Recreation and hunting differentially affect deer behaviour and sapling performance

Humans are increasingly acknowledged as apex predators that shape landscapes of fear to which herbivores adapt their behaviour. Here, we investigate how humans modify deer space-use and their effects on vegetation at two spatial scales; zones with different types of human use (largescale risk factor) and, nested within that, trails (fine-scale risk factor). In zones with three contrasting types of human activities: 1) no recreation, no hunting, 2) with recreation, no hunting and 3) with recreation and hunting, we linked deer space-use (dropping counts) to browsing intensity, relative growth and survival of planted saplings. Plots were located at two distances to trails (20 versus 100 m) to test how trails affect deer space-use and sapling performance. Additionally, plots were distributed over forest and heathland as risk effects are habitat-dependent. Deer space-use was highest in the zone without recreation or hunting, resulting in higher browsing levels and lower sapling growth and survival, but only in heathland. In contrast, deer space-use and sapling performance did not differ between zones with recreation only and zones with recreation and hunting. Deer dropping counts were lower near trails used for recreation, but this was not associated with browsing impact or sapling performance. Our results show that recreational use modifies deer space-use which is associated with browsing impact on woody vegetation, while seasonal hunting activities in zones with recreation did not have additive year-round effects. Yet, effects were only observed at the larger scale of recreation zones and not near trails. Furthermore, deer space-use was only associated with sapling performance in open heathland, where high visibility presumably increases avoidance behaviour because it increases detectability and decreases escape possibilities. This suggests that recreation creates behaviourally mediated cascading effects that influence vegetation development, yet these effects are context-dependent. We advocate incorporating human-induced fear effects in conservation, management and research

Impediments affect deer foraging decisions and sapling performance

Impediments, such as tree logs, can prevent access to saplings for deer, and can increase perceived predation risk by blocking view and escape possibilities for deer in areas with large carnivores. Therefore, impediments can influence deer foraging decisions and the trade-off between safety and food of different quality indirectly influencing tree regeneration. The aim of our study was to test how the presence of an impediment affects deer foraging behavior and tree sapling performance of eight species that differ in preference by deer. We planted saplings without, nearby and inside impediments and followed their fate for three consecutive years in Białowieża forest, Poland. We constructed 1 m high impediments of 5 × 5 m that would still allow deer to enter the impediment and forage from the saplings planted inside. However, we never recorded deer inside the impediments. Near the impediments deer visitation rate and cumulative visitation time was reduced. As a result, browsing intensity of all tree species was lower nearby and especially inside the impediment. Deer did not select different tree species without, near or inside the impediment. Due to the overall lower browsing intensity, tree saplings increased in height near the impediment and heights increased significantly within the impediment. The palatable, but browsing intolerant, Acer platanoides benefited most from the impediment as this species was highly selected and heavily browsed without an impediment. Followed by the palatable, but more browse tolerant Tilia cordata and Pyrus pyraster. In comparison, the presence of an impediment had a smaller effect on the less preferred Alnus glutinosa, Picea abies and Pinus sylvestris which survived well without an impediment. Our study showed that impediments modified deer behavior as they visited these plots less frequently and thereby indirectly reduced the browsing impact on the preferred tree species. Therefore, the potential for successful recruitment of preferred tree species is higher near an impediment, and especially when surrounded by impediment structures. In the long-term, the presence of natural impediments like tree logs allows browsing intolerant tree species to escape browsing, ultimately leading to a more diverse forest composition

Long-term decline in a salt marsh hare population largely driven by bottom-up factors

The widespread decline of the European brown hare (Lepus europaeus) in Europe has been attributed to both bottom-up and top-down factors, as well as climate change. Few studies have attempted to study the relative importance of these factors considered simultaneously. In this study we tested the hypotheses that hare population density is regulated by bottom-up (food), top-down (predation) or abiotic factors including tidal floods and climatic conditions. We related data on hare population density on a relatively isolated island to changes in surface area of suitable vs. unsuitable vegetation for forage, predator densities, flooding parameters and climatic variables. During the study period (1996–2012), hare numbers decreased from 580 to 219. Estimated population density was positively correlated with the cover of short, intermediate successional vegetation types and was negatively correlated with the cover of tall, late successional vegetation types. These findings corroborate results from earlier experimental studies that reported a strong aversion of hares to tall vegetation. Additionally, we found indications that raptor population density and unusually high floods also exerted negative effects on hare population density. We conclude that bottom-up factors (the availability of suitable forage) are the main regulators of the studied hare population. This suggests that the importance of bottom-up effects has been underestimated and could explain leporid population decline in areas that have experienced a similar increase in tall, unsuitable vegetation

Data from: Do wild ungulates experience higher stress with humans than with large carnivores?

Predation is a major selective pressure for prey; however, the stress response to predation risk and the relative importance of natural versus anthropogenic stress factors in wild populations of animals have rarely been studied. We investigated the level of fecal glucocorticoid metabolites (FGMs) in 6 populations of red deer and roe deer exposed to potentially different levels of stress, resulting from both natural (predator presence, forest cover, undergrowth, ungulate density, and temperature) and anthropogenic (hunting harvest, percentage of build-up areas, and road density) factors. We found the highest and most variable FGM concentrations in both ungulates in areas without large carnivores, and the lowest and least variable FGM levels in areas with wolf and lynx. Anthropogenic factors (hunting harvest, roads, and built-up area) positively correlated with the gradient of FGM levels in both species. Both the mean and the variance of the FGM concentrations measured within populations of both red deer and roe deer were affected positively by variation in hunting harvest and negatively by the minimum temperature. The variance in the roe deer FGM was also positively influenced by the percentage of built-up areas. The results indicate that stress in wild ungulate populations is lower and less variable in areas utilized by large carnivores than in carnivore-free areas where human-related factors predominate. This may be explained by evolutionary adaptations of prey animals constantly exposed to the risk of natural predation and their inability of adapting to the risk from humans probably due to its high intensity and erratic occurrence

Red deer allocate vigilance differently in response to spatio-temporal patterns of risk from human hunters and wolves

Ungulate prey can use increased vigilance to reduce their risk of predation, but little is known of the combined and interactive risk effects from humans and wolves in determining ungulate behaviour across time and space. Understanding the interplay between these risk effects is increasingly important, considering the recolonisation of several large carnivores to more human-dominated landscapes in Europe. Aim: The aim of the present study was to assess the vigilance behaviour expressed by red deer (Cervus elaphus) in response to both humans and wolves in the Polish Białowieża Forest. Methods: Using a camera-trap transect, the effect of distance to human settlements, hunting season, patterns of space use by wolves (Canis lupus), canopy openness, canopy height, time of day, as well as sex/age of individuals, on the vigilance behaviour observed in red deer was studied using a model-selection approach. Key results: We did not find a clear effect of patterns of space use by wolves or distance to human settlements on red deer vigilance behaviour at the landscape scale. However, red deer showed increased vigilance during the hunting season and during the day outside of protected areas and reserves, because disturbance from human hunters is highest. Conversely, we also found that red deer were more vigilant at night within more protected areas, which is likely to be explained by the increased activity of wolves because human activity is strictly limited. Conclusions: Our study showed that vigilance behaviour of red deer in Białowieża Primeval Forest is more driven by human hunting than by the frequency of wolf presence at a landscape scale. This could be explained by the higher temporal and spatial predictability of human hunting activities than wolf risk. We found that patterns of wolf space use, as opposed to the omnipresent fear effects from humans, had only localised effects by increasing vigilance levels during night hours in non-hunting areas of the forest. The reverse was observed outside of protected reserves. Understanding how prey species respond to this new combination of risk from natural predators and humans, is increasingly important in a landscape where human risk is becoming ever more potent and carnivores recolonise

What Cues Do Ungulates Use to Assess Predation Risk in Dense Temperate Forests?

Anti-predator responses by ungulates can be based on habitat features or on the near-imminent threat of predators. In dense forest, cues that ungulates use to assess predation risk likely differ from half-open landscapes, as scent relative to sight is predicted to be more important. We studied, in the Białowieża Primeval Forest (Poland), whether perceived predation risk in red deer (Cervus elaphus) and wild boar (Sus scrofa) is related to habitat visibility or olfactory cues of a predator. We used camera traps in two different set-ups to record undisturbed ungulate behavior and fresh wolf (Canis lupus) scats as olfactory cue. Habitat visibility at fixed locations in deciduous old growth forest affected neither vigilance levels nor visitation rate and cumulative visitation time of both ungulate species. However, red deer showed a more than two-fold increase of vigilance level from 22% of the time present on control plots to 46% on experimental plots containing one wolf scat. Higher vigilance came at the expense of time spent foraging, which decreased from 32% to 12% while exposed to the wolf scat. These behavioral changes were most pronounced during the first week of the experiment but continuous monitoring of the plots suggested that they might last for several weeks. Wild boar did not show behavioral responses indicating higher perceived predation risk. Visitation rate and cumulative visitation time were not affected by the presence of a wolf scat in both ungulate species. The current study showed that perceived predation risk in red deer and wild boar is not related to habitat visibility in a dense forest ecosystem. However, olfactory cues of wolves affected foraging behavior of their preferred prey species red deer. We showed that odor of wolves in an ecologically equivalent dose is sufficient to create fine-scale risk factors for red deer.

What cues do ungulates use to assess predation risk in dense temperate forests?

Anti-predator responses by ungulates can be based on habitat features or on the near-imminent threat of predators. In dense forest, cues that ungulates use to assess predation risk likely differ from half-open landscapes, as scent relative to sight is predicted to be more important. We studied, in the Białowieża Primeval Forest (Poland), whether perceived predation risk in red deer (Cervus elaphus) and wild boar (Sus scrofa) is related to habitat visibility or olfactory cues of a predator. We used camera traps in two different set-ups to record undisturbed ungulate behavior and fresh wolf (Canis lupus) scats as olfactory cue. Habitat visibility at fixed locations in deciduous old growth forest affected neither vigilance levels nor visitation rate and cumulative visitation time of both ungulate species. However, red deer showed a more than two-fold increase of vigilance level from 22% of the time present on control plots to 46% on experimental plots containing one wolf scat. Higher vigilance came at the expense of time spent foraging, which decreased from 32% to 12% while exposed to the wolf scat. These behavioral changes were most pronounced during the first week of the experiment but continuous monitoring of the plots suggested that they might last for several weeks. Wild boar did not show behavioral responses indicating higher perceived predation risk. Visitation rate and cumulative visitation time were not affected by the presence of a wolf scat in both ungulate species. The current study showed that perceived predation risk in red deer and wild boar is not related to habitat visibility in a dense forest ecosystem. However, olfactory cues of wolves affected foraging behavior of their preferred prey species red deer. We showed that odor of wolves in an ecologically equivalent dose is sufficient to create fine-scale risk factors for red deer

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