Density-dependent switches in diet:a likely mechanism for negative feedbacks on goose population increase?

Goose grazing on arctic tundra vegetation has shown both positive and negative effects on subsequent foraging conditions. To understand the potential of a density-dependent feedback on herbivore population size, the relation between grazing pressure and future foraging conditions is essential. We studied the effect of increasing grazing pressure of barnacle geese (Branta leucopsis) on Spitsbergen. During the establishment of a breeding colony in the period 1992-2004, the proportion of graminoids decreased in the diet of wild geese, while the percentage of mosses increased. Grazing trials with captive geese in an unexploited area showed a similar shift in diet composition. High-quality food plants were depleted within years and over years. Intake rate declined too and as consequence, metabolisable energy intake rate (MEIR) decreased rapidly with increasing grazing pressure. During three successive years of experimental grazing, MEIR decreased at all levels of grazing pressure and declined below minimal energetic requirements when grazing exceeded natural levels of grazing pressure. This suggests that foraging conditions rapidly decline with increasing grazing pressure in these low-productive habitats. The potential for density-dependent feedbacks on local population increase is discussed

Behavioral response of naïve and non-naïve deer to wolf urine

Large carnivores are recolonizing many regions in Europe, where their ungulate prey have lived without them for >150 years. Whether the returning large carnivores will modify ungulate behavior and indirectly affect lower trophic levels, depends on the ability of ungulates to recognize risk based on past encounters and cues indicating carnivore presence. In two case studies, we tested, by means of camera trapping, the behavioral response of deer to wolf urine. The first case study was in the Netherlands where deer (still) live in absence of wolves, and the second in Poland with long-term wolf presence. As controls we used water (no scent) and all-purpose soap (unfamiliar scent). Deer vigilance level on control plots was 20% in both case studies indicating that wolf occupancy per se does not lead to a consistent difference in behavior. Placing wolf urine did not significantly affect deer behavior in either the wolf-absent or the wolf-present area. More intense cues, or a combination of cues, are likely needed to affect deer behavior. Moreover, we found an unexpected reaction of deer towards all-purpose soap of reduced foraging (and tendency for increased vigilance) in the wolf-present area, whereas it did not affect deer behavior in the wolf-absent area. We hypothesize that deer associate all-purpose soap with human presence, causing no response in human-dominated landscapes (the Netherlands), but triggering a behavioral reaction in more remote areas (Poland). This illustrates attention should be paid to controls used in scent experiments as they may be associated differently than intended

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

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

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

Small herbivores slow down species loss up to 22 years but only at early successional stage

The long-term influence of persistent small herbivores on successional plant community configuration is rarely studied. We used a herbivore exclusion experiment along the successional gradient in a salt-marsh system, to investigate the effects of hares and geese, and hares alone, on plant diversity at five successional stages (the earliest, two early, the intermediate and the late successional stages) in the short and long term, i.e. 7 and 22 years, respectively. Plant diversity declined over time at all successional stages except for the earliest one. Small herbivores slowed down species decline, but only at one early successional stage. Small herbivores slowed down species decline via decreasing dominance of preferred grass Festuca rubra in the short term, and less preferred Elytrigia atherica in the long term. The effects of hares and geese were more pronounced than hares alone, indicating an important additive role of geese, especially in the long term. Synthesis. Small herbivores can have a strong and long-lasting impact on plant diversity, but it highly depends on the abundance of small herbivores, which in turn depends on the quality and abundance of forage plants. A diverse herbivore community may have more positive effects on regulating plant communities