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, Białowieża 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.

Aim Several large‐mammal species in Europe have recovered and recolonized parts of their historical ranges. Knowing where suitable habitat exists, and thus where range expansions are possible, is important for proactively promoting coexistence between people and large mammals in shared landscapes. We aimed to assess the opportunities and limitations for range expansions of Europe's two largest herbivores, the European bison (Bison bonasus) and moose (Alces alces). Location Central Europe. Methods We used large occurrence datasets from multiple populations and species distribution models to map environmentally suitable habitats for European bison and moose across Central Europe, and to assess human pressure inside the potential habitat. We then used circuit theory modeling to identify potential recolonization corridors. Results We found widespread suitable habitats for both European bison (>120,000 km2) and moose (>244,000 km2), suggesting substantial potential for range expansions. However, much habitat was associated with high human pressure (37% and 43% for European bison and moose, respectively), particularly in the west of Central Europe. We identified a strong east–west gradient of decreasing connectivity, with major barriers likely limiting natural recolonization in many areas. Main conclusions We identify major potential for restoring large herbivores and their functional roles in Europe's landscapes. However, we also highlight considerable challenges for conservation planning and wildlife management, including areas where recolonization likely leads to human–wildlife conflict and where barriers to movement prevent natural range expansion. Conservation measures restoring broad‐scale connectivity are needed in order to allow European bison and moose to recolonize their historical ranges. Finally, our analyses and maps indicate suitable but isolated habitat patches that are unlikely to be colonized but are candidate locations for reintroductions to establish reservoir populations. More generally, our work emphasizes that transboundary cooperation is needed for restoring large herbivores and their ecological roles, and to foster coexistence with people in Europe's landscapes.

Many medium‐sized carnivores are fossorial and use burrow systems to reduce predation risk or avoid predators. But fossorial species cannot stay safely underground forever, and they must also risk emerging overground, to forage and find mates. To make this trade‐off effectively and maximize their own fitness, it is imperative they assess how predation risk varies in space and time, and adapt their denning behaviour accordingly. We used European badger (Meles meles) burrows (setts) in Białowieża Forest, Poland as a model for investigating how the denning behaviour of a fossorial mesocarnivore varies across gradients of landscape scale perceived risk imposed by wolves (Canis lupus) and humans, defined as wolf space use and distance to settlements, respectively. We monitored seventeen setts with varying levels of perceived human and wolf risk with camera traps for two months to study three denning behaviours: frequency of badger sett use, frequency of badger sett sharing with other mesocarnivore species and badger emergence time. Frequency of sett use varied relative to perceived wolf but not human risk. Setts in the highest risk areas were used ca. 60% less often than those in the lowest perceived risk areas, and setts with juveniles were only present in areas of lower perceived risk. Food availability, quantified as earthworm abundance in an area of 2.1 km2 around setts, did not affect the frequency setts were used. Emergence time and frequency of sett sharing with other mesocarnivores, which have been proposed to be anti‐predator strategies, did not vary with either perceived risk factor. These results show fossorial species can adapt their use of burrows to the prevailing risk landscape and suggest burrows deserve more attention in studies on the ecological impacts of apex predators.

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.

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 Białowieża 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.

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Introduced alien species can negatively affect native competitors by reducing their populations or eliminating them from ecosystems. However, studies do not always find evidence for anticipated impacts, and changes in native populations can be difficult to estimate. Interactions between the invasive American mink Neovison vison and native European polecat Mustela putorius have been studied in several countries, but the mink’s impact on polecat populations at a large spatiotemporal scale remains unclear. In the years 1995–2018, we live-trapped mink and polecats at 60 study sites in Poland, and we analysed hunting bags of mink and polecats from the years 2009–2018. During 13,766 trap-nights, we captured 905 individuals. Mink comprised 91.2% and polecats 8.8% of trapped animals. The mean mink and polecat trappability was 6 and 0.6 individuals per 100 trap-nights, respectively. At rivers, polecat and mink trappability were negatively correlated, whereas at lakes, they were not correlated. The sex ratio of trapped polecats was more skewed toward males than that of mink. Mink comprised 63.6% and polecats 36.4% of 59,831 animals killed by hunters. Over 10 years, the numbers of mink shot annually increased slightly, whereas the numbers of polecat decreased slightly. There was a positive correlation between numbers of mink and polecats shot annually. We found weak evidence that at a large spatiotemporal scale, the invasion of mink has led to a decline in polecat numbers. Although the datasets we analysed were based on large samples, they were insufficient to show evidence of competitive interactions between these two mustelids.

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