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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.
As wolves recolonize areas of Europe ranging from moderate to high anthropogenic impact, fear of wolves is a recurring source of conflict. Shared tools for evaluating wolf responses to humans, and comparing such responses across their range, can be valuable. Experiments in which humans approach wild wolves can increase our understanding of how wolves respond to humans, facilitating human-wolf coexistence. We have developed the first standardized protocol for evaluating wolf responses to approaching humans using high-resolution GPS data, and tested it on wild wolves. We present a field protocol for experimentally approaching GPS-collared wolves, a descriptive comparison of two statistical methods for detecting a measurable flight response, a tutorial for identifying wolf flight initiation and resettling positions, and an evaluation of the method when reducing GPS positioning frequency. The field protocol, a data collection form, and the tutorial with R code for extracting flight parameters are provided. This protocol will facilitate studies of wolf responses to approaching humans, applicable at a local, national, and international level. Data compiled in a standardized way from multiple study areas can be used to quantify the variation in wolf responses to humans within and between populations, and in relation to predictors such as social status, landscape factors, or human population density, and to establish a baseline distribution of wolf response patterns given a number of known predictors. The variation in wolf responses can be used to assess the degree to which results can be generalized to areas where GPS studies are not feasible, e.g., for predicting the range of likely wolf behaviors, assessing the likelihood of wolf-human encounters, and complementing existing tools for evaluating reports of bold wolves. Showing how wolves respond to human encounters should help demystify the behavior of wild wolves toward humans in their shared habitat.