Badgers remain fearless in the face of simulated wolf presence near their setts.

Many mesocarnivores are fossorial and use burrow systems to avoid predators. But fossorial animals cannot stay safely underground forever; they must also risk emerging overground to forage and find mates. To make this trade-off effectively and maximise their own fitness, it is imperative they assess how risk varies in space and time and adapt their denning behaviour accordingly. We used the badger in Bialowieza Forest, Poland, as a model for investigating how the denning behaviour of a fossorial mesocarnivore varies in response to short-term large carnivore risk. To this end, we experimentally simulated perceived wolf presence outside 10 badger setts using audio playbacks of wolves (their howls). We assayed two behavioural measures of fear: badger emergence time from setts on the day playbacks were broadcast and their presence in setts on the day after. We found that neither badger emergence time nor next-day sett use varied in response to wolf playbacks. The results of the present study contrast with a previous study of ours that found badgers used setts in areas with high landscape level perceived wolf risk less often than those in lower-risk areas. Together, these papers' findings suggest that different spatiotemporal scales of perceived risk can have differential effects on badger behaviour. We conclude that rather than take risk avoidance measures at all risky times and places, badgers likely display a diversity of reactions to large carnivore presence that depend on the context and spatiotemporal scale of the risk being perceived.

Wolf risk fails to inspire fear in two mesocarnivores suggesting facilitation prevails.

Large carnivores not only supress mesocarnivores via killing and instilling fear, but also facilitate them through carrion provisioning. Hence, mesocarnivores frequently face a trade-off between risk avoidance and food acquisition. Here we used the raccoon dog and red fox in Bialowieza Forest, Poland as models for investigating how large carnivores shape mesocarnivore foraging behaviour in an area with widespread large carnivore carrion provisioning. Using a giving up density experiment we quantified mesocarnivore foraging responses to wolf body odour across a landscape-scale gradient in wolf encounter rates. At locations with higher wolf encounter rates, raccoon dogs depleted feeding trays more than at plots with lower wolf encounter rates. Simulating wolf presence by adding wolf body odour caused raccoon dogs to deplete feeding trays more at locations with low wolf encounter rates, but less at locations with high wolf encounter rates. Fox foraging costs did not vary with the application of wolf body odour or wolf encounter rates. The frequency that the mesocarnivores visited experimental foraging patches was unaffected by wolf body odour or landscape level encounter rates. These results provide further evidence that large carnivore suppression can play a subordinate role to facilitation in determining mesocarnivore behaviour. The varying raccoon dog response to wolf odour across the landscape-scale gradient in wolf encounter rates shows how mesocarnivore-large carnivore interactions can be context-dependent. We suggest that rather than testing the effects of single risk cues on prey behaviour, future studies should focus on understanding how context modifies the ecological impacts of large carnivores.

Linking spatial patterns of terrestrial herbivore community structure to trophic interactions.

Large herbivores influence ecosystem functioning via their effects on vegetation at different spatial scales. It is often overlooked that the spatial distribution of large herbivores results from their responses to interacting top-down and bottom-up ecological gradients that create landscape-scale variation in the structure of the entire community. We studied the complexity of these cascading interactions using high-resolution camera trapping and remote sensing data in the best-preserved European lowland forest, Bialowieza Forest, Poland. We showed that the variation in spatial distribution of an entire community of large herbivores is explained by species-specific responses to both environmental bottom-up and biotic top-down factors in combination with human-induced (cascading) effects. We decomposed the spatial variation in herbivore community structure and identified functionally distinct landscape-scale herbivory regimes ('herbiscapes'), which are predicted to occur in a variety of ecosystems and could be an important mechanism creating spatial variation in herbivory maintaining vegetation heterogeneity.

Deficiencies in Natura 2000 for protecting recovering large carnivores: A spotlight on the wolf Canis lupus in Poland.

If protected areas are to remain relevant in our dynamic world they must be adapted to changes in species ranges. In the EU one of the most notable such changes is the recent recovery of large carnivores, which are protected by Natura 2000 at the national and population levels. However, the Natura 2000 network was designed prior to their recent recovery, which raises the question whether the network is sufficient to protect the contemporary ranges of large carnivores. To investigate this question we evaluated Natura 2000 coverage of the three wolf Canis lupus populations in Poland. Wolf tracking data showed that wolves have recolonised almost all suitable habitat in Poland (as determined by a recent habitat suitability model), so we calculated the overlap between the Natura 2000 network and all wolf habitat in Poland. On the basis of published Natura 2000 criteria, we used 20% as the minimum required coverage. At the national level, wolves are sufficiently protected (22% coverage), but at the population level, the Baltic and Carpathian populations are far better protected (28 and 47%, respectively) than the endangered Central European Lowland population (12%). As Natura 2000 insufficiently protects the most endangered wolf population in Poland, we recommend expansion of Natura 2000 to protect at least an additional 8% of wolf habitat in western Poland, and discuss which specific forests are most in need of additional coverage. Implementation of these actions will have positive conservation implications and help Poland to fulfil its Habitats Directive obligations. As it is likely that similar gaps in Natura 2000 are arising in other EU member states experiencing large carnivore recoveries, particularly in Central Europe, we make the case for a flexible approach to Natura 2000 and suggest that such coverage evaluations may be beneficial elsewhere.