Knowledge concerning the relative importance of biotic and abiotic factors associated with the long-term settlement of large carnivores in human-dominated landscapes is important for effective species conservation. However, identifying the major drivers of large carnivore occurrences at a continental scale is challenging as several factors can act in concert to affect large carnivore distributions. Thus, we investigated the relationship between the permanent and sporadic occurrences of large carnivores (brown bear, Eurasian lynx, grey wolf and wolverine) and three groups of predictor variables (i.e., landscape attributes, human disturbance and prey availability) on the continental scale in Europe. Specifically, we used generalized linear models (GLMs) and variation partitioning to estimate the independent and cumulative effects of the three predictor groups on large carnivore occurrences. The explained variance for permanent vs. sporadic occurrence was highest for the wolverine (42.8% and 29.7%), followed by the brown bear (20.5% and 16.4%), Eurasian lynx (15.2% and 11.6%) and grey wolf (15.8% and 6.1%). Landscape attributes, such as forest cover, shrub-land cover, altitude and slope, were positively related to the occurrence of grey wolf, brown bear and Eurasian lynx, whereas human disturbance (human population density, distance to roads and to human settlements) was negatively related to wolverine occurrence. For all species, shared effects between landscape attributes and human disturbance accounted for a considerable portion of the explained variation in both permanent and sporadic occurrence, and landscape attributes were generally more important for explaining permanent than sporadic occurrence, except for the wolverine. Prey availability was marginally associated with the permanent occurrence of the grey wolf but we found no statistical effect of prey availability on the occurrence of the other large carnivores. In conclusion, the sporadic occurrence of large carnivores in Europe is more stochastic and less predictable than their permanent occurrence. Landscape attributes and their joint effects with human disturbance are the most important factors related to the recolonization dynamics of large carnivores, although wolverine distributions appear strongly limited by human disturbance itself. Domestic prey availability seems to play a subordinate role in driving the recent distribution dynamics of large carnivores at the continental scale. Thus, our results are relevant also in light of the ongoing recolonization of large carnivores which, in many rural areas, caused concerns among the locals, especially in relation to conflicts with human activities. For these reasons, we stress the need to continue and improve large carnivore monitoring across Europe.
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