Large carnivores can be found in different scenarios of cohabitation with humans. Behavioral adaptations to minimize risk from humans are expected to be exacerbated where large carnivores are most vulnerable, such as at breeding sites. Using wolves as a model species, along with data from 26 study areas across the species´ worldwide range, we performed a meta-analysis to assess the role of humans in breeding site selection by a large carnivore. Some of the patterns previously observed at the local scale become extrapolatable to the entire species range provided that important sources of variation are taken into account. Generally, wolves minimised the risk of exposure at breeding sites by avoiding human-made structures, selecting shelter from vegetation and avoiding agricultural lands. Our results suggest a scaled hierarchical habitat selection process across selection orders by which wolves compensate higher exposure risk to humans within their territories via a stronger selection at breeding 3 sites. Dissimilar patterns between continents suggest that adaptations to cope with human-associated risks are modulated by the history of coexistence and persecution. Although many large carnivores persisting in human-dominated landscapes do not require large-scale habitat preservation, habitat selection at levels below occupancy and territory should be regarded in management and conservation strategies aiming to preserve these species in such contexts. In this case, we recommend providing shelter from human interference at least in small portions of land in order to fulfill the requirements of the species to locate their breeding sites.

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Highly mobile mammalian carnivores are expected to have the capability to maintain high levels of gene flow across large geographic scales. Nonetheless, surprising levels of genetic structure have been found in many such populations. We combined genetic and spatial behavioural information from wolves (Canis lupus) in the Iberian Peninsula (Western Europe) during the last two decades to present a particular case of low dispersal levels in a large carnivore population persisting in human-dominated landscapes. We found an exceptionally reticulated pattern of cryptic population structure emerging at two hierarchical levels, in which four or eleven meaningful genetic clusters can be recognized, respectively. These clusters were characterized by moderate-high levels of differentiation (average pairwise F = 0.09-0.19), low levels of admixture and varying degrees of genetic diversity. The number of dispersers identified among the 11 clusters was very low (<4% out of 218 wolves). Spatial information of tracked wolves further confirmed the geographical genetic patterns (only 2 out of 85 collared wolves overlapped with more than one genetic cluster). The high levels of genetic structure in this population may be determined by the recent demographic history of this population, among other factors. The identification of meaningful genetic clusters has implications for the delineation of conservation units and, consequently, on the conservation and management actions for Iberian wolves.

Despite severe population declines and an overall range contraction, some populations of large carnivores have managed to survive in human-modified landscapes. From a conservation perspective, it is important to identify the factors allowing for this coexistence, including the relevant habitat characteristics associated with the presence of large carnivores. We evaluated the role of several environmental factors describing habitat quality for wolves Canis lupus in the humanised Iberian Peninsula, which currently holds an important wolf population at European level. We used maximum entropy and generalized linear model approaches in a nestedscale design to identify the environmental factors that are related to wolf presence at three spatial scales and resolutions: (1) distribution range: wolf presence on a 10 9 10 km grid resolution, (2) wolf habitat use: wolf occurrence on a 2 9 2 km grid and (3) dens/rendezvous sites: breeding locations on a 1 9 1 km grid. Refuge availability, as defined by topography, seemed to be the key factor determining wolf presence at the multiple scales analysed. As a result, wolf populations may coexist with humans in modified landscapes when the topography is complex. We found that a significant amount of favourable habitat is not currently occupied, suggesting that the availability of suitable habitat is not the limiting factor for wolves in the Iberian Peninsula. Habitat suitability outside the current range indicates that other factors, such as direct persecution and other sources of anthropogenic mortality, may be hampering its expansion. We suggest that priorities for conservation should follow two general lines: (1) protect good quality habitat within the current range; and (2) allow dispersal to unoccupied areas of good quality habitat by reducing human-induced mortality rates. Finally, we still need to improve our understanding of how wolves coexist with humans in modified landscapes at fine spatiotemporal scales, including its relationship with infrastructures, land uses and direct human presence.

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