red deer spatial behavior when conservation grazing practice ends. Using GPS-telemetry data on red deer from a German heathland area, we modelled their use of areas grazed by sheep and goats, using mixed-effect logistic regression. Additionally, we developed seasonal resource selection functions (use-availability design) to depict habitat selection by red deer before, during, and after conservation grazing. Red deer used areas less during conservation grazing throughout all times of the day and there was no compensatory use during nighttime. This effect mostly persisted within 21 days after conservation grazing. Effects on habitat selection of red deer were detectable up to 3000 meters away from the conservation grazing sites, with no signs of either habituation or adaption. For the first time, we demonstrate that conservation grazing can affect the spatio-temporal behavior of wild herbivores. Our findings are relevant for optimizing landscape and wildlife management when conservation grazing is used in areas where wild herbivores are present.

The products of the genes of the major histocompatibility complex (MHC) are known to be drivers of pathogen resistance and sexual selection enhancing offspring genetic diversity. The MHC further influences individual odour types and social communication. However, little is known about the receptors and their volatile ligands that are involved in this type of chemical communication. Here, we have investigated chemosensory receptor genes that ultimately enable females to assess male genes through odour cues. As a model, we used an invasive population of North American raccoons (Procyon lotor) in Germany. We investigated the effect of two groups of chemosensory receptor genes—trace amine-associated receptors (TAARs) and olfactory receptors (ORs)—on MHC-dependent mate choice. Females with more alleles of the TAAR or OR loci were more likely to choose a male with a diverse MHC. We additionally found that MHC class I genes have a stronger effect on mate choice than the recently reported effect for MHC class II genes, probably because of their immunological relevance for viral resistance. Our study is among the first to show a genetic link between behaviour and chemosensory receptor genes. These results contribute to understanding the link between genetics, olfaction and associated life-history decisions.