For wolves that were not observed for more than 1 year, we assumed that their continued presence was highly unlikely, and estimated the likely day of disappearance as the last observation date plus the average observation interval until its disappearance, adding those days to the last observation (Sunde et al., 2021). Accordingly, if the second registration in grid cell B was not based on a predation event (e.g., feces), we recorded 5 days without livestock depredation for grid cell B. If the observation in grid cell A was based on a DNA profile from a sheep kill, we entered the data as one observation day with livestock depredation and nine observation days without livestock depredation. For example, if a wolf was first observed in grid cell A, and 10 days later in grid cell B to appear in grid cell C after another 5 days, we allocated ten observation days to grid cell A and five observation days to grid cell B. 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. Despite successful reproductions, we conclude the region is a wolf population sink, primarily driven by cryptic mortality, most likely illegal killing.

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. 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. 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). Large carnivores are currently recolonizing Europe following legal protection, but increased mortality in landscapes highly impacted by humans may limit further population expansion.

Wolf gone wild