Moose (Alces alces) browsing patterns in recently planted clear-cut areas in relation to predation risk of the gray wolf (Canis lupus) in Sweden

The aim of this study was to test whether behaviorally mediated effects of gray wolf (Canis lupus) presence were visible in terms of browsing intensity by moose (Alces alces) on tree saplings in recently planted clear-cut areas in Sweden. The study involved two different spatial scales; the clear-cut scale and tree scale. At the clear-cut scale the influence of wolf predation risk, tree density and tree height on moose browsing intensity was evaluated. At the individual tree scale, the factor distance to the nearest forest edge was also included. Browsing intensity was measured as apical shoot browsing and lateral shoot browsing in 24 different recently planted clear-cuts. Due to differences in preference between tree species and qualitative differences between plant parts, the same analyses were conducted for the five main tree species and for apical and lateral shoot browsing separately. Based on pellet group counts, moose is most likely the main browser in this study. Further, I found that both tree density and tree height were lower in the wolf core areas, suggesting multicollinearity between wolf utilization, tree density and tree height at the clear-cut scale, the effect of these factors could not be separated in the multivariate models. At the clear-cut scale browsing intensity decreased with tree density. At the tree scale, although wolf utilization was an important factor explaining the variation in browsing intensity, the browsing probability decreased with tree height. Overall, this suggests that moose foraging and thus its browsing intensity was more influenced by tree density and tree height than predation risk by wolves. It also seemed that moose browse through a clear-cut area irrespectively of distance to the closest forest edge

Olfactory cues of large carnivores modify red deer behavior and browsing intensity

This study examined the effect of perceived predation risk imposed by lynx (Lynx lynx) and wolf (Canis lupus) on red deer (Cervus elaphus) foraging behavior under experimental conditions. We hypothesized that in response to large carnivore scent red deer would increase their vigilance, although reducing the frequency and duration of visits to foraging sites. Consequently, browsing intensity on tree saplings was expected to decrease, whereas a higher proportion of more preferred species was expected to be browsed to compensate for higher foraging costs. We expected stronger responses towards the ambush predator lynx, compared with the cursorial predator wolf. These hypotheses were tested in a cafeteria experiment conducted within three red deer enclosures, each containing four experimental plots with olfactory cues of wolf, lynx, cow, and water as control. On each plot, a camera trap was placed and browsing intensity was measured for one consecutive week, repeated three times. Red deer reduced their visitation duration and browsing intensity on plots with large carnivore scent. Despite red deer showing a clear preference for certain tree species, the presence of large carnivore scent did not change selectivity towards different tree species. Contrary to our hypothesis, we found more pronounced effects of wolf (cursorial) compared with lynx (ambush). This study is the first to experimentally assess the perceived risk effects on the red deer foraging behavior of large carnivores differing in hunting modes. Our findings provide insights into the role of olfactory cues in predator-prey interactions and how they can modify fine-scale herbivore-plant interactions.publishedVersio

Humans rather than Eurasian lynx (Lynx lynx) shape ungulate browsing patterns in a temperate forest

The recolonization of human-dominated landscapes by large carnivores has been followed with considerable scientific interest; however, little is known about their interactive effect on ungulate foraging behavior. This study compared the risks imposed by humans and lynx on ungulate foraging behavior by examining the effects of browsing intensity (at two spatial scales), diet quality, and tree species selection. We hypothesized that: (1) in areas with high risk imposed by humans and lynx browsing intensity would be reduced; (2) risk effects would interact with habitat visibility at a fine scale, resulting in contrasting browsing patterns in response to humans versus lynx risk; (3) ungulates compensate for the higher costs incurred in high-risk areas by switching to a higher diet quality, and (4) browse a higher proportion of more-preferred tree species. These hypotheses were tested by measuring browsing intensity along 48 transects located at different distances from human settlements within the hunted and nonhunted areas of the Bavarian Forest. Dung samples were collected and analyzed as a proxy of diet quality (C:N ratio, fiber). The spatial patterns of browsing intensity, diet quality, and tree species selection were then linked to lynx risk, hunting intensity, recreation intensity, and distance to human settlements. Our results showed that (1) browsing intensity strongly decreased with increasing recreational activities, whereas it increased with lynx risk; (2) only in close proximity to human settlements tree browsing was higher in dense habitats and (3) a higher diet quality was obtained. (4) We found a stronger avoidance of the less preferred tree species in high-hunting intensity areas. In conclusion, our results indicate that the risk effects of human activities outweigh those of a natural large carnivore. Thus, highlighting the importance of taking those activities into account in predicting the impacts of large carnivores on ungulates and their plant-food choices.publishedVersio

Mammal responses to predator scents across multiple study areas

Antipredator behaviors allow prey to mitigate the impacts of their predators. We investigated antipredator responses of two herbivore species, roe deer and European hare, and one mesopredator, red fox, toward predation risk imposed by lynx and wolf. We collected data (using camera traps) on visitation frequency and vigilance behavior to olfactory predator stimuli during 158 standardized scent trials in five areas across Europe, where lynx and wolves either occurred or had been absent for centuries. After a period without scent, trial sites were either marked with lynx or wolf urine, or butyric acid (unspecific scent used to contrast species-specific scent responses). We expected the two herbivores to respond aversively (reduced visitation frequency, and increased vigilance) to predator urine, while red foxes (scavengers) might adopt a risk-sensitive exploration strategy by increasing vigilance near predator urine without reducing visitation frequency. For all species, we expected stronger responses toward the ambush predator lynx than to the pursue predator wolf (cryptic predator hypothesis). If prey responds more strongly to predator stimuli when coexisting with the predator, we expected stronger reactions to predator urine in areas where predators occurred (naïve prey hypothesis). Roe deer significantly avoided lynx urine and butyric acid, but not wolf urine. However, roe deer visitation frequency did not differ between scent treatment with large carnivore urine and butyric acid (suggesting that roe deer generally avoid areas with unfamiliar scent), or between areas where large carnivores were present and absent. Hares did not significantly avoid predator urine. Red foxes were attracted to lynx urine in sympatry, but not in allopatry with large carnivores. They increased vigilance rates in the presence of lynx urine independent of sympatry/ allopatry with large carnivores. These findings generally confirm our expectations of predator avoidance by herbivores, and attraction combined with increased vigilance of mesopredators. In all species, lynx urine elicited stronger responses than wolf urine (although not significantly different), which to some extent conforms to the cryptic predator hypothesis. We found no support for the naïve prey hypothesis. However, higher attraction of red foxes to lynx urine in sympatric situations might indicate that positive responses by scavengers are learning based.publishedVersio

Analysis of differences and commonalities in wildlife hunting across the Africa-Europe South-North gradient

Hunting and its impacts on wildlife are typically studied regionally, with a particular focus on the Global South. Hunting can, however, also undermine rewilding efforts or threaten wildlife in the Global North. Little is known about how hunting manifests under varying socioeconomic and ecological contexts across the Global South and North. Herein, we examined differences and commonalities in hunting characteristics across an exemplary Global South-North gradient approximated by the Human Development Index (HDI) using face-to-face interviews with 114 protected area (PA) managers in 25 African and European countries. Generally, we observed that hunting ranges from the illegal, economically motivated, and unsustainable hunting of herbivores in the South to the legal, socially and ecologically motivated hunting of ungulates within parks and the illegal hunting of mainly predators outside parks in the North. Commonalities across this Africa-Europe South-North gradient included increased conflict-related killings in human-dominated landscapes and decreased illegal hunting with beneficial community conditions, such as mutual trust resulting from community involvement in PA management. Nevertheless, local conditions cannot outweigh the strong effect of the HDI on unsustainable hunting. Our findings highlight regional challenges that require collaborative, integrative efforts in wildlife conservation across actors, while identified commonalities may outline universal mechanisms for achieving this goal.publishedVersio

Data_file_manuscript

This data file contains raw data collected at the plantation scale, within-plantation scale and the raw data from the number of pellet groups collected

Olfactory cues of large carnivores modify red deer behavior and browsing intensity

This study examined the effect of perceived predation risk imposed by lynx (Lynx lynx) and wolf (Canis lupus) on red deer (Cervus elaphus) foraging behavior under experimental conditions. We hypothesized that in response to large carnivore scent red deer would increase their vigilance, although reducing the frequency and duration of visits to foraging sites. Consequently, browsing intensity on tree saplings was expected to decrease, whereas a higher proportion of more preferred species was expected to be browsed to compensate for higher foraging costs. We expected stronger responses towards the ambush predator lynx, compared with the cursorial predator wolf. These hypotheses were tested in a cafeteria experiment conducted within three red deer enclosures, each containing four experimental plots with olfactory cues of wolf, lynx, cow, and water as control. On each plot, a camera trap was placed and browsing intensity was measured for one consecutive week, repeated three times. Red deer reduced their visitation duration and browsing intensity on plots with large carnivore scent. Despite red deer showing a clear preference for certain tree species, the presence of large carnivore scent did not change selectivity towards different tree species. Contrary to our hypothesis, we found more pronounced effects of wolf (cursorial) compared with lynx (ambush). This study is the first to experimentally assess the perceived risk effects on the red deer foraging behavior of large carnivores differing in hunting modes. Our findings provide insights into the role of olfactory cues in predator-prey interactions and how they can modify fine-scale herbivore-plant interactions

Data from: Does wolf presence reduce moose browsing intensity in young forest plantations?

Large carnivores can be a key factor in shaping their ungulate prey’s behavior, which may affect lower trophic levels. While most studies on trade-offs between food acquisition and risk avoidance by ungulate prey species have been conducted in areas with limited human impact, carnivores are now increasingly returning to highly anthropogenic landscapes. Many of these landscapes are dominated by forestry, and ungulate-forestry conflicts are an increasing issue. The aim of this study was to test if the indirect effects of a re-colonizing large predator, the wolf (Canis lupus), results in a change in browsing intensity by moose (Alces alces) in young forest plantations in a boreal forest in Sweden. We selected 24 different forest plantations, with 12 located in low-wolf and 12 in high-wolf utilization areas. In each plantation, we measured browsing intensity, tree height, tree density, distance to the closest forest edge and we counted the number of moose pellet groups. In contrast to our predictions, wolf utilization was not the main driver of moose browsing patterns. Instead, moose browsing intensity declined with tree density and height. Separate analyses on the main tree species showed that wolf utilization had an influence, but browsing intensity was in fact higher in the high-wolf utilization areas for three out of five tree species. This pattern seemed to be driven by a strong confounding relationship between wolf utilization, tree density and height, which were both lower in the high-wolf utilization areas. We argue that this confounding effect is due to wolves being pushed towards the less productive parts of the landscape away from human activity centers. Therefore, we concluded that in order to better understand carnivore driven risk- mediated effects on herbivore behavior in anthropogenic landscapes we need to better understand the complexity of human-carnivore-prey-ecosystem interactions

Humans rather than Eurasian lynx (Lynx lynx) shape ungulate browsing patterns in a temperate forest

The recolonization of human-dominated landscapes by large carnivores has been followed with considerable scientific interest; however, little is known about their interactive effect on ungulate foraging behavior. This study compared the risks imposed by humans and lynx on ungulate foraging behavior by examining the effects of browsing intensity (at two spatial scales), diet quality, and tree species selection. We hypothesized that: (1) in areas with high risk imposed by humans and lynx browsing intensity would be reduced; (2) risk effects would interact with habitat visibility at a fine scale, resulting in contrasting browsing patterns in response to humans versus lynx risk; (3) ungulates compensate for the higher costs incurred in high-risk areas by switching to a higher diet quality, and (4) browse a higher proportion of more-preferred tree species. These hypotheses were tested by measuring browsing intensity along 48 transects located at different distances from human settlements within the hunted and nonhunted areas of the Bavarian Forest. Dung samples were collected and analyzed as a proxy of diet quality (C:N ratio, fiber). The spatial patterns of browsing intensity, diet quality, and tree species selection were then linked to lynx risk, hunting intensity, recreation intensity, and distance to human settlements. Our results showed that (1) browsing intensity strongly decreased with increasing recreational activities, whereas it increased with lynx risk; (2) only in close proximity to human settlements tree browsing was higher in dense habitats and (3) a higher diet quality was obtained. (4) We found a stronger avoidance of the less preferred tree species in high-hunting intensity areas. In conclusion, our results indicate that the risk effects of human activities outweigh those of a natural large carnivore. Thus, highlighting the importance of taking those activities into account in predicting the impacts of large carnivores on ungulates and their plant-food choices

The blame game: Using eDNA to identify species-specific tree browsing by red deer (Cervus elaphus) and roe deer (Capreolus capreolus) in a temperate forest

Increasing deer populations in many temperate regions can affect tree regeneration, resulting in severe long-term impacts on forest structure, composition and diversity. Of the most common deer species in Europe — red deer (Cervus elaphus) and roe deer (Capreolus capreolus) — roe deer are generally thought to have the highest impact on palatable tree species owing to their feeding niche. Although browsing and its potential consequences are well researched, less is known about the influence of specific deer species within multi-species ungulate communities on specific tree species. Environmental DNA (eDNA) allows the determination of species-specific browsing habits without the need for direct observations, facilitating effective targeting of management interventions. In this study eDNA was used to elucidate the browsing patterns of these two deer species in the temperate forest of the Bavarian Forest National Park, Germany and analysed the influence of tree species, management type and height of browsing, on the success rate of the method. Samples were collected from twigs used in feeding trials from enclosures containing red deer or roe deer and from naturally browsed twigs in three different management types within the national park. eDNA was successfully amplified from 98% of the feeding trial samples, and the correct deer species was identified for all samples. eDNA was successfully amplified from approximately 50% of the naturally browsed samples. Neither management type, tree species, nor height of browsing had any significant influence on the success of the method. For silver fir and rowan, no significant difference was found in the proportion of browsing events attributable to roe or red deer, when the two deer species occur at similar densities. These results indicate that roe deer might not always be disproportionately responsible for the browsing of palatable tree species as expected from its food niche. Roe deer were significantly more responsible for browsing at lower heights than red deer. Although not statistically significant, roe deer were more responsible for browsing in intact forest compared to bark-beetle-impact forest, with the opposite relationship for red deer